^/'/ illary circulation — Rapidity of entire circulation — Local variations in different parts . 229-264 CHAPTER XV. SECRETION. Nature of secretion — Variations in activity — Mucus — Sebaceous matter — Per- spiration — Tears — Milk — Secretion of bile — Anatomical peculiarities . 265-281 CHAPTER XYI. EXCRETION. Nature of excretion — Excrementitious substances — Effect of their retention — Urea — Its source — Conversion into carbonate of ammonia — Daily quan- tity of urea — Creatine — Creatinine— Urate of soda — Urates of potass and ammonia — General characters of the urine — Its composition — Variations — Accidental ingredients of the urine — Acid and alkaline fermentktions — Final decomposition of the urine ..... 282-304 CONTENTS. Xlll SECTION II. NERVOUS SYSTEM. CHAPTER I. GENERAL CHARACTER AND FUNCTIONS OF THE NERVOUS SYSTEM. PAGE Nature of the function performed by nervous system — Two kinds of nervous tissue — Fibres of white substance — Their minute structure — Division and inosculation of nerves — Gray substance — Nervous system of radiata — Of mollusca — Of articulata — Of mammalia and human subject — Structure of encepbalon — Connections of its different parts .... 305-327 CHAPTER II. OF NERVOUS IRRITABILITY, AND ITS MODE OP ACTION. Irritability of muscles — How esliibited — Influences wliich exbaust and destroy it — Nervous irritability — How exhibited — Continues after death — Exhausted by repeated excitement — Influence of direct and inverse electrical currents — Nervous irritability distinct from muscular irritability — Nature of the nervous force — Its resemblance to electricity — Difi'erences between the two 328-339 CHAPTER III. THE SPINAL CORD. Distinct seat of sensation and motion in nervous system — Sensibility and excitability — Distinct seat of sensibility and excitability in spinal cord — Crossed action of spinal cord — Independent and associated action of motor and sensitive filaments — Reflex action of spinal cord — How manifested dur- ing disease — Influence in health on sphincters, voluntary muscles, urinary bladder, &c. . . . . . . . . 340-356 CHAPTER lY. THE BRAIN. Seat of sensibility and excitability in difi'erent parts of the encephalon — Olfac- tory ganglia — Optic thalami — Corpora striata — Hemispheres — Remarkable cases of injury of hemispheres — Efi'ect of their removal — Imperfect develop- ment in idiots — Aztec children — Theory of phrenology — Cerebellum — Efi"ect of its injury or removal — Comparative development in different classes — Tubercula quadrigemina — Tuber annulare — Medulla oblongata — Three kinds of reflex action in nervous system ..... 357-384 xiv CONTENTS. CHAPTER Y. THE CRANIAL NERVES. PAGE Olfactory nerves — Optic nerves — Auditory nerves — Classification of cranial nerves — Motor Nerves of the Head — Motor ocull communis — Patheticus — Motor externtis — Small root of fifth pair— Facial nerve — Sublingual nerve — Spinal accessory — Sensitive Nerves of the Head — Fifth pair — Its sensi- bility — Effect of division — Influence on the organ of sight — Glosso-pharyn- geal nerve — Pneumogastric — Its distribution — Influence on pharynx and oesophagus — On larynx — On lungs — On stomach and digestion . 385-413 CHAPTER YI. SYSTEM OF THE GREAT SYMPATHETIC. Ganglia of the great sympathetic — Distribution of its nerves — Sensibility and excitability of sympathetic — Sluggish action of this nerve — Influence over organs of special sense — Elevation of temperature after division of sympa- thetic — Contraction of pupil following the same operation — Reflex actions taking place through the great sympathetic .... 414-425 SECTION III. KEPRODUCTION. CHAPTER I. ON THE NATURE OF REPRODUCTION, AND THE ORIGIN OF PLANTS AND ANIMALS. Nature and objects of the function of reproduction — Mode of its accomplish- ment — By generation from parents — Spontaneous generation — Mistaken in- stances of this mode of generation — Production of infusoria— Conditions of their development — Schultze's experiment on generation of infusoria — Pro- duction of animal and vegetable parasites— Encysted entozoa — Trichina spiralis — Tsnia — Cysticercus— Production of taenia from cysticercus — Of cysticercus from eggs of taenia — Plants and animals always produced by generation from parents ....... 427-441 CHAPTER II. ON SEXUAL GENERATION, AND THE MODE OF ITS ACCOMPLISHMENT. Sexual apparatus of plants— Fecundation of the germ — Its development into a new plant — Sexual apparatus of animals — Ovaries and testicles — Uni- sexual and bisexual species — Distinctive characters of the two sexes 442-445 CONTENTS. XV CHAPTER III. ON THE EGG, AND THE FEMALE ORGANS OF GENERATION. PAGE Size and appearance of the egg — Vitelline membrane — Vitellus — Germinative vesicle — Germinative spot — Ovaries — Graafian Follicles — Oviducts — Female generative organs of frog — Ovary and oviduct of fowl — Changes in the egg, while passing through the oviduct — Complete fowl's egg — Uterus and ova- ries of the sow — Female generative apparatus of the human subject — Fal- lopian tubes— Body of the uterus— Cervix of the uterus . . 446-457 CHAPTER lY. ON THE SPERMATIC FLUID, AND THE MALE ORGANS OF GENERATION, The spermatozoa — Their varieties in different species — Their movement — For- mation of spermatozoa in the testicles — Accessory male organs of generation — Epididymis — Vas deferens — Vesiculae seminales — Prostate — Cowper's glands — Function of spermatozoa — Physical conditions of fecundation 458-464 CHAPTER y. ON PERIODICAL OVULATION, AND THE FUNCTION OP MENSTRUATION. Periodical Ovulation — Pre-existence of eggs in the ovaries of all animals — Their increased development at the period of puberty — Their successive ripening and periodical discharge — Discharge of eggs independently of sexual intercourse — Rupture of Graafian follicle, and expulsion of the egg — Pheno- mena of cestruation — Menstruation — Correspondence of menstrual periods with periods of ovulation in the lower animals — Discharge of egg during menstrual period — Conditions of its impregnation, after leaving the ovary 465-477 CHAPTER Y I. ON THE CORPUS LUTEUM OF MENSTRUATION AND PREGNANCY. Corpus Luteum of Menstruation — Discharge of blood into the ruptured Graafian follicle — Decolorization of the clot, and hypertrophy of the membrane of the vesicle — Corpus luteum of menstruation, at the end of three weeks — Yellow coloration of convoluted wall — Corpus luteum of menstruation at the end of four weeks — Shrivelling and condensation of its tissues — Its condition at the end of nine weeks — Its final atrophy and disappearance — Corpus Luteum OF Pregnancy — Its continued development after the third week — Appearance at the end of second month — Of fourth month — At the termination of preg- nancy — Its atrophy and disappearance after delivery — Distinctive characters of corpora lutea of menstruation and pregnancy . . . 478-487 Xvi CONTENTS. CHAPTER VII. ON THE DEVELOPMENT OF THE IMPREGNATED EGG. PAGE Segmentation of the vitellus — Formation of blastodermic membrane. — Two layers of blastodermic membrane — Thickening of external layer — Formation of primitive trace — Dorsal plates — Abdominal plates — Closure of dorsal and abdominal plates on the median line — Formation of intestine — Of mouth and anus — Of organs of locomotion — Continued development of organs, after leaving the egg ....... 488-497 CHAPTER YIII. THE UMBILICAL VESICLE. Separation of vitelline sac into two cavities — Closure of abdominal walls, and formation of umbilical vesicle in fish — Mode of its disappearance after hatch- ing — Umbilical vesicle in human embryo — Formation and growth of pedicle — Disappearance of umbilical vesicle during embryonic life . . 498-500 CHAPTER IX. AMNION AND ALLANTOIS — DEVELOPMENT OF THE CHICK. Necessity for accessory organs in the development of birds and quadrupeds — Formation of amniotic folds — Their union and adhesion — Growth of allantois from lower part of intestine — Its vascularity — Allantois in the egg of the fowl — Respiration of the egg — Absorption of calcareous matter from the shell — Ossification of skeleton — Fracture of egg-shell — Casting off of amnion and allantois . . . . . . . 501-509 C HAPTER X. DEVELOPMENT OF THE EGG IN THE HUMAN SPECIES — FORMATION OF THE CHORION. Conversion of allantois into chorion — Subsequent changes of the chorion — Its villosities — Formation of bloodvessels in villosities — Action of villi of chorion in providing for nutrition of foetus — Proofs that the chorion is formed from the allantois — Partial disappearance of villosities of chorion, and changes in its external surface . ..... 510-515 CHAPTER XI. DEVELOPMENT OF UTERINE MUCOUS MEMBRANE — FORMATION OF THE DECIDUA. Structure of uterine mucous membrane — Uterine tubules — Thickening of uterine mucous membrane after impregnation — Decidua vera — Entrance of egg into uterus — Decidua reflexa— Inclosure of egg by decidua reflexa — Union of chorion with decidua — Changes in the relative development of dif- ferent portions of chorion and decidua ..... 516-522 CONTENTS. XVll CHAPTER XII. THE PLACENTA. PAGE Nourisliment of foetus by maternal and foetal vessels — Arrangement of the vascular membranes in different species of animals — Membranes of foetal pig — Cotyledon of cow's uterus — Development of foetal tufts in liuman pla- centa — Development of uterine sinuses — Relation of foetal and maternal bloodvessels in tlie placenta — Proofs that the maternal sinuses extend through the whole thickness of the placenta — Absorption and exhalation by the placental vessels . ..... 523-531 CHAPTER XIII. DISCHARGE OF THE OVUM, AND INVOLUTION OF THE UTERUS. Enlargement of amniotic cavity — Contact of amnion and chorion — Amniotic fluid — Movements of foetus — Union of decidua vera and reflexa — Expulsion of the ovum and discharge of decidual membrane — Separation of the pla- centa — Formation of new mucous membrane underneath the old decidua — Fatty degeneration and reconstruction of muscular walls of uterus 532-538 CHAPTER XIV. DEVELOPMENT OF THE EMBRYO — NERVOUS SYSTEM, ORGANS OF SENSE, SKELETON AND LIMBS. Formation of spinal cord and cerebro-spinal axis — Three cerebral vesicles — Hemispheres — Optic thalami — Tubercula quadrigemina — Cerebellum — Me- dulla oblongata — Eye — Pupillary membrane — Skeleton — Chorda dorsalis — Bodies of the vertebrae — Laminae and ribs — Spina bifida — Anterior and pos- terior extremities — Tail — Integument — Hair — Vernix caseosa — Exfoliation of epidermis ........ 539-545 CHAPTER XY. DEVELOPMENT OP THE ALIMENTARY CANAL AND ITS APPENDAGES. Formation of intestine — Stomach — Duodenum — Convolutions of intestine — Large and small intestine — Caput coli and appendix vermiformis — Umbi- lical hernia — Formation of urinary bladder — Urachus — Vesico-rectal septum — Perineura — Liver — Secretion of bile — Gastric juice — Meconium — Glyco- genic function of liver — Diabetes of foetus — Pharynx and oesophagus — Dia- phragm — Diaphragmatic hernia — Heart and pericardium — Ectopia cordis — Development of the face ...... 546-555 1 XVlll CONTENTS. CHAPTER XVI. DEVELOPMENT OF THE KIDNEYS, WOLFFIAN BODIES, AND INTERNAL ORGANS OF GENERATION. PAGE Wolffian bodies — Their structure — First appearance of kidneys — Growth of kidneys, and atrophy of Wolffian bodies — Testicles and ovaries — Descent of the testicles — Tunica vaginalis testis — Congenital inguinal hernia — Descent of the ovaries — Development of the uterus .... 556-565 CHAPTER XVII. DEVELOPMENT OF THE CIRCULATORY APPARATUS. First, or vitelline circulation — Area vasculosa — Sinus terminalis — Vitelline circulation of fish — Arrangement of arteries and veins in body of foetus — Second, or placental circulation — Omphalo-mesenteric arteries and veins — Circulation of the umbilical vesicle — Of the allantois and placenta — Umbi- lical arteries and veins — Third, or adult circulation — Portal and pulmonary systems — Development of the arterial system — Development of the venous system — Changes in the hepatic circulation — Portal vein — Umbilical vein — Ductus venosus — Changes in the cardiac circulation — Division of heart into right and left cavities — Aorta and pulmonary artery — Ductus arteriosus — Foramen ovale and Eustachian valve — Changes in circulation at the period of birth . . . . . . . . 566-587 C HAPTER XVIII. DEVELOPMENT OF THE BODY AFTER BIRTH. Condition of foetus at birth — Gradual establishment of respiration — Inactivity of the animal functions — Preponderance of reflex actions in the nervous system — Peculiarities in the action of drugs on infant — DiflTerence in rela- tive size of organs, in infaiat and adult — Withering and separation of umbi- lical cord — Exfoliation of epidermis — First and second sets of teeth — Sub- sequent changes in osseous, muscular and tegumentary systems, and gene- ral development of the body ...... 588-591 LIST OF ILLUSTRATIONS, ALL OF WHICH HAVE BEEN PREPARED FROM ORIGINAL DRAWINGS, WITH THE EXCEPTION OF ELEVEN, CREDITED TO THEIR AUTHORITIES. FIG. 1. Fibula tied iu a knot, after maceration in a dilute acid 2. Grains of potato starcli 3. Starch grains of Bermuda arrowroot 4. Starcli grains of wheat flour 5. Starch grains of Indian corn 6. Starch grains from wall of lateral ventricle 7. Stearine 8. Oleaginous principles of human fat 9. Human adipose tissue 10. Chyle 11. Globules of cow's milk 12. Cells of costal cartilages 13. Hepatic cells 14. Uriniferous tubules of dog 15. Muscular fibres of human uterus 16. Alimentary canal of fowl . 17. Compound stomach of ox 18. Human alimentary canal 19. Skull of rattlesnake 20. Skull of polar bear 21. Skull of the horse 22. Molar tooth of the horse . 23. Human teeth — upper jaw 24. Buccal and glandular epithelium deposited from saliva 25. Gastric mucous membrane, viewed from above 26. Gastric mucous membrane, in vertical section 27. Mucous membrane of pig's stomach 28. Gastric tubules from pig's stomach, pyloric portion 29. Gastric tubules from pig's stomach, cardiac portion 30. Confervoid vegetable, growing in gastric juice 31. Follicles of Lieberkiihn . 32. Brunner's duodenal glands 33. Contents of stomach, during digestion of meat 34. From duodenum of dog, during digestion of meat 35. From middle of small intestine . From Rymer Jones From Achille Richard PAGE 43 48 48 49 49 50 55 56 . 58 58 59 59 60 60 61 85 86 87 89 90 90 90 91 92 101 101 101 102 102 108 119 120 126 126 127 XX LIST OF ILLUSTRATIONS. FIG. 36. From last quarter of small intestine 37. One of the closed follicles of Peyer's patches 38. Glandulse agminatse 39. Extremity of intestinal villus 40. Panizza's experiment on absorption by bloodvessels 41. Chyle, from commencement of thoracic duct 42. Lacteals, thoracic diict, &c. 43. Lacteals and lymphatics . 44. Intestinal epithelium, in intervals of digestion 45. Intestinal epithelium, during digestion . 46. Cholesteriu .... 47. Ox-bile, crystallized 48. Glyko-cholate of soda from ox-bile 49. Glyko-cholate and tauro-cholate of soda, from ox-bile 50. Dog's bile, crystallized 51. Human bile, showing resinous matters . 52. Crystalline and resinous biliary substances, from dog's intestine 53. Duodenal iistula .... 54. Human blood-globules 55. The same, seen out of focus 56. The same, seen within the focus . 57. The same, adhering together in rows 58. The same, swollen by addition of water . 59. The same, shrivelled by evaporation 60. Blood-globules of frog 61. White globules of the blood 62. Coagulated fibrin . 63. Coagulated blood .... 64. Coagulated blood, after separation of clot and serum 65. Recent coagulum .... 66. Coagulated blood, clot buffed and cupped 67. Head and gills of men'obranchus . 68. Lung of frog .... 69. Human larynx, trachea, bronchi, and lungs 70. Single lobule of human lung 71. Diagram illustrating the respiratory movements 72. Small bronchial tube 73. Human larynx, with glottis closed 74. The same, with glottis open 75. Human larynx ; posterior view . 76. Circulation of fish 77. Circulation of reptiles 78. Circulation of mammalians 79. Human heart, anterior view 80. Human heart, posterior view 81. Right auricle and ventricle, tricuspid valve open, arterial valves closed 82. Right auricle and ventricle, tricuspid valve closed, arterial valves open 83. Course of blood through the heart 84. Illustrating production of valvular sounds 85. Heart of frog, in relaxation LIST OF ILLUSTRATIONS. XXI FIG. PAGE 86. Heart of frog, in contraction ..... 241 87. Simple looped fibres ..... 241 88. Bullock's heart, showing superficial muscular fibres 242 89. Left ventricle of bullock's heart, showing deep fibres . 242 90. Diagram of circular fibres of the heart . 243 91. Converging fibres of the apex of the heart 243 92. Arterial circulation ..... 248 93. Arteiy in pulsation ..... 248 94. Volkmann's apparatus ..... 251 95. The same ...... 251 96. Vein, with valves open ..... 253 97. Vein, with valves closed .... 253 98. Small artery, with capillary branches . 255 99. Capillary network . . . 256 100. Capillary circulation .... 257 101. Diagram of the circulation 264 102. Follicles of a compound mucous glandule . From Kolliker 268 103. Meibomian glands .... From Ludovic 270 104. Perspiratory gland . . . From Todd and Bowman 271 105. Glandular structure of mamma . . . • 274 106. Colostrum corpuscles .... 275 107. Milk-globules ..... 276 108. Division of portal vein in liver . 279 109. Lobule of liver . , 280 110. Hepatic cells ..... 281 111. Urea .... From Lehmann (Funke's Atlas) 285 112. Creatine .... From Lehmann (Funke's Atlas) 287 113. Creatinine . . . From Lehmann (Funke's Atlas) 288 114. Urate of soda ........ 289 115. Uric acid ..... . 296 116. Oxalate of lime .... . 302 117. Phosphate of magnesia and ammonia . . 304 118. Nervous filaments, from brain . . 309 119. Nervous filaments, from sciatic nerve . . 310 120. Division of a nerve . 311 121. Inosculation of nerves . . 312 122. Nerve cells .... . 312 123. Nervous system of starfish . 313 124. Nervous system of aplysia . 315 125. Nervous system of centipede . 316 126. Cerebro-spinal system of man . . 319 127. Spinal cord .... . 320 128. Brain of alligator . 322 129. Brain of rabbit .... . 323 130. Medulla oblongata of human brain . 324 131. Diagram of human brain . 326 132. Experiment showing irritability of muscles . 329 133. Experiment showing irritability of nerve . 331 134. Action of direct and inverse currents . 334 135. Diagram of spinal cord and nerves . 342 XXll LIST OF ILLUSTKATIONS. FIG. 136. Spinal cord in vertical section . 137. Experiment, showing effect of poisons on nerves 138. Pigeon, after removal of the hemispheres 139. Aztec children 140. Brain in situ 141. Transverse section of brain 142. Pigeon, after removal of the cerebellum 143. Inferior surface of brain of cod . 144. Inferior surface of brain of fowl . 145. Course of optic nerves in man . 146. Facial nerve 147. Distribution of fifth nerve upon the face 148. Pneumogastric nerve 149. Great sympathetic 150. Cat, after division of sympatlietic in the neck 151. Different kinds of infusoria 152. Experiment on spontaneous generation . . From Schultze 153. Trichina spiralis . 154. Taenia .... 155. Cysticercus, retracted 156. Cysticercus, unfolded 157. Blossom of Convolvulus purpureus 158. Single articulation of Taenia crassicollis 159. Human ovum 160. Human ovum, ruptured by pressure 161. Female generative organs of frog 162. Mature frogs' eggs 163. Female generative organs of fowl 164. Fowl's egg 165. Uterus and ovaries of the sow . 166. Generative organs of human female 167. Spermatozoa 168. Graafian follicle . 169. Ovary with Graafian follicle ruptured 170. Graafian follicle, ruptured and filled with blood 171. Corpus luteum, three weeks after menstruation 172. Corpus luteum, four weeks after menstruation 173. Corpus luteum, nine weeks after menstruation 174. Corpus luteum of pregnancy, at end of second month 175. Corpus luteum of pregnancy, at end of fourth month 176. Corpus luteum of pregnancy, at term 177. Segmentation of the vitellus 178. Impregnated egg, showing embryonic spot 179. Impregnated egg, showing two layers of blastodermic membrane 180. Impregnated egg, farther advanced 181. Frog's egg, at an early period 182. Egg of frog, in process of development 183. Egg of frog, farther advanced 184. Tadpole, fully developed 185. Tadpole, changing into frog LIST OF ILLUSTRATIONS. FIG. 186. Perfect frog .... 187. Egg of fish .... 188. Young fish, with umbilical vesicle 189. Human embryo, with umbilical vesicle 190. Fecundated egg, showing formation of amnion 191. Fecundated egg, showing commencement of allantois 192. Fecundated egg, with allantois nearly complete 193. Fecundated egg, with allantois fully formed 194. Egg of fowl, showing area vasculosa 195. Egg of fowl, showing allantois, amnion, &c. 196. Human ovum, showing formation of chorion 197. Human chorion . 198. Villosity of chorion 199. Human ovum, at end of third month 200. Uterine mucous membrane 201. Uterine tubules . 202. Impregnated uterus, showing formation of decidua . 203. Impregnated uterus, showing formation of decidua reflexa 204. Impregnated uterus, with decidua reflexa complete 205. Impregnated uterus, showing union of chorion and decidua 206. Pregnant uterus, showing formation of placenta 207. Foetal pig, with membranes 208. Cotyledon of cow's uterus 209. Foetal tuft of human placenta . 210. Vertical section of placenta 211. Human ovum, at end of first month 212. Human ovum, at end of third month 213. Gravid human uterus and contents 214. Muscular fibres of unimpregnated uterus 215. Muscular fibres of human uterus, ten days after parturition 216. Muscular fibres of human uterus, three weeks after parturition 217. Formation of cerebro-spinal axis 218. Formation of cerebro-spinal axis 219. Foetal pig, showing brain and spinal cord 220. Foetal pig, showing brain and spinal cord 221. Head of foetal pig 222. Brain of adult pig 223. Formation of alimentary canal . 224. Head of human embryo, at twenty days . . From Longet 225. Head of human embryo, at end of first month . . From Longet 226. Head of human embryo, at end of second month 227. Foetal pig, showing Wolfiian bodies 228. Foetal pig, showing first appearance of kidneys 229. Internal organs of generation 230. Internal organs of generation 231. Formation of tunica vaginalis testis 232. Congenital inguinal hernia 233. Egg of fowl, showing area vasculosa 234. Egg of fish, showing vitelline circulation 235. Young embryo and its vessels . PAGE 496 498 499 499 502 503 503 504 505 506 510 512 513 514 517 517 519 519 519 521 522 524 524 527 527 532 53a 534 537 537 538 539 540 540 541 541 541 547 553 554 554 556 558 558 560 561 562 567 567 568 XXIV LIST OF ILLUSTRATIOXS. FIG. 236. Embryo and its vessels, farther advanced 237. Arterial system, embryonic form 238. Arterial system, adult form 239. Early condition of venous system 240. Venous system, farther advanced 241. Continued development of venous system 242. Adult condition of venous system 243. Early form of hepatic circulation 244. Hepatic circulation, farther advanced . 245. Hepatic circulation, during latter part of foetal life 246. Adult form of hepatic circulation 247. Foetal heart 248. Foetal heart 249. Foetal heart 250. Foetal heart 251. Heart of infant 252. Heart of human foetus, showing Eustachian valve 253. Circulation through the fcetal heart 254. Adult circulation through the heart PAGE 569 571 571 573 574 574 575 576 577 577 578 579 579 579 580 580 582 583 586 HUMAN PHYSIOLOGY. INTRODUCTION. I. Physiology is the study of the phenomena presented by organized bodies, animal and vegetable. These phenomena are different from those presented by inorganic substances. They require, for their production, the existence of peculiarly formed animal and vegetable organisms, as well as the presence of various external conditions, such as warmth, light, air, moisture, &c. They are accordingly more complicated than the phenomena of the inorganic world, and require for their study, not only a pre- vious acquaintance with the laws of chemistry and physics, but, in addition, a careful examination of other characters which are pecu- liar to them. These peculiar phenomena, by which we so readily distinguish living organisms from inanimate substances, are called Vital j^^^eno- mena., or the phenomena of Life. Physiology consequently includes the study of all these phenomena, in whatever order or species of organized body they may originate. We find, however, upon examination, that there are certain general characters by which the vital phenomena of vegetables re- semble each other, and by which they are distinguished from the vital phenomena of animals. Thus, vegetables absorb carbonic acid, and exhale oxygen ; animals absorb oxygen, and exhale car- bonic acid. Vegetables nourish themselves by the absorption of unorganized liquids and gases, as water, ammonia, saline solutions, &c. ; animals require for their support animal or vegetable sub- stances as food, such as meat, fruits, milk, &c. Physiology, theu, 2 18 INTEODUCTIOX. is naturally divided into two parts, viz., Vegetable Physiology, and Animal Physiology. Again, the different groups and species of animals, while they resemble each other in their general characters, are distinguished by certain minor diff'erences, both of structure and function, which require a special study. Thus, the physiology of fishes is not ex- actly the same with that of reptiles, nor the physiology of birds with that of c[uadrupeds. Among the warm-blooded quadrupeds, the carnivora absorb more oxygen, in proportion to the carbonic acid exhaled, than the herbivora. Among the herbivorous quad- rupeds, the process of digestion is comparatively simple in the horse, while it is complicated in the ox, and other ruminating ani mals. There is, therefore, a special physiology for every distinct species of animal. Human Physiology treats of the vital phenomena of the human species. It is more practically important than the physiology of the lower animals, owing to its connection with human pathology and therapeutics. But it cannot be made the exclusive subject of our study ; for the special physiology of the human body cannot be properly understood without a previous acquaintance with the vital phenomena common to all animals, and to all vegetables; besides which, there are many phj^siological questions that require for their solution experiments and observations, which can onl}^ be made upon the lower animals. While the following treatise, therefore, has for its principal sub- ject the study of Human Physiology, this will be illustrated, when- ever it may be required, by what we know in regard to the vital phenomena of vegetables and of the lower animals. II. Since Physiology is the study of the active phenomena of livincr bodies, it requires a previous acquaintance with their struc- ture, and with the substances of which they are composed ; that is, with their anatomy. Anatomy, again, requires a previous acquaintance with inorganic substances; since some of these inorganic substances enter into the composition of the body. Chloride of sodium, for example, water and phosphate of lime, are component parts of the animal frame, and therefore require to be studied as such by the anatomist. Now these inorganic substances present certain active phenomena, when placed under the requisite external conditions, which are characteristic of them, and by which they may be recognized. INTRODUCTION. 19 Thus lime, dissolved in water, if brought into contact with car- bonic acid, alters its condition, and takes part in the formatiou of an insoluble substance, carbonate of lime, which is thrown down as a deposit. A knowledge of such chemical reactions as these is necessary to the anatomist, since it is by them that he is enabled to recognize the inorganic substances, forming a part of the animal body. It is important to observe, however, that a knowledge of these reactions is necessary to the anatomist only in order to enable him to judge of the presence or absence of the inorganic substances to which they belong. It is the object of the anatomist to make him- self acquainted with every constituent part of the body. Those parts, therefore, which cannot be recognized by their form and texture, he distinguishes by their chemical reactions. But after- ward, he has no occasion to decompose them further, or to make them enter into new combinations; for he only wishes to know these substances as they exist in the hody, and not as they may exist under other conditions. The unorganized substances which exist in the body as compo- nent parts of its structure, such as chloride of sodium, water, phos- phate of lime, &c., are called the proximate principles of the body. Mingled together in certain proportions, they make up the animal fluids, and associated also in a solid form, they constitute the tissues and organs, and in this way make up the entire frame. Anatomy makes us acquainted with all these component parts of the body, both solid and fluid. It teaches us the structure of the body in a state of rest; that is, just as it would be after life had suddenly ceased, and before putrefaction had begun. On the other hand. Physiology is a description of the body in a state of activity. It shows us its movements, its growth, its reproduction, and the chemical changes which go on in its interior; and in order to com- prehend these, we must know, beforehand, its entire mechanical, textural, and chemical structure. It is evident, therefore, that the description of the proximate prin- ciples^ or the chemical substances entering into the constitution of the body, is, strictly speaking, a part of Anatomy. But there are many reasons why this study is more conveniently pursued in con- nection with Physiology ; for some of the proximate principles are derived directly, as we shall hereafter show, from the external world, and some are formed from the elements of the food in the process of digestion ; while most of them undergo certain changes in the 20 IXTEODUCTION. interior of the body, whicli result in the formation of new sub- stances ; all these active phenomena belonging necessarily to the domain of Physiology. The description of the proximate principles of animals and vege- tables will therefore be introduced into the following pages. The description of the minute structures of the bodj', or Micro- scopic Anatomy, is also so closely connected with some parts of Phy- siology as to make it convenient to speak of them together ; and this will accordingly be done, whenever the nature of the subject may make it desirable. III. The study of Physiology, like that of all the other natural sciences, is a study of phenomena, and of phenomena alone. The essential nature of the vital processes, and their ultimate causes, are questions which are beyond the reach of the physiologist, and cannot be determined by the means of investigation which are at his disposal. Consequently, all efforts to solve them Avill only serve to mislead the investigator, and to distract his attention from the real subject of examination. Much time has been lost, for example, in discuss- ing the probable reason why menstruation returns, in the human female, at the end of every four weeks. But the observation of nature, which is our only means of scientific investigation, cannot throw any light on this point, but only shows us the fact that men- struation does really recur at the above periods, together with the phenomena which accompany it, and the conditions under which it is hastened or retarded, and increased or diminished, in intensity, duration, &c. If we employ ourselves, consequently, in the discus- sion of the reason above mentioned, we shall only become involved in a network of hypothetical surmises, which can never lead to any definite result. Our time, therefore, will be much more profitabl}^ devoted to the study of the above phenomena, which can be learned from nature, and which constitute, afterward, a permanent acquisi- tion. The physiologist, accordingly, confines himself strictly to the study of the vital phenomena themselves, their characters, their frequency, their regularity or irregularity, and the conditions under which they originate. When he has discovered that a certain phenomenon always takes place in the presence of certain conditions, he has established what is called a general principle, or a Law of Physiology. INTRODUCTION. 21 As, for example, wben lie has ascertained that sensation and motion occupy distinct situations in every part of the nervous system. This " Law," however, it must be remembered, is not a discovery by itself, nor does it give him any new information, but is simply the expression, in convenient and comprehensive language, of the facts with which he was already previously acquainted. It is very dangerous, therefore, to make these laws or general principles the subjects of our study instead of the vital phenomena, or to suppose that they have any value, except as the expression of previously ascertained facts. Such a misconception would lead to bad practi- cal results. For if we were to observe a phenomenon in discord- ance with a " law" or " principle," we might be led to neglect or misinterpret the phenomenon, in order to preserve the law. But this would be manifestly incorrect. For the law is not superior to the phenomenon, but, on the contrary, depends upon it, and derives its whole authority from it. Such mistakes, however, have been repeatedly made in Physiology, and have frequently retarded its advance. IV. There is only one means by which Physiology can be studied : that is, the observation of nature. Its phenomena cannot be reasoned out by themselves, nor inferred, by logical sequence, from any original principles, nor from any other set of phenomena whatever. In Mathematics and Philosophy, on the other hand, certain truths are taken for granted, or perceived by intuition, and the remainder afterward derived from them by a process of reasoning. But in Physiology, as in all the other natural sciences, there is no such starting point, and it is impossible to judge of the character of a phenomenon until after it has been observed. Thus, the only way to learn what action is exerted by nitric acid upon carbonate of soda is to put the two substances together and observe the changes which take place ; for there is nothing in the general characters of these two substances which could guide us in anticipating the result. Neither can we infer the truths of Physiology from those of Anatomy, nor the truths of one part of Physiology from those of another part ; but all must be ascertained directly and separately by observation. For, although one department of natural science is almost always a necessary preliminary to the study of another, yet the facts of the 22 INTEODUCTION. latter can never he in the least degree inferred from those of the former^ hut must he studied hy themselves. Thus Chemistry is essential to Anatomy, because certain sub- stances, as we have already shown, belonging to Chemistry, such as chloride of sodium, occur as constituents of the animal body. Chemistry teaches us the composition, reactions, mode of crystal- lization, solubility, &c., of chloride of sodium ; and if we did not know these, we could not extract it, or recognize it when extracted from the body. But, however well we might know the chemistry of this substance, we could never, on that account, irfer its presence in the body or otherwise, nor in what quantities nor in what situa- tions it would present itself. These facts must be ascertained for themselves, by direct investigation, as a part of anatomy proper. So, again, the structure of the body in a state of rest, or its ana- tomy, is to be first understood; but its active phenomena or its physiology must then be ascertained by direct observation and experiment. The most intimate knowledge of the minute struc- ture of the muscular and nervous fibres could not teach us any- thing of their physiology. It is only by experiment that we ascer- tain one of them to be contractile, the other sensitive. Many of the phenomena of life are chemical in their character, and it is requisite, therefore, that the physiologist know the or- dinary chemical properties of the substances composing the animal frame. But no amount of previous chemical knowledge will enable him to foretell the reactions of any chemical substance in the interior of the body ; because the peculiar conditions, under which it is there placed, modify these reactions, as an elevation or depression of temperature, or other external circumstance, might modify them outside the body. We must not, therefore, attempt to deduce the chemical phe- nomena of physiology from any previously established facts, since these are no safe guide ; but must study them by themselves, and depend for our knowledge of them upon direct observation alone. V. By the term Vital phenomena^ we mean those phenomena which are manifested in the living body, and which are characteristic of its functions. Some of these phenomena are physical or mechanical in their character; as, for example, the play of the articulating surfaces upon each other, the balancing of the spinal column Avith its ap- pendages, the action of the elastic ligaments. Nevertheless, these INTRODUCTION. 23 ]>heuomena, tbougli strictly physical in character, are often entirely peculiar and different from those seen elsewhere, because the me- chanism of their production is peculiar in its details. Thus the liuman voice and its modulations are produced in the larynx, in accordance with the general physical laws of sound ; but the arrangement of the elastic and movable vocal chords, with the columns of air above and below, the moist and flexible mucous membrane, and the contractile muscles outside of it, are of such a special character, that the entire apparatus, as well as the sounds produced by it, is peculiar; and its action cannot be properly compared with that of any other known musical instrument. In the same manner, the movements of the heart are so com- plicated and remarkable that they cannot be comprehended, even by one who is acquainted with the anatomy of the organ, without a direct examination. This is not because there is anything essen- tially obscure or mysterious in their nature, for they are purely mechanical in character; but because their conditions are so pecu- liar, owing to the tortuous course of the muscular fibres, their ar- rangement in interlacing layers, their attachments and relations, that their combined action produces an effect altogether peculiar, and one which is not similar to anything outside the living body. A very large and important class of the vital phenomena are those of a chemical character. It is one of the characteristics of living bodies that a succession of chemical actions, combinations and decompositions, is constantly going on in their interior. It is one of the necessary conditions of the existence of every animal and every vegetable, that it should constantly absorb various sub- stances from without, which undergo different chemical alterations i n its interior, and are finally discharged from it under other forms. If these changes be prevented from taking place, life is immediately extinguished. Thus animals constantly absorb, on the one hand, water, oxygen, salts, albumen, oil, sugar, &c., and give up, on the other hand, to the surrounding media, carbonic acid, water, ammonia, urea, and the like; while between these two extreme points, of ab- sorption and exhalation, there take place a multitude of different transformations which are essential to the continuance of life. Some of these chemical actions are the same with those which are seen outside the body ; but most of them are entirely peculiar, and do not take place, and cannot be made to take place, anywhere else. This, again, is not because there is anything particularly mysterious or extraordinary in their nature^ but because the con- 24 INTEODUCTION. ditions necessary for their accomplisliment exist in the body, and do not exist elsewhere. All chemical phenomena are liable to be modified by surrounding conditions. Many reactions, for example, which will take place at a high temperature, will not take place at a low temperature, and vice versa. Some will take place in the light, but not in the dark; others will take place in the dark, but not in the light. If a hot concentrated solution of sulphate of soda be allowed to cool in contact with the atmosphere, it crystallizes: covered with a film of oil, it remains fluid. Because a chemical reaction, therefore, takes place under one set of conditions, we can- not be at all sure that it will also take place under others, which are different. The chemical conditions of the living body are exceedingly com- plicated. In the animal solids and fluids there are many substances mingled together in varying quantities, which modify or interfere with each other's reactions. New substances are constantly entering by absorption, and old ones leaving by exhalation ; while the circu- lating fluids are constantly passing from one part of the body to another, and coming in contact with different organs of different texture and composition. All these conditions are peculiar, and so modify the chemical actions taking place in the body, that they are unlike those met with anywhere else. If starch and iodine be mingled together in a watery solution, they unite with each other, and strike a deep opaque blue color ; but if they be mingled in the blood, no such reaction takes place, because it is prevented by the presence of certain organic substances which interfere with it. If dead animal matter be exposed to warmth, air, and moisture, it putrefies ; but if introduced into the living stomach, even after putrefaction has commenced, this process is arrested, because the fluids of the stomach cause the animal substance to undergo a peculiar transformation (digestion), after which the bloodvessels immediately remove it by absorption. There are also certain sub- stances which make their appearance in the living body, both of animals and vegetables, and which cannot be formed elsewhere ; such as fibrin, albumen, casein, pneumic acid, the biliary salts, mor phine, &c. These substances cannot be manufactured artificially, simply because the necessary conditions cannot be imitated. They require for their production the presence of a living organism. The chemical phenomena of the living body are, therefore, not different in their nature from any other chemical phenomena ; but INTRODUCTION. 25 they are different in tlieir conditions and in tbeir results, and are consequently peculiar and characteristic. Another set of vital phenomena are those whicli are manifested in the processes of reproduction and development. They are again entirely distinct from any phenomena which are exhibited by matter not endowed with life. An inorganic substance, even when it has a definite form, as, for example, a crystal of fluor-spar, has no particular relation to any similar form which has preceded, or any other which is to follow it. On the other hand, every animal and every vegetable owes its origin to preceding animals or vege- tables of the same kind ; and the manner in which this production takes place, and the different forms through which the new body successively passes in the course of its development, constitute the phenomena of reproduction. These phenomena are mostly de- pendent on the chemical processes of nutrition and growth, which take place in a particular direction and in a particular manner; but their results, viz., the production of a connected series of different forms, constitute a separate class of phenomena, which cannot be explained in any manner by the preceding, and require, therefore, to be studied by themselves. Another set of vital phenomena are those which belong to the nervous system. These, like the processes of reproduction and development, depend on the chemical changes of nutrition and growth. That is to say, if the nutritive processes did not go on in a healthy manner, and keep the nervous system in a healthy condi- tion, the peculiar phenomena which, are characteristic of it could not take place. The nutritive processes are necessary conditions of the nervous phenomena. But there is no other connection between them ; and the nervous phenomena themselves are distinct from all others, both in their nature and in the mode in which they are to be studied. A troublesome confusion might arise if we were to neglect the distinction that really exists between these different sets of phe- nomena, and confound them together under the expectation of thereby simplifying our studies. Since this can only be done by overlooking real points of difference, its effect will merely be to introduce erroneous ideas and suggest unfounded similarities, and will therefore inevitably retard our progress instead of advancing it. It has been sometimes maintained, for example, that all the vital phenomena, those of the nervous system included, are to be reduced to the chemical chansjes of nutrition, and that these asain are to be 26 INTRODUCTION. regarded as not at all different in any respect from the ordinary chemical changes taking place outside the body. This, however, is not only erroneous in theory, but conduces also to a vicious mode of study. For it draws away our attention from the phe- nomena themselves and their real characteristics, and leads us to deduce one set of phenomena from what we know of another; a method which we have already shown to be unsafe and pernicious. It has also been asserted that the phenomena of the nervous system are identical with those of electricity ; for no other reason than that there exist between them certain general resemblances. But when we examine the phenomena in detail, we find that, beside these general resemblances, there are many essential points of dis- similarity, which must be suppressed and kept out of sight in order to sustain the idea of the assumed identity. This assumption is consequently a forced and unnatural one, and the simplicity which it was intended to introduce into our physiological theories is imaginary and deceptive, and is attained only by sacrificing a part of those scientific truths, which are alone the real object of our study. We should avoid, therefore, making auy such unfounded comparisons ; for the theoretical simplicity which results from them does not compensate for the loss of essential scientific details. VI. The study of Physiology is naturally divided into three distinct Sections : — The first of these includes everything which relates to the Nutri- tion of the body in its widest sense. It comprises the history of the proximate principles, their source, the manner of their produc- tion, the proportions in which they exist in different kinds of food and drink, the processes of digestion and absorption, and the con- stitution of the circulating fluids; then the physical phenomena of the circulation and the forces by which it is accomplished ; the changes which the blood undergoes in different parts of the body ; all the phenomena, both physical and chemical, of respiration; those of secretion and excretion, and the character and destination of the secreted and excreted fluids. All these processes have reference to a common object, viz., the preservation of the internal structure and healthy organization of the individual. With certain modifi- cations, they take place in vegetables as well as in animals, and are consequently known by the name of the vegetative functions. The Second Section, in the natural order of study, is devoted to the phenomena of the Nervous System. These phenomena are INTRODUCTION. 27 Dot exhibited by vegetables, but belong exclusively to animal or- ganizations. They bring the animal body into relation with the external world, and preserve it from external dangers, through the means of sensation, movement, consciousness, and volition. They are more particularly distinguished by the name of the animal functions. Lastly comes the study of the entire process of Reproduction. Its phenomena, again, with certain modifications, are met with in both animals and vegetables ; and might, therefore, with some pro- priety, be included under the head of vegetative functions. But their distinguishing peculiarity is, that they have for their object the production of new organisms, which take the place of the old and remain after they have disappeared. These phenomena do not, therefore, relate to the preservation of the individual, but to that of the species; and any study which concerns the species comes properly after we have finished everything relating to the individual. SECTIOI( I. NUTRITION. CHAPTER I. PROXIMATE PRINCIPLES IN GENERAL. The study of Nuteition begins naturally with tliat of the 2^roxi- mate princijjies, or the substances entering into the composition of the different parts of the body, and the different kinds of food. In examining the body, the anatomist finds that it is composed, first, of various parts, which are easily recognized by the eye, and which occupy distinct situations. In the case of the human body, for example, a division is easily made of the entire frame into the head, the neck, the trunk, and extremities. Each of these regions, again, is found, on examination, to contain several distinct parts, or "organs," which require to be separated from each other by dissec- tion, and which are distinguished by their form, color, texture, and consistency. In a single limb, for example, every bone and every muscle constitutes a distinct organ. In the trunk, we have the heart, the lungs, the liver, spleen, kidneys, spinal cord, &c., each of which is also a distinct organ. When a number of organs, differing in size and form, but similar in texture, are found scattered through- out the entire frame, or a large portion of it, they form a connected set or order of parts, which is called a " system." Thus, all the muscles taken together constitute the muscular system; all the bones, the osseous system; all the arteries, the arterial system. Several entirely different organs may also be connected with each other, so that their associated actions may tend to accomplish a single object, and they then form an " apparatus." Thus the heart, arteries, capillaries, and veins, together, form the circulatory appa- ratus; the stomach, liver, pancreas, intestine, &c., the digestive apparatus. Every organ, again, on microscopic examination, is 30 PEOXIMATE PRIXCIPLES IX GEXEPvAL. seen to be made up of minute bodies, of definite size and figure, which, are so small as to be invisible to the naked eye, and which, after separation from each other, cannot be further subdivided with- out destroying their organization. They are, therefore, called "ana- tomical elements." Thus, in the liver, there are hepatic cells, capil- lary bloodvessels, the fibres of Glisson's capsule, and the ultimate filaments of the hepatic nerves. Lastly, two or more kinds of ana- tomical elements, interwoven with each other in a particular manner, form a "tissue." Adipose vesicles, with capillaries and nerve tubes, form adipose tissue. White fibres and elastic fibres, with capillaries and nerve tubes, form areolar tissue. Thus the solid parts of the entire body are made up of anatomical elements, tissues, organs, systems, and apparatuses. Every organized frame, and even every apparatus, every organ, and every tissue, is made up of different parts, variously interwoven and connected with each other, and it is this character which constitutes its organization. But besides the above solid forms, there are also certain fluids, which are constantly present in various parts of the body, and which, from their peculiar constitution, are termed "animal fluids." These fluids are just as much an essential part of the body as the solids. The blood and the lymph, for example, the pericardial and synovial fluids, the saliva, which always exists more or less abundantly in the ducts of the parotid gland, the bile in the biliary ducts and the gall-bladder: all these go to make up the entire body, and are quite as necessary to its structure as the muscles or the nerves. Now, if these fluids be examined, they are found to be made up of many different substances, which are mingled together in certain propor- tions ; these proportions being constantly maintained at or about the same standard by the natural processes of nutrition. Such a fluid is termed an organized fluid. It is organized by virtue of the numerous ingredients which enter into its composition, and the regular proportions in which these ingredients are maintained. Thus, in the plasma of the blood, we have albumen, fibrin, water, chlorides, carbonates, phosphates, &c. In the urine, we find water, urea, urate of soda, creatine, creatinine, coloring matter, salts, &c. These substances, which are mingled together so as to make up, in each instance, by their intimate union, a homogeneous liquid, are called the proximate peikciples of the animal fluid. In the solids, however, even in those parts which are apparently homogeneous, there is the same mixture of different ingredients. In the hard substance of bone, for example, there is, first, water, TKOXIMATE PRINCIPLES IN GENERAL, 31 •\vliich may be expelled bj evaporation; second, phosphate and car- bonate of lime, which may be extracted by the proper solvents; third, a peculiar animal matter, with which these calcareous salts are in union ; and fourth, various other saline substances, in special proportions. In the muscular tissue, there is chloride of potassium, lactic acid, water, salts, albumen, and an animal matter termed mus- culine. The difference in consistenc}^ between the solids does not, therefore, indicate any radical difference in their constitution. Both solids and fluids are equallj' made up of proximate principles, min- gled together in various proportions. It is important to understand, however, exactly what are proxi- mate principles, and what are not such ; for since these principles are extracted from the animal solids and fluids, and separated from each other by the help of certain chemical manipulations, such as evaporation, solution, crystallization, and the like, it might be sup- posed that every substance which could be extracted from an organ- ized solid or fluid, by chemical means, should be considered as a proximate principle. That, however, is not the case. A proximate principle is properly defined to be any substance, luhether simple or compound, chemically speaking, ichich exists, under its own form, in the animal solid or fluid, and which can be extracted by means which do not alter or destroy its chemical properties. Phosphate of lime, for example, is a proximate principle of bone, but phosphoric acid is not so, since it does not exist as such in the bony tissue, but is pro- duced only by the decomposition of the calcareous salt ; still less phosphorus, which is obtained only by the decomposition of the phosphoric acid. Proximate principles may, in fact, be said to exist in all solids or fluids of mixed composition, and may be extracted from them by the same means as in the case of the animal tissues or secretions. Thus, in a watery solution of sugar, we have two pi'oximate princi- ples, viz : first, the water, and second, the sugar. The water may be separated by evaporation and condensation, after which the sugar remains behind, in a crystalline form. These two substances have, therefore, been simply separated from each other by the pro- cess of evaporation. They have not been decomposed, nor their chemical properties altered. On the other hand, the oxygen and hydrogen of the water were not proximate principles of the original solution, and did not exist in it under their own forms, but only in a state of combination; forming, in this condition, a fluid substance (water), endowed with sensible properties entirely different from 32 PROXIMATE PRINCIPLES IN GENERAL. theirs. If we wisli to ascertain, accordingly, the nature and proper- ties of a saccharine solution, it will afford us but little satisfaction to extract its ultimate chemical elements; for its nature and properties depend not so much on the presence in it of the ultimate elements, oxygen, hydrogen, and carbon, as on the particular forms of com- bination, viz., water and sugar, under which they are present. It is very essential, therefore, that in extracting the proximate principles from the animal body, only such means should be adopted as will isolate the substances already existing in the tissues and fluids, without decomposing them, or altering their nature. A neglect of this rule has been productive of much injury in the pur- suit of organic chemistry; for chemists, in subjecting the animal tissues to the action of acids and alkalies, of prolonged boiling, or of too intense heat, have often obtained, at the end of the analj^sis, many substances which were erroneously described as proximate principles, while they were only the remains of an altered and dis- organized material. Thus, the fibrous tissues, if boiled steadily for thirty- six hours, dissolve, for the most part, at the end of that time, in the boiling water ; and on cooling the whole solution solidifies into a homogeneous, jelly-like substance, which has received the name of gelatine. But this gelatine does not really exist in. the body as a proximate principle, since the fibrous tissue which produces it is not at first soluble, even in boiling water, and its ingredients become altered and converted into a gelatinous matter only by pro- longed ebullition. So, again, an animal substance containing ace- tates or lactates of soda or lime will, upon incineration in the open air, yield carbonates of the same bases, the organic acid having been destroyed, and replaced by carbonic acid ; or sulphur and phospho- rus, in the animal tissue, may be converted by the same means into sulphuric and phosphoric acids, which, decomposing the alkaline carbonates, become sulphates and phosphates. In either case, the analysis of the tissues, so conducted, will be a deceptive one, and useless for all anatomical and physiological purposes, because its real ingredients have been decomposed, and replaced by others, in the process of manipulation. It is in this way that different chemists, operating upon the same animal solid or fluid, by following different plans of analysis, have obtained different results; enumerating as ingredients of the body many artificially formed substances, which are not, in reality, proximate principles, thereby introducing much confusion into phy- siological chemistry. PEOXIMATE PRINCIPLES IN GENERAL. 33 It is to be kept constantly in view, in the examination of an ani- mal tissue or fluid, that the object of the operation is simply the separation of its ingredients from each other, and not their decomposi- tion or ultimate analysis. Only the simplest forms of chemical manipulation should, therefore, be employed. The substance to be examined should first be subjected to evaporation, in order to extract and estimate its water. This evaporation must be conducted at a heat not above 212° F., since a higher temperature would de- stroy or alter some of the animal ingredients. Then, from the dried residue, chloride of sodium, alkaline sulphates, carbonates, and phos- phates may be extracted with water. Coloring matters may be separated by alcohol. Oils may be dissolved out by ether, &c. &c. When a chemical decomposition is unavoidable, it must be kept in sight and afterward corrected. Thus the glyko-cholate of soda of the bile is separated from certain other ingredients by precipitating it with acetate of lead, forming glyko-cholate of lead ; but this is afterward decomposed, in its turn, by carbonate of soda, reproduc- ing the original glyko-cholate of soda. Sometimes it is impossible to extract a proximate principle in an entirely unaltered form. Thus the fibrin of the blood can be separated only by allowing it to coagulate ; and once coagulated, it is permanently altered, and can no longer present all its original characters of fluidity, &c., as it existed beforehand in the blood. In such instances as this, we can only make allowance for an unavoidable difficulty, and be care- ful that the substance suffers no further alteration. By bearing in mind the above considerations, we may form a tolerably correct estimate of the nature and quantity of all of the proximate princi- ples existing in the substance under examination. The manner in which the proximate principles are associated together, so as to form the animal tissues, is deserving of notice. In every animal solid and fluid, there is a considerable number of proximate principles, which are present in certain proportions, and which are so united with each other that the mixture presents a homogeneous appearance. But this union is of a complicated cha- racter ; and the presence of each ingredient depends, to a certain extent, upon that of the others. Some of them, such as the alkaline carbonates and phosphates, are in solution directly in the water. Some, which are insoluble in water, are held in solution by the presence of other soluble substances. Thus, phosphate of lime is held in solution in the urine by the bi-phosphate of soda. In the blood, it is dissolved by the albumen, which is itself fluid by 3 34 PEOXIMATE PRINCIPLES IN GENEEAL. union with the water. The same substance may be fluid in one part of the body, and solid in another part. Thus in the blood and secretions the water is fluid, and holds in solution other sub- stances, both animal and mineral, while in the bones and cartilages it is solid — not crystallized, as in the case of ice or of saline sub- stances which contain water of crystallization, but amorphous and solid, by the fact of its intimate union with the animal and saline ingredients, which are abundant in. quantity, and which are them- selves present in the solid form. Again, the phosphate of lime in the blood is fluid by solution in the albumen; but in the bones it forms a solid substance with the animal matter of the osseous tissue; and yet the union of the two is as intimate and. homo- geneous in the bones as in the blood. A proximate principle, therefore, never exists alone in any part of the body, but is always intimately associated with a number of others by a kind of homo- geneous mixture or solution. Every animal tissue and fluid contains a number of proximate principles which are present, as we have already mentioned, in certain characteristic proportions. Thus, water is present in very large quantity in the perspiration and the saliva, but in very small quantity in the bones and teeth. Chloride of sodium is compara- tively abundant in the blood and deficient in the muscles. On the other hand, chloride of potassium is more abundant in the muscles, less so in the blood. But these proportions, it is important to ob- serve, are nowhere absolute or invariable. There is a great differ- ence in this respect between the chemical composition of an inor- ganic substance and the anatomical constitution of an animal flaid. The former is always constant and definite; the latter is always subject to certain variations. Thus, water is always composed of exactly the same relative quantities of oxygen and hydrogen ; and if these proportions be altered in the least, it thereby ceases to be water, and is converted into some other substance. But in the urine, the proportions of water, urea, urate of soda, phosphates, &c., vary within certain limits in different individuals, and even in the same individual, from one hour to another. This variation, which is almost constantly taking place,, within the limits of health, is characteristic of all the animal solids and fluids ; for they are com- posed of different ingredients which are supplied by absorption or formed in the interior, and which are constantly given up again, under the same or different forms, to the surrounding media by the unceasing activity of the vital processes. Every variation, then, in PROXIMATE PRINCIPLES IN GENERAL. 35 the general condition of the body, as a whole, is accompanied by a corresponding variation, more or less pronounced, in the consti- tution of its different parts. This constitution is consequently of a very different character from the chemical constitution of an oxide or a salt. Whenever, therefore, we meet with the quantita- tive analysis of an animal fluid, in which the relative quantity of its different ingredients is represented in numbers, we must under- stand that such an analysis is always approximative, and not abso- lute. The proximate principles are naturally divided into three differ- ent classes. The first of these classes comprises all the proximate principles which are purely inorganic in their nature. These principles are derived mostly from the exterior. They are found everywhere, in unorganized as well as in organized bodies; and they present them- selves under the same forms and with the same properties in the interior of the animal frame as elsewhere. They are crystallizable, and have a definite chemical composition. They comprise sucb substances as water, chloride of sodium, carbonate and phosphate of lime, &c. The second class of proximate principles is known as crystal- lizable SUBSTANCES OF ORGANIC ORIGIN. This is the name given to them by Robin and Yerdeil,' whose classification of the proxi- mate principles is the best which, has yet been offered. They are crystallizable, as their name indicates, and have a definite chemical composition. They are said to be of "organic origin," because they first make their appearance in the interior of organized bodies, and are not found in external nature as the ingredients of inorganic substances. Such are the different kinds of sugar, oil, and starch. The third class comprises a very extensive and important order of proximate principles, which go by the name of the Organic Substances proper. They are sometimes known as "albuminoid" substances or "protein compounds." The name organic substances is given to them in consequence of the striking difference which exists between them and all the other ingredients of the body. The substances of the second class differ from those of the first by their ' Chimie Anatomique et Physiologique. Paris, 1853. 36 PROXIMATE PRINCIPLES IN GENERAL. exclusively organic origin, but they resemble the latter in their crys- tallizability and their definite chemical composition; in consequence of which their chemical investigation may be pursued in nearly the same manner, and their chemical changes expressed in nearly the same terms. But the proximate principles of the third class are in every respect peculiar. They have an exclusively organic origin ; not being found except as ingredients of living or recently dead animals or vegetables. They have not a definite chemical composition, and are consequently not crystallizable; and the forms which they present, and the chemical changes which they undergo in the body, are such as cannot be expressed by ordinary chemical phraseology. This class includes such substances as albumen, fibrin, casein, &c. PROXIMATE PRINCIPLES OF THE FIRST CLASS. 37 CHAPTER II. PROXIMATE PRINCIPLES OF THE FIRST CLASS. The proximate principles of the first class, or those of an inor- ganic nature, are very numerous. Their most prominent characters have already been stated. They are all crystallizable, and have a definite chemical composition. They are met with extensively in the inorganic world, and form a large part of the crust of the earth. They occur abundantly in the different kinds of food and drink ; and are necessary ingredients of the food, since they are necessary ingredients of the animal frame. Some of them are found universally in all parts of the body, others are met with only in particular regions; but there are hardly any which are not present at the same time in more than one animal solid or fluid. The following are the most prominent of them, arranged in the order of their re- spective importance. 1. Water. — Water is universally present in all the tissues and fluids of the body. It is abundant in the blood and secretions, where its presence is indispensable in order to give them the fluid- ity which is necessary to the performance of their functions ; for it is by the blood and secretions that new substances are intro- duced into the body, and old ingredients discharged. And it is a necessary condition both of the introduction and discharge of substances naturally solid, that they assume, for the time being, a fluid form ; water is therefore an essential ingredient of the fluids, for it holds their solid materials in solution, and enables them to pass and repass through the animal frame. But water is an ingredient also of the solids. For if we take a muscle or a cartilage, and expose it to a gentle heat in dry air, it loses water by evaporation, diminishes in size and weight, and be- comes dense and stiff. Even the bones and teeth lose water by evaporation in this way, though in smaller quantity. In all these solid and semi-solid tissues, the water which they contain is useful 38 PEOXIMATE PRINCIPLES OF THE FIRST CLASS. 37 Bile . . 880 100 Milk . 887 130 Pancreatic juice . 900 550 Urine . 936 750 Lymph. . 960 768 Gastric juice . 975 789 Perspiration . 986 795 Saliva . 995 805 bj giving them the special consistency which is characteristic of them, and which would be lost without it. Thus a tendon, in its natural condition, is white, glistening, and opaque ; and though very strong, perfectly flexible. If its water be expelled by evaporation it becomes yellowish in color, shrivelled, semi-transparent, inflexi- ble, and totally unfit for performing its mechanical functions. The same thing is true of the skin, muscles, cartilages, &c. The following is a list, compiled by Eobin and Yerdeil from various observers, showing the proportion of water per thousand parts, in different solids and fluids: — Quantity of Water in 1,000 parts in Epidermis Teeth Bones Cartilage Muscles . Ligaments Brain Blood Synovial fluid According to the best calculations, water constitutes, in the human subject, between two-thirds and three-quarters of the entire weight of the body. The water which thus forms a part of the animal frame is derived from without. It is taken in the different kinds of drink, and also forms an abundant ingredient in the various articles of food. For no articles of food are taken in an absolutely dry state, but all contain a larger or smaller quantity of water, which may readily be expelled, by evaporation. The quantity of water, therefore, which is daily taken into the system, cannot be ascertained in any case by simply measuring the quantity of drink, but its proportion in the solid food, taken at the same time, must also be determined by experiment, and this ascertained quantity added to that which is taken in with the fluids. The entire quantity of water so intro- duced during twenty-four hours varies according to the researches of M. Barral' from 3f to 4J pounds. After forming a part of the animal solids and fluids, and taking part in the various physical and chemical processes of the body, the water is again discharged ; for its presence in the body, like that of all the other proximate principles, is not permanent, but only ' In Robin and Verdeil, vol. ii. p. 139. CHLORIDE OF SODIUM. 39 temporary. After being taken in with the food and drink, it is associated with other principles in the fluids and solids, passing from the intestine to the blood and from the blood to the tissues and secretions. It afterwards makes its exit from the body, from which it is discharged by four different passages, viz., in a liquid form with the urine and the feces, and in a gaseous form with the breath and the perspiration. Of all the water which is expelled in this way, about 48 per cent, is discharged Avith the urine and feces, ^ and about 52 per cent, by the lungs and skin. This estimate, how- ever, is an average, calculated from the observations of different authors upon different individuals. The absolute and relative amount of water discharged, both in a liquid and gaseous form, varies according to circumstances. There is particularly a com- pensating action in this respect between the kidneys and the skin, so that when the cutaneous perspiration is very abundant the urine is less so, and vice versa. The quantity of water exhaled from the lungs varies also with the state of the pulmonary circulation, and with the temperature and dryness of the atmosphere. The water is not discharged at any time in a state of purity, but is mingled in the urine and feces with saline substances which it holds in solution, and in the cutaneous and pulmonary exhalations with animal vapors and odoriferous substances of various kinds. In the perspiration it is also mingled with saline substances, which it leaves behind on evaporation. 2. Chloeide of Sodium. — This substance is found, like water, throughout the different tissues and fluids of the body. The only exception to this is perhaps the enamel of the teeth, where it has not yet been discovered. Its presence is important in the body, as regulating the phenomena of endosmosis and exosmosis in different parts of the frame. For we know that a solution of common salt passes through animal membranes much less readily than pure water; and tissues which have been desiccated will absorb pure water more abundantly than a saline solution. It must not be sup- posed, however, that the presence or absence of chloride of sodium, or its varying quantity in the animal fluids, is the only condition which regulates their transudation through the animal membranes. The manner in which endosmosis and exosmosis take place in the animal frame depends upon the relative quantity of all the ingre- ' Op. cit., vol. ii. pp. 143 and 145. 40 PROXIilATE PEIXCIPLES OF THE FIRST CLASS. clients of the fluids, as well as on the constitution of the solids them- selves; and the chloride of sodium, as one ingredient among many, influences these phenomena to a great extent, though it does not regulate them exclusively. It exerts also an important influence on the solution of various other ingredients, with which it is associated. Thus, in the blood it increases the solubility of the albumen, and perhaps also of the earthy phosphates. The blood-globules, again, which become dis- integrated and dissolved in a solution of pure albumen, are main- tained in a state of integrity by the presence of a small quantity of chloride of sodium. It exists in the following proportions in several of the solids and fluids :^ — QCA>'TITY OF ChLOEIDE OF SoiJirjI IS 1.000 PAETS !>' THE Muscles . 2 Bile 3.5 Bones 2.5 Blood . 4.5 Milk 1 Mucus . 6 Saliva . 1.5 Aqueous humor '. 11 Urine 3 Vitreous hurnor . 14 In the blood it is rather more abundant than all the other saline ingredients taken together. Since chloride of sodium is so universally present in all parts of the body, it is an important ingredient also of the food. It occurs, of course, in all animal food, in the quantities in which it naturally exists in the corresponding tissues; and in vegetable food also, though in smaller amount. Its proportion in muscular flesh, however, is much less than in the blood and other fluids. Conse- quently, it is not supplied in sufficient quantity as an ingredient of animal and vegetable food, but is taken also by itself as a condi- ment. There is no other substance so universally used by all races and conditions of men, as an addition to the food, as chloride of sodium. This custom does not simply depend on a fancy for grati- fying the palate, but is based upon an instinctive desire for a sub- stance which is necessary to the proper constitution of the tissues and fluids. Even the herbivorous animals are greedy of it, and if freely supplied with it, are kept in a much better condition than when deprived of its use. The importance of chloride of sodium in this respect has been well demonstrated by Boussingault, in his experiments on the fattening of animals. These observations were made upon six ' Robin and Verdeil. CHLOKIDE OF SODIUM. 41 bullocks, selected, as nearly as possible, of the same age and vigor, and subjected to comparative experiment. They were all supplied with an abundance of nutritious food ; but three of them (lot No. 1) received also a little over 500 grains of salt each per day. The remaining three (lot No, 2) received no salt, but in other respects were treated like the first. The result of these experiments is given by Boussingault as follows : — ' "Though salt administered with the food has but little effect in increasing the size of the animal, it appears to exert a favorable influence upon his qualities and general aspect. Until the end of March (the experiment began in October) the two lots experimented on did not present any marked difference in their appearance ; but in the course of the following April, this difference became quite manifest, even to an unpractised eye. The lot No. 2 had then been without salt for six months. In the animals of both lots the skin had a fine and substantial texture, easily stretched and separated from the ribs; but the hair, which was tarnished and disordered in the bullocks of the second lot, was smooth and glistening in those of the first. As the experiment went on, these characters became more marked ; and at the beginning of October the animals of lot No. 2, after going without salt for an entire year, presented a rough and tangled hide, with patches here and there where the skin was entirely uncovered. The bullocks of lot No. 1 retained, on the contrary, the ordinary aspect of stall-fed animals. Their vivacity and their frequent attempts at mounting contrasted strongly with the dull and unexcitable aspect presented by the others. No doubt, the first lot would have commanded a higher price in the market than the second." Chloride of sodium acts also in a favorable manner by exciting the digestive fluids, and assisting in this way the solution of the food. For food which is tasteless, however nutritious it may be in other respects, is taken with reluctance and digested with difficulty; while the attractive flavor which is developed by cooking and by the addition of salt and other condiments in proper proportion excites the secretion of the saliva and gastric juice, and facilitates consequently the whole process of digestion. The chloride of sodium is then taken up by absorption from the intestine, and is deposited in various quantities in different parts of the body. It is discharged with the urine, mucus, cutaneous perspiration, ' Chimie Agricole, p. 271. Paris, 1854. 42 PEOXIMATE PRINCIPLES OF THE FIRST CLASS, &c., in solution in tlie water of these fluids. According to the esti- mates of M. Barral/ a small quantity of chloride of sodium dis appears in the body; since he finds by accurate comparison that all the salt introduced with the food is not to be found in the excreted fluids, but that about one-fifth of it remains unaccounted for. This portion is supposed to undergo a double decomposition in the blood with phosphate of potass, forming chloride of potassium and phos- phate of soda. By far the greater part of the chloride of sodium, however, escapes under its own form with the secretions. 3, Chloride of Potassium. — This substance is found in the muscles, the blood, the milk, the urine, and various other fluids and tissues of the body. It is not so universally present as chlo- ride of sodium, and not so important as a proximate principle. In some parts of the body it is more abundant than the latter salt, in others less so. Thus, in the blood there is more chloride of sodium than chloride of potassium, but in the muscles there is more chloride of potassium than chloride of sodium. This sub- stance is always in a fluid form, by its ready solubility in water, and is easily separated by lixiviation, It is introduced mostly with the food, but is probably formed partly in the interior of the body from chloride of sodium by double decomposition, as already men- tioned. It is discharged with the mucus, the saliva, and the urine. 4. Phosphate of Lime. — This is perhaps the most important of the mineral ingredients of the body next to chloride of sodium. It is met with universally, in every tissue and every fluid. Its quantity, however, varies very much in different parts, as will be seen by the following list: — Quantity of Phosphate of Lime in 1,000 parts in the Enamel of the teeth . . 885 Muscles . . . .2.5 Dentine . . . .643 Blood . . . .0.3 Bones .... 550 Gastric juice . • • 0.4 Cartilages ... 40 It occurs also under different physical conditions. In the bones, teeth, and cartilages it is solid, and gives to these tissues the resist- ance and solidity which are characteristic of them. The calcareous salt is not, however, in these instances, simply deposited mechani- cally in the substance of the bone or cartilage as a granular powder, but is intimately united with the animal matter of the tissues, like ' In Robin and Verdeil, op. cit., vol. ii. p. 193. PHOSPHATE OF LIME. 43 a coloring matter in colored glass, so as to present a more ov less homogeneous appearance. It can, however, be readily dissolved out by maceration in dilute muriatic acid, leaving behind the animal substance, which still retains the original form of the bone or cartilage. It is not, therefore, united with the animal matter so as to lose its identity and form a new chemical substance, as where an acid combines with an alkali to form a salt, but in the same manner as salt unites with water in a saline solution, both sub- stances retaining their original character and composition, but so intimately associated that they cannot be separated by mechanical means. In the blood, phosphate of lime is in a liquid form, notwithstand- ing its insolubility in water and in alkaline fluids, being held in solution by the albuminous matters of the circulating fluid. In the urine, it is retained in solution by the bi-phosphate of soda. In all the solid tissues it is useful by giving to them their proper consistence and solidity. For example, in the enamel of the teeth, the hardest tissue of the body, it predominates very much over the animal matter, and is present in greater abun- dance there than in any other part of the frame. In the dentine, a softer tissue, it is in somewhat smaller quantity, and in the bones smaller still ; though in the bones it continues to form more than one-half the entire mass of the osseous sub- stance. The importance of phosphate of lime in communicating to bones their natural stiffness and consistency may be readily shown by the altera- tion which they suffer from its removal. If a long bone be macerated in dilute muriatic acid, the earthy salt, as already mentioned, is entirely dis- solved out, when the bone loses its rigidity, and may be bent or twisted in any direction without breaking. (Fig. 1.) Whenever the nutrition of the bone during life is interfered with from any pathological cause, so that its phosphate of lime becomes deficient in amount, a softening of the osseous tissue is the consequence, by which the bones yield to external pressure, and become more or less distorted. (Osteo- malakia.) After forming, for a time, a part of the tissues and fluids, the PlBtTLA TIED IN A KNOT, after macera- tion in a dilute acid. (From a specimen in tlie museum of the College of Physicians and Surgeons.) 44 PEOXIMATE PEIN'CIPLES OP THE FIEST CLASS. phosphate of lime is discharged from the body by the urine, the perspiration, mucus, &c. Much the larger portion is discharged by the urine. A small quantity also occurs in the feces, but that is pro- bably only the superfluous residue of what is taken in with the food. 5. Caebonate of Lime. — Carbonate of lime is to be found in the bones, and sometimes in the urine. The concretions of the internal ear are almost entirely formed of it. It very probably occurs also in the blood, teeth, cartilages, and sebaceous matter; but its presence here is not quite certain, since it may have been produced from the lactate, or other organic combination, by the process of incineration. In the bones, it is in much smaller quan- tity than the phosphate. Its solubility in the blood and the urine 13 accounted for by the presence of free carbonic acid, and also of chloride of potassium, both of which substances exert a soluble action on carbonate of lime. 6. Caeboxate of Soda. — This substance exists in the bones, blood, saliva, lymph, and urine. As it is readily soluble in water, it naturally assumes the liquid form in the animal fluids. It is important principally as giving to the blood its alkalescent reaction, by which the solution of the albumen is facilitated, and various other chemico-physiological processes in the blood accomplished. The alkalescence of the blood is, in fact, necessary to life ; for it is found that, in the living animal, if a mineral acid be gradually injected into the blood, so dilute as not to coagulate the albumen, death takes place before its alkaline reaction has been completely neutralized.^ The carbonate of soda of the blood is partly introduced as such with the food ; but the greater part of it is formed within the body by the decomposition of other salts, introduced with certain fruits and vegetables. These fruits and vegetables, such as apples, cherries, grapes, potatoes, &c., contain malates, tartrates, and citrates of soda and potass. Now, it has been often noticed that, after the use of acescent fruits and vegetables containing the above salts, the urine becomes alkaline in reaction from the presence of the alkaline carbonates. Lehmann^ found, by experiments upon his own person, that, within thirteen minutes after taking half an ounce ' Cl. Bernard. Lectures on tlie Blood ; reported by W. F. Atlee, M. D. Pliila- delpMa, 1854, p. 31. ^ Physiological Cliemistry. Philadelpliia ed., vol. i. p. 97. PHOSPHATES OF MAGNESIA, SODA, AND POTASS, 45 of lactate of soda, the urine had an alkaline reaction. He also ob- served that, if a solution of lactate of soda were injected into the jugular vein of a dog, the urine became alkaline at the end of five, or, at the latest, of twelve minutes. The conversion of these salts into carbonates takes place, therefore, not in the intestine but in the blood. The same observer' found that, in many persons living on a mixed diet, the urine became alkaline in two or three hours after swallowing ten grains of acetate of soda. These salts, therefore, on being introduced into the animal body, are decomposed. Their organic acid is destroyed and replaced by carbonic acid ; and they are then discharged under the form of carbonates of soda and potass. 7. Carbonate of Potass. — -This substance occurs in very nearly tbe same situations as the last. In the blood, however, it is in smaller quantity. It is mostly produced, as above stated, by the decomposition of the malate, tartrate, and citrate, in the same manner as the carbonate of soda. Its function is also the same as that of the soda salt, and it is discharged in the same manner from the body. 8, Phosphates of Magnesia, Soda, and Potass. — All these substances exist universally in all the solids and fluids of the body, but in very small quantity. The phosphates of soda and potass are easily dissolved in the fluids, owing to their ready solubility in water. The phosphate of magnesia is held in solution in the blood by the alkaline chlorides and phosphates ; in the urine, by the acid phosphate of soda. A peculiar relation exists between the alkaline phosphates and carbonates in different classes of animals. For while the fluids of carnivorous animals contain a preponderance of phosphates, those of the herbivora. contain a preponderance of the carbonates: a peculiarity readily understood when we recollect that muscular flesh and the animal tissues generally are comparatively abundant in phosphates ; while vegetable substances abound in salts of the organic acids, which give rise, as already described, by decomposi- tion in the blood, to the alkaline carbonates. The proximate principles included in the above list resemble ' Physiological Chemistry, vol. ii. p. 130. 46 PROXIMATE PRIXCIPLES OF THE FIRST CLASS. each other not only in their inorganic origin, their crystallizability, and their definite chemical composition, but also in the part which they take in the constitution of the animal frame. They are distinguished in this respect, first, by being derived entirely from without. There are a few exceptions to this rule; as, for example, in the case of the alkaline carbonates, which partly originate in the body from the decomposition of malates, tartrates, &c. These, however, are only exceptions ; and in general, the proximate prin- ciples belonging to the first class are introduced with the food, and taken up by the animal tissues in precisely the same form under which they occur in external nature. The carbonate of lime in the bones, the chloride of sodium in the blood and tissues, are the same substances which are met with in the calcareous rocks, and in solution in sea water. They do not suffer any chemical alteration in becoming constituent parts of the animal frame. They are equally exempt, as a general rule, from any alteration while they remain in the body, and during their passage through it. The exceptions to this rule are very few ; as, for example, where a small part of the chloride of sodium suffers double decomposition with phosphate of potass, giving rise to chloride of potassium and phosphate of soda ; or where the phosphate of soda itself gives up a part of its base to an organic acid (uric), and is converted in this way into a bi-phosphate of soda. Nearly the whole of these substances, finally, are taken up un- changed from the tissues, and discharged unchanged with the excre- tions. Thus we find the phosphate of lime and the chloride of so- dium, which were taken in with the food, discharged again under the same form in the urine. They do not, therefore, for the most -part, participate directly in the chemical changes going on in the body ; but only serve by their presence to enable those changes- to be accomplished in the other ingredients of the animal frame, which are necessary to the process of nutrition. PROXIMATE PRINCIPLES OF THE SECOND CLASS, 47 CHAPTER III. PROXIMATE PRINCIPLES OF THE SECOND CLASS. The proximate principles belonging to tlie second class are divided into three principal groups, viz. : starch, sugar, and oil. Thej are distinguished, in the first place, bj their organic origin. Unlike the principles of the first class, they do not exist in external nature, but are only found as ingredients of organized bodies. They exist both in animals and in vegetables, though in somewhat different proportions. All the substances belonging to this class have a definite chemical composition ; and are further distinguished by the fact that they are composed of oxygen, hydro- gen, and carbon alone, without nitrogen, whence they are sometimes called the " non-nitrogenous" substances. 1. Starch (C^JI^fi^^). The first of these substances seems to form an exception to the general rule in a very important particular, viz., that it is not crystallizable. Still, since it so closely resembles the rest in all its general properties, and since it is easily converti- ble into SQgar, which is itself crystallizable, it is naturally included in the second class of proximate principles. Though not crystal- lizable, furthermore, it still does assume a distinct form, by which it differs from substances that are altogether amorphous. Starch occurs in some part or other of almost all the flowering plants. It is very abundant in corn, wheat, rye, oats, and rice, in the parenchyma of the potato, in peas and beans, and in most vege- table substances used as food. It constitutes almost entirely the different preparations known as sago, tapioca, arrowroot, &c., which are nothing more than varieties of starch, extracted from different species of plants. The following is a list showing the percentage of starch occurring in different kinds of food : — ^ ' Pereira on Food and Diet, p. 39. New York, 1843. 48 PKOXIMATE PRINCIPLES OF THE SECOND CLASS. Quantity of Starch IN 100 PARTS IN 85.07 Wheat flour 56.50 80.92 Iceland moss 44.60 67.18 Kidney bean 35.94 61.07 Peas . 32.45 59.00 Potato 15.70 Rice . Maize Barley meal Rye meal . Oat meal . When purified from foreign substances, starch is a white, liglit powder, which gives rise to a peculiar crackling sensation when rubbed between the fingers. ■^"S- ^- It is not amorphous, as we have already stated, but is composed of solid granules, which, while they have a general resemblance to each other, differ somewhat in va- rious particulars. The starch grains of the potato (Fig. 2), vary considerably in size. The smallest have a diameter of lohoo, the largest ^^o of an inch. They are irregu- larly pear-shaped inform, and are marked by concentric la- minse, asifthematter of which they are composed had been deposited in successive layers. At one point on the surface of every starch grain, there is a minute pore or depression, called the hilus, Grains of Potato Starch. Fig. 3. Starch Grains of Bermcda Arrowroot. around which the circular markings are arranged in a concentric form. The starch granules of arrowroot (Fig. 3) are gene- rally smaller, and more uni- form in size than those of the potato. They vary from 2o'oo to 5-^0 of an inch in diameter. They are elongated and cylindrical in form, and the concentric markings are less distinct than in the pre- ceding variety. The hilus STARCH. 49 Starch Grains op Wheat Flour. bas here sometimes the form of a circular pore, and sometimes that of a transverse fissure or slit. The grains of wheat starch (Fig. 4) are still smaller than those t)f arrowroot. They vary from TOO 00 to ^lo of an inch in diameter. They are nearly circular in form, with a round or transverse hilus, but without any distinct appearance of lamination. Many of them are flattened or compressed laterally, so that they present a broad surface in one position, and a narrow edge when viewed in the opposite direction. The starch grains of In- dian corn (Fig. 5), are of nearly the same size with those of wheat flour. They are somewhat more irregular and angular in shape ; and are often marked with crossed or radiating lines, as if from partial fracture. Starch is also an ingre- dient of the animal body. It was first observed by Purkinje, and afterward by Koiliker, ' that certain bodies are to be found in the interior of the brain, about the lateral ventricles, in the fornix, septum lucidum, and other parts which present a cer- tain resemblance to starch grains, and which have there- fore been called "corpora amylacea." Subsequently Virchow^ corroborated the above observations, and ascertained the corpora amylacea to be ' Handbuch der Gewebelehre, Leipzig, 1852, p. 311. * In American Journal Med. Sci., April, 1854, p. 466. Starch Grains of Indian Corn. 50 PROXIMATE PRINCIPLES OF THE SECOND CLASS. Starch Grains from Wall of Lateral Ventricle ; from a woman aged 35. really substances of a starchy nature ; since they exhibit the usual chemical reactions of vegetable starch. The starch granules of the human brain (Fig. 6), are transparent and colorless, like those from plants. They refract the light strongly, and vary in size from 4^Vu to yyVo of an inch. Their average is ygViy of an inch. They are some- times rounded or oval, and sometimes angular in shape. They resemble considerably in appearance the starch granules of Indian corn. The largest of them present a very faint concentric lamina- tion, but the greater number are destitute of any such appearance. They have nearly always a distinct hilus, which is sometimes circular and sometimes slit-shaped. They are also often marked with delicate radiating lines and shadows. On the addition of iodine, they become colored, first purple, afterward of a deep blue. They are less firm in consistency than vegetable starch grains, and can be more readily disintegrated by pressing or rubbing them upon the glass. Starch, derived from all these different sources, has, so far as known, the same chemical composition, and may be recognized by the same tests. It is insoluble in cold water, but in boiling water its granules first swell, become gelatinous and opaline, then fuse together, and finally liquefy altogether, provided a sufficient quantity of water be present. After that, they cannot be made to resume their original form, but on cooling and drying merely solidify into a homogeneous mass or paste, more or less consistent, according to the quantity of water which remains in union witb it. The starch is then said to be amorphous or "hydrated." By this process it is not essentially altered in its chemical properties, but only in its physical condition. Whether in granules, or in solution, or in an amorphous and hydrated state, it strikes a deep blue color on the addition of free iodine. Starch may be converted into sugar by three different methods. First, by boiling with a dilute acid. If starch be boiled with dilute SUGAR. 51 nitric, sulphuric, or muriatic acid during thirty-six hours, it first changes its opalescent appearance, and becomes colorless atid trans- parent; losing at the same time its power of striking a blue color with iodine. After a time, it begins to acquire a sweet taste, and is finally altogether converted into a peculiar species of sugar. Secondly, by contact with certain animal and vegetable sub- stances. Thus, boiled starch, mixed with human saliva and kept at the temperature of 100° F., is converted in a few minutes into sugar. Thirdly, by the processes of nutrition and digestion in animals and vegetables. A large part of the starch stored up in seeds and other vegetable tissues is, at some period or other of the growth of the plant, converted into sugar by the molecular changes going on in the vegetable fabric. It is in this way, so far as we know, that all the sugar derived from vegetable sources has its origin. Starch, as a proximate principle, is more especially important as entering largely into the composition of many kinds of vegetable food. With these it is introduced into the alimentary canal, and there, during the process of digestion, is converted into sugar. Consequently, it does not appear in the blood, nor in any of the secreted fluids. 2. Sugar, — This group of proximate principles includes a con- siderable number of substances, which differ in certain minor details, while they resemble each other in the following particulars: They are readily soluble in water, and crystallize more or less perfectly on evaporation ; they have a distinct sweet taste ; and finally, by the process of fermentation, they are converted into alcohol and carbonic acid. These substances are derived from both animal and vegetable sources. Those varieties of sugar which are most familiar to us are the following six, three of which are of vegetable and three of animal origin. fCane sugar, r Milk sugar, ^ Animal ^ . Grape sugar, < Liver sugar, Sugar of starch. ° V Sugar of honey. The cane and grape sugars are held in solution in the juices of the plants from which they derive their name. Sugar of starch, or glucose^ is produced by boiling starch for a long time with a dilute acid. Liver sugar and the sugar of milk are produced in the tissues of the liver and the mammary gland, and the sugar oi 52 PROXIMATE PRINCIPLES OF THE SECOND CLASS. honey is prepared in some way by the bee from materials of vege- table origin. These varieties cliflfer but little in their ultimate chemical compo- sition. The following formulae have been established for three of them. Cane sugar ......= C24H220^ Milk siigar = Cj^Hj^Oj^ Glucose = C^jH28023 Cane sugar is sweeter than most of the other varieties, and more soluble in water. Some sugars, such as liver sugar and sugar of honey, crystallize only with great difficulty; but this is probably owing to their being mingled with other substances, from which it is difficult to separate them completely. If they could be obtained in a state of purity, they would doubtless crystallize as perfectly as cane sugar. The different sugars vary also in the readiness with which they undergo fermentation. Some of them, as grape sugar and liver sugar, enter into fermentation very promptly ; others, such as milk and cane sugar, with considerable difficulty. The above are not to be regarded as the only varieties of sugar existing in nature. On the contrary, it is probable that nearly every different species of animal and vegetable produces a distinct kind of sugar, differing slightly from the rest in its degree of sweet- ness, its solubility, its crystallization, its aptitude for fermentation, and perhaps in its elementary composition. Nevertheless, there is so close a resemblance between them that they are all properly regarded as belonging to a single group. The test most commonly employed for detecting the presence of suffar is that known as Trommer's test. It depends upon the fact that the saccharine substances have the power of reducing the persalts of copper when heated with them in an alkaline solution. The test is applied in the following manner : A very small quantity of sulphate of copper in solution should be added to the suspected liquid, and the mixture then rendered distinctly alkaline by the addition of caustic potass. The whole solution then takes a deep blue color. On boiling the mixture, if sugar be present, the insoluble suboxide of copper is thrown down as an opaque red, yellow, or orange colored deposit ; otherwise no change of color takes place. This test requires some precautions in its application. In the first place, it is hot applicable to all varieties of sugar. Cane sugar, for example, when pure, has no power of reducing the salts SUGAR. 53 of copper, even when present in large quantity. Maple sugar, also, which resembles cane sugar in some other respects, reduces the copper, in Trommer's test, but slowly and imperfectly. Beet- root sugar, according to Bernard, presents the same peculiarity. If these sugars, however, be boiled for two or three minutes with a trace of sulphuric acid, they become converted into glucose, and acquire the power of reducing the salts of copper. Milli sugar, liver sugar, and sugar of honey, as well as grape sugar and glucose all act promptly and perfectly with Trommer's test in their natural condition. Secondly, care must be taken to add to the suspected liquid only a small quantity of sulphate of copper, just sufficient to give to the whole a distinct blue tinge, after the addition of the alkali. If a larger quantity of the copper salt be used, the sugar in solution may not be sufficient to reduce the whole of it ; and that which remains as a blue sulphate will mask the yellow color of the sub- oxide thrown down as a deposit. By a little care, however, in managing the test, this source of error may be readily avoided. Thirdly, there are some albuminous substances which have the power of interfering with Trommer's test, and prevent the reduc- tion of the copper, even when sugar is present. Certain animal matters, to be more particularly described hereafter, which are liable to be held in solution in the gastric juice, have this effect. This source of error may be avoided, and the substances in ques- tion eliminated when present, by treating the suspected fluid with animal charcoal, or by evaporating and extracting it with alcohol before the application of the test. A less convenient but somewhat more certain test for sugar is that o^ fermentation. The' saccharine fluid is mixed with a little yeast, and kept at a temperature of 70° to 100° F. until the fer- menting process is completed. By this process, as already men-J tioned, the sugar is converted into alcohol and carbonic acid. The gas, which is given off in minute bubbles during fermentation, should be collected and examined. The remaining fluid is purified by distillation and also subjected to examination. If the gas be found to be carbonic acid, and the remaining fluid contain alcohol, there can be no doubt that sugar was present at the commencement of the operation. The following list shows the percentage of sugar in various articles of food.' ' Pereira, op. cit., p. 55. 54 PROXIMATE PRINCIPLES OF THE SECOND CLASS. Quantity of Sugar in 100 parts in Figs . 62.50 Wheat flour 4.20 to 8.48 Cherries 18.12 Rye meal . 3.28 Peaches 16.48 Indian meal 1.45 Tamarinds 12.50 Peas . 2.00 Pears 11.52 Cow's milk 4.77 Beets 9.00 Ass's milk 6.08 Sweet almonds 6.00 Human milk 6.50 Barley meal . 5.21 Beside the sugar, therefore, which is taken into the alimentary canal in a pure form, a large quantity is also introduced as an in- gredient of the sweet-flavored fruits and vegetables. All the starchy substances of the food are also converted into sugar in the process of digestion. Two of the varieties of sugar, at least, originate in the interior of the body, viz., sugar of milk and liver sugar. The former exists in a solid form in the substance of the mammary gland, from which it passes in solution into the milk. The liver sugar is found in the substance of the liver, and almost always also in the blood of the hepatic veins. The sugar which is introduced with the food, as well as that which is formed in the liver, disappears by decomposition in the animal fluids, and does not appear in any of the excretions. 8. Fats. — These substances, like the sugars, are derived from both animal and vegetable sources. There are three principal varieties of them, which may be considered as representing the class, viz : — Oleine ' . . = Cg^ H^^ 0,5 Margarine ......= C^g H^j 0,2 Stearine ......= Ci^gH^iO^y The principal difference between the oleaginous and saccharine substances, so far as regards their ultimate chemical composition, is that in the sugars the oxygen and hydrogen always exist together in the proportion to form water; while in the fats the proportions of carbon and hydrogen are nearly the same, but that of oxygen is considerably less. The fats are all fluid at a high temperature, but assume the solid form on cooling. Stearine, which is the most solid of the three, liquefies only at 148° F.; margarine at 118° F.; while oleine remains fluid considerably below 100° F., and even very near the freezing point of water. The fats are all insoluble in water, but readily soluble in ether. When treated with a solu- tion of a caustic alkali, they are decomposed, and as the result of FATS. 55 the decomposition tliere are formed two new bodies ; first, glycerine, which is a neutral fluid substance, and secondly, a fatty acid, viz : oleic, margaric, or stearic acid, corresponding to the kind of fat which has been used in the experiment. The glycerine remains in a free state, while the fatty acid unites with the alkali employed, forming an oleate, margarate, or stearate. This combination is termed a soap^ and the process by which it is formed is called saponification. This process, however, is not a simple decomposition of the fatty body, since it can only take place in the presence of water; several equivalents of which unite with the elements of the fatty body, and enter into the composition of the glycerine, &c., so that the fatty acid and the glycerine together weigh more than the original fatty substance which was decomposed. It is not proper, therefore, to regard an oleaginous body as formed by the union of a fatty acid with glycerine. It is formed, on the contrary, in all pro- bability, by the direct combination of its ultimate chemical elements. The different kinds of oil, fat, lard, suet, &c., contain the three oleaginous matters mentioned above, mingled together in different proportions. The more solid fats contain a larger quantity of stearine and margarine; the less consistent varieties, a larger propor- tion of oleine. Neither of the oleaginous matters, stearine, mar- garine, or oleine, ever occur separately ; but in every fatty substance they are mingled together, so that the more fluid of them hold in solution the more solid. Generally speaking, in the living body, these mixtures are fluid or nearly so ; for though both stearine and margarine are solid, when pure, at the ordinary tem- perature of the body, they are held in solution, during life, by the oleine with which they are associated. After death, however, as the body cools, tlie stearine and mar- garine sometimes separate from the mixture in a crys- talline form, since the oleine can no longer hold in solution so large a quantity of them as it had dissolved at a higher temperature. Steakine crystallized from a Warm Solution in Oleine. 56 PEOXIMATE PEINCIPLES OF THE SECOND CLASS. These substances crystallize in very slender needles, which are sometimes straight, but more often somewhat curved or wavy in their outline. (Fig. 7.) They are always deposited in a more or less radiated form ; and have sometimes a very elegant, branched, or arborescent arrange- ment. When in a fluid state, the fatty substances present themselves under the form of drops or globules, which vary indefi- nitely in size, but which may be readily recognized by their optical properties. They are circular in shape, and have a faint amber color, distinct in the larger globules, less so in the smaller. They have a sharp, well defined outline (Fig. 8); and as they refract the light strongly, and act therefore as double convex lenses, they present a brilliant centre, surrounded by a dark border. These marks will generally be sufficient to distinguish them under the microscope. The following list shows the percentage of oily matter present in various kinds of animal and vegetable food.* Oleaginous Principles of Human Fat. Stearine and Margarine crystallized; Oleine Fluid. Quantity of Fat in 100 parts in Filberts . Walnuts Cocoa-nuts Olives . Linseed . Indian corn Yolk of eggs 60.00 50.00 47.00 32.00 22.00 9.00 28.00 Ordinary meat Liver of the ox Cow's milk . Human milk . Asses' milk . Goat's milk . 14.30 3.89 3.13 3.55 0.11 3.32 The oleaginous matters present a striking peculiarity as to the form under which they exist in the animal body ; a peculiarity which distinguishes them from all the other proximate principles. The rest of the proximate principles are all intimately associated together by molecular union, so as to form either clear solutions or Pereira, op. cit., p. 81. FATS. 67 homogeneous solids. Thus, the sugars of the blood are in solution in water, in company with the albumen, the phosphate of lime, chloride of sodium, and the like ; all of them equally distributed throughout the entire mass of the fluid. In the bones and car- tilages, the animal matters and the calcareous salts are in similarly intimate union with each other ; and in every other part of the body the animal and inorganic ingredients are united in the same Avay. But it is different with the fats. For, while the three prin- cipal varieties of oleaginous matter are always united with each other, they are not united with any of the other kinds of proximate principles; that is, with water, saline substances, sugars, or albu- minous matters. Almost the only exception to this is in the nerv- ous tissue; in which, according to Robin and Verdeil, the oily matters seem to be united with an albuminoid substance. Another exception is, perhaps, in the bile ; since some of the biliary salts have the power of dissolving a certain quantity of fat. Every- where else, instead of forming a homogeneous solid or fluid with the other proximate principles, the oleaginous matters are found in distinct masses or globules, which are suspended in serous fluids, interposed in the interstices between the anatomical elements, in- cluded in the interior of cells, or deposited in the substance of fibres or membranes. Even in the vegetable tissues, the oil is always deposited in this manner in distinct drops or granules. Owing to this fact, the oils can be easily extracted from the organized tissues by the employment of simply mechanical pro- cesses. The tissues, animal or vegetable, are merely cut into small pieces and subjected to pressure, by which the oil is forced out from the parts in which it was entangled, and separated, without any further manipulation, in a state of purity. A moderately elevated temperature facilitates the operation by increasing the fluidity of the oleaginous matter ; but no other chemical agency is required for its separation. Under the microscope, also, the oil drops and granules can be readily perceived and distinguished from the remaining parts of the tissue, and can, moreover, be easily re- cognized by the dissolving action of ether, which acts upon them, as a general rule, without attacking the other proximate principles. Oils are found, in the animal body, most abundantly in the adipose tissue. Here they are contained in the interior of the adipose vesicles, the cavities of which they entirely fill, in a state 58 PEOXIMATE PRINCIPLES OF THE SECOND CLASS. Human Adipose Tissue. of health. These vesicles are transparent, and have a somewhat angular form, owing to their mutual compression. (Fig. 9.) They vary in diameter, in the hu- man subject, from ^^^ to ^^j^ of an inch, and are composed of a thin, structureless ani- mal membrane, forming a closed sac, in the interior of which the oily matter is con- tained. There is here, ac- cordingly, no union whatever of the oil with the other proximate principles, but only a mechanical inclusion of them by the walls of the vesicles. Sometimes, when emaciation is going on, the oil partially disappears from the cavity of the adipose vesicle, and its place is taken by a watery serum ; but the serous and oily fluids always remain distinct, and occupy different parts of the cavity of the vesicle. In the chyle, the oleaginous matter is in a state of emulsion or suspension in the form of minute particles in a serous fluid. Its subdivision is here more com- Fig. 10. plete, and its molecules more minute than anywhere else in the body. It presents the ap- pearance of a fine granular dust, which has been known by the name of the " molecu- lar base of the chyle." A few of these granules are to be seen which measure xo^oo of an inch in diameter ; but they are generally much less than this, and the greater part are so small that they cannot be accurately measured. (Fig. 10.) For the same reason they do not present the bril- liant centre and dark border of the larger oil-globules ; but appear Chyle, from commencemeut of Thoracic Duct, from the Dog. FATS. 59 o° o%^. by transmitted light only as minute dark granules. The white color and opacity of the chyle, as of all other fatty emulsions, depend upon this molecular condition of the oily ingredients. The albumen, salts, &c., which are in intimate union with each other, and in solution in the water, would alone make a colorless and transparent fluid; but the oily matters, suspended in distinct par tides, which have a different refractive power from the serous fluid, interfere with its transparency and give it the white color and opaque appearance which are characteristic of emulsions. The oleasfinous nature of these particles is readily shown by their solubility in ether. In the milk, the oily matter occurs in larger masses than in the chyle. In cow's milk (Fig. 11), these oil-drops, or " milk-globules," are not quite fluid, but have a pasty con- sistency, owing to the large quantity of margarine which they contain, in proportion to the oleine. When forcibly amalgamated with each other and collected into a mass by prolonged beating or churning, they con- stitute butter. In cow's milk, the globules vary somewhat in size, but their average diameter is 40V0 of ^^ inch. They are simply suspended in the serous fluid of the milk, and are not covered with any albuminous mem- brane. In the cells of the laryn- geal, tracheal, and costal car- tilages (Fig. 12), there is always more or less fat de- posited in the form of round- ed globules, somewhat simi- , , PI •11 Cells OF Costal Cartilages, containing Oil-Globules. lar to those 01 the milk. Human. Globules of Cow's Milk. Fia;. 12. 60 PROXIMATE PRINCIPLES OF THE SECOND CLASS. Hepatic Cells. Human. In the glandular cells of the liver, oil occurs constantly, in a state of health. It is here deposited in the substance of the cell (Fig. 13), generally in smaller globules than the preceding. In some cases of disease, it accumulates in excessive quantity, and produces the state known as fatty degene- ration of the liver. This is consequently only an ex- aggerated condition of that which normally exists in health. In the carnivorous ani- mals, oil exists in considera- ble quantity in the convo- luted portion of the urini- ferous tubules. (E'ig. 14.) It is here in the form of granules and rounded drops, which some- times appear to fill nearly the whole calibre of the tubules. It is found also in the secreting cells of the sebaceous and other glandules, deposited in the same manner as in those of the liver, but in smaller quantity. It exists, beside, in large proportion, in a granular form, in the secre- tion of the sebaceous gland- ules. It occurs abundantly in the marrow of the bones, both under the form of free oil-globules and inclosed in the vesicles of adipose tissue. It is found in considerable quantity in the substance of the yellow wall of the corpus luteum, and is the immediate cause of the peculiar color of this body. It occurs also in the form of granules and oil-drops in the muscular fibres of the uterus (Fig. 15), in which it begins to be Uriniferods Tubules of Dog, from Cortical Portion of Kidney. FATS. 61 MrscnLAR Fibres of Human TJterits three weeks after parturition. deposited sood after delivery, and where it continues to be present during the whole period of the resorption or involution of this organ. lu all these instances, the oleaginous matters remain distinct in form and situation from the other ingredients of the ani- ^ig- 15. mal frame, and are only me- chanically entangled among its fibres and cells, or im- bedded separately in their interior. A large part of the fat which is found in the body may be accounted for by that which is taken in with the food, since oily matter occurs in both animal and vegetable substances. Fat is, however, formed in the body, independ- ently of what is introduced with the food. This im- portant fact has been definitely ascertained by the experiments of MM. Dumas and Milne-Edwards on bees,* M. Persoz on geese,^ and finally by those of M. Boussingault on geese, ducks, and pigs.^ The observers first ascertained the quantity of fat existing in the whole body at the commencement of the experiment. The animals were then subjected to a definite nutritious regimen, in which the quantity of fatty matter was duly ascertained by analysis. The experiments lasted for a period varying, in different instances, from thirty-one days to eight months; after which the animals were killed and all their tissues examined. The result of tbese investi- gations showed that considerably more fat had been accumulated by the animal during the course of the experiment than could be accounted for by that which existed in the food; and placed it beyond a doubt that oleaginous substances may be, and actually are, formed in the interior of tbe animal body by the decomposition or metamorphosis of other proximate principles. It is not known from what proximate principles the fat is pro- duced, when it originates in this way in the interior of the body. Particular kinds of food certainly favor its production and accu- ' Annales de China, et de Phys., 3d series, vol. xiv. p. 400. ^ Chimie Agdcole, Paris, 1854. 2 Ibid., p. 408. 62 proxi:mate peixciples of the secoxd class, mulation to a considerable degree. It is well known, for instance, that in sugar-growing countries, as in Louisiana and the West Indies, during the few weeks occupied in gathering the cane and extracting the sugar, all the negroes employed on the plantations, and even the horses and cattle, that are allowed to feed freely on the saccharine juices, grow remarkably fat; and that they again lose their superabundant flesh when the season is past. Even in these instances, however, it is not certain whether the saccharine substances are directly converted into fat, or whether they are first assimilated and only afterward supply the materials for its production. The abundant accumulation of fat in certain regions of the body, and its absence in others; and more particularly its constant occurrence in certain situations to which it could not be transported by the blood, as for example the interior of the cells of the costal cartilages, the substance of the muscular fibres of the uterus after parturition, &c,, make it probable that under ordinary conditions the oily matter is formed by decomposition of the tissues upon the very spot where it subsequently makes its appearance. In the female during lactation a large part of the oily matter in- troduced with the food, or formed in the body, is discharged with the milk, and goes to the support of the infant. But in the female in the intervals of lactation, and in the male at all times, the oily matters almost entirely disappear by decomposition in the interior of the body ; since the small quantity which is discharged with the sebaceous matter by the skin bears only an insignificant proportion to that which is introduced daily with the food. The most important characteristic, in a physiological point of view, of the proximate principles of the second class, relates to their origin and their final destination. Not only are they all of a purely organic origin, making their appearance first in the interior of vege- tables ; but the sugars and the oils are formed also, to a certain ex- tent, in the bodies of animals; continuing to make their appearance when no similar substances, or only an insufficient quantity of them, have been taken with the food. Furthermore, when introduced with the food, or formed in the body and deposited in the tissues, these substances do not reappear in the secretions. They, therefore, for the most part disappear by decomposition in the interior of the body. They pass through a series of changes by which their es- sential characters are destroyed ; and they are finally replaced in the circulation by other substances, which are discharged with the excreted fluids. PROXIMATE PRINCIPLES OF THE THIRD CLASS. 63 CHAPTER IV. PROXIMATE PRINCIPLES OF THE THIRD CLASS. The substances belonging to this class are very important, and form by far the greater part of the entire mass of the body. They are derived both from animal and vegetable sources. They have been known by the name of the "protein compounds" and the " albuminoid substances." The name organic substances was given to them by Robin and Verdeil, by whom their distinguishing pro- perties were first accurately described. They have not only an organic origin, in common with the proximate principles of the second class, but their chemical constitution, their physical struc- ture and characters, and the changes which they undergo, are all so different from those met with in any other class, that the term "or- ganic substances" proper, appears particularly appropriate to them. Their first peculiarity is that they are not crystallizable. They always, when pure, assume an amorphous condition, which is some- times solid (organic substance of the bones), sometimes fluid (albu- men of the blood), and sometimes semi-solid in consistency, midway between the solid and fluid condition (organic substance of the mus- cular fibre). Their chemical constitution differs from that of bodies of the second class, first in the fact that they all contain the four chemical elements, oxygen, hydrogen, carbon, and nitrogen ; while the starches, sugars, and oils are destitute of the last named ingredient. The or- ganic matters have therefore been sometimes known by the name of the " nitrogenous substances," while the sugars, starch, and oils have been called "non-nitrogenous." Some of the organic matters, viz., albumen, fibrin, and casein, contain sulphur also, as an ingre- dient; and others, viz., the coloring matters, contain iron. The remainder consist of oxygen, hydrogen, carbon, and nitrogen alone. The most important peculiarity, however, of the organic sub- stances, relating to their chemical composition, is that it is not defi- nite. That is to say, they do not always contain precisely the same 64 PEOXIMATE PEIISrCIPLES OF THE THIED CLASS. proportions of oxygen, hydrogen, carbon, and nitrogen ; but the re- lative quantities of these elements vary within certain limits, in different individuals and at different times, without modifying, in any essential degree, the peculiar properties of the animal matters which they constitute. This fact is altogether a special one, and characteristic of organic substances. No substance having a definite chemical composition, like phosphate of lime, starch, or olein, can suffer the slightest change in its ultimate constitution without being, by that fact alone, totally altered in its essential properties. If phosphate of lime, for example, were to lose one or two equivalents of oxygen, an entire destruction of the salt would necessarily result, and it would cease to be phosphate of lime. For its properties as a salt depend entirely upon its ultimate chemical constitution; and if the latter be changed in any way, the former are necessarily lost. But the properties which distinguish the organic substances, and which make them important as ingredients of the body, do not depend immediately upon their ultimate chemical constitution, and are of a peculiar character ; being such as are only manifested in the interior of the living organism. Albumen, therefore, though it may contain a few equivalents more or less of oxygen or nitrogen, does not on that account cease to be albumen, so long as it retains its fluidity and its aptitude for undergoing the processes of absorption and transformation, which characterize it as an ingredient of the living body. It is for this reason that considerable discrepancy has existed at various times among chemists as to the real ultimate composition of these substances, different experimenters often obtaining different analytical results. This is not owing to any inaccuracy in the analyses, but to the fact that the organic substance itself really has a different ultimate constitution at different times. The most ap- proved formulae are those which have been established by Liebig for the following substances: — Fibrin = C29gH228N4o092S2 Albumen = C2,6H,g9N270g8S2 Casein ......= C288H22sN3gOggS2 Owing to the above mentioned variations, however, the same degree of importance does not attach to the quantitative ultimate analysis of an organic matter, as to that of other substances. This absence of a definite chemical constitution in the organic sub- stances is undoubtedly connected with their incapacity for crystalli- zation. It is also connected with another almost equally peculiar OEGANIC SUBSTANCES. 65 fact, viz., that although the organic substances unite with acids and with alkalies, they do not play the part of an acid toward the base, or of a base toward the acid ; for the acid or alkaline reaction of the substance employed is nqt neutralized, but remains as strong- after the combination as before. Furthermore, the union does not take place, so far as can be ascertained, in any definite proportions. The organic substances have, in fact, no combining equivalent; and their molecular reactions and the changes which they undergo in the body cannot therefore be expressed by the ordinary chemical phrases which are adapted to inorganic substances. Their true characters, as proximate principles, are accordingly to be sought for in other properties than those which depend upon their exact ulti- mate composition. One of these characters is that they are hygroscopic. As met with in different parts of the body, they present different degrees of con- sistency; some being nearly solid, others more or less fluid. But on being subjected to evaporation they all lose water, and are reduced to a perfectly solid form. If after this desiccation they are exposed to the contact of moisture, they again absorb water, swell, and regain their original mass and consistency. This phenomenon is quite different from that of capillary attraction, by which some in- organic substances become moistened when exposed to the contact of water; for in the latter case the water is simply entangled me- chanically in the meshes and pores of the inorganic body, while that which is absorbed by the organic matter is actually united with its substance, and diffused equally throughout its entire mass. Every organic matter is naturally united in this way with a certain quantity of water, some more and some less. Thus the albumen of the blood is in union with so much water that it has the fluid form, while the organic substance of cartilage contains less and is of a firmer con- sistency. The quantity of water contained in each organic sub- stance may be diminished by artificial desiccation, or by a deficient supply ; but neither of them can be made to take up more than a certain amount. Thus if the albumen of the blood and the organic substance of cartilage be both reduced by evaporation to a similar degree of dryness and then placed in water, the albumen will absorb so much as again to become fluid, but the cartilaginous substance only so much as to regain its usual nearly solid consistency. Even where the organic substance, therefore, as in the case of albumen, becomes fluid under these circumstances, it is not exactly a solution 5 66 PEOXIMATE PRINCIPLES OF THE THIRD CLASS. of it in water, but only a reabsorption by it of that quantity of fluid with which it is naturally associated. Another peculiar phenomenon characteristic of organic substances is their coagulation. Those which are naturally fluid suddenly as- sume, under certain conditions, a solid or semi-solid consistency. They are then said to be coagulated; and after coagulation they cannot be made to reassume their original condition. Thus fibrin coagulates on being withdrawn from the bloodvessels, albumen on being subjected to the temperature of boiling water, casein on being placed in contact with an acid. When an organic substance thus coagulates, the change which takes place is a peculiar one, and has no resemblance to the precipitation of a solid substance from a watery solution. On the contrary, the organic substance merely assumes a special condition; and in passing into the solid form it retains all the water with which it was previously united. Albumen, for example, after coagulation, retains the same quantity of water in union with it, which it held before. After coagulation, accordingly, this water may be driven off by evaporation, in the same manner as previously; and on being again exposed to moisture, the organic matter will again absorb the same quantity, though it will not re- sume the fluid form. By coagulation, an organic substance is permanently altered ; and though it may be afterwards dissolved by certain chemical re- agents, as, for example, the caustic alkalies, it is not thereby restored to its original condition, but only suffers a still further alteration. In many instances we are obliged to resort to coagulation in order to separate an organic substance from the other proximate principles with which it is associated. This is the case, for example, with the fibrin of the blood, which is obtained in the form of flocculi, by beating freshly drawn blood with a bundle of rods. But when separated in this way, it is already in an unnatural condition, and no longer represents exactly the original fluid fibrin, as it existed in the circulating blood. Nevertheless, this is the only mode in which it can be examined, as there are no means of bringing it back to its previous condition. Another important property of the organic substances is that they readily excite, in other proximate principles and in each other, those peculiar indirect chemical changes which are termed catalyses or catalytic transformations. That is to say, they produce the changes referred to, not directly, by combining with the substance which suffers alteration, or with any of its ingredients; but simply by their #* ORGANIC SUBSTANCES. 67 presence, which induces the chemical change in an indirect manner. Thus, the organic substances of the intestinal fluids induce a cata- lytic action by which starch is converted into sugar. The albumen of the blood, by contact with the organic substance of the muscular fibre, is transformed into a substance similar to it. The entire process of nutrition, so far as the organic matters are concerned, consists of such catalytic transformations. Many crystallizable substances, which when pure remain unaltered in the air, become changed if mingled with organic substances, even in small quantity. Thus the casein of milk, after being exposed for a short time to a warm atmosphere, becomes a catalytic body, and converts the sugar of the milk into lactic acid. In this change there is no loss nor addition of any chemical element, since lactic acid has precisely the same ultimate composition with sugar of milk. It is simply a transformation induced by the presence of the casein. Oily matters, which are entirely unalterable when pure, readily become rancid at warm temperatures, if mingled with an organic impurity. Fourthly, The organic substances, when beginning to undergo decay, induce in certain other substances the phenomenon of fer- mentation. Thus, the mucus of the urinary bladder, after a short exposure to the atmosphere, causes the urea of the urine to be con- verted into carbonate of ammonia, with the development of gaseous bubbles. The organic matters of grape juice, under similar circum- stances, give rise to fermentation of the sugar, by which it is con- verted into alcohol and carbonic acid. Fifthly, The organic substances are the only ones capable of undergoing the process of 'putrefaction. This process is a compli- cated one, and is characterized by a gradual liquefaction of the ani- mal substance, by many mutual decompositions of the saline matters which are associated with it, and by the development of peculiarly fetid and unwholesome gases, among which are carbonic acid, nitrogen, sulphuretted, phosphoretted, and carburetted hydrogen, and ammoniacal vapors. Putrefaction takes place constantly after death, if the organic tissue be exposed to a moist atmosphere at a moderately warm temperature. It is much hastened by the presence of other organic substances, in which decomposition has already commenced. The organic substances are readily distinguished, by the above general characters, from all other kinds of proximate principles. They are quite numerous; nearly every animal fluid and tissue con- taining at least one which is peculiar to itself. They have not as yet been all accurately described. The following list, however, 68 PEOXIMATE PRINCIPLES OF THE THIRD CLASS. comprises the most important of them, and those with which we are at present most thoroughly acquainted. The first seven are fluid, or nearly so, and either colorless or of a faint yellowish tinge. 1. Fibrin. — Fibrin is found in the blood ; where it exists, in the human subject, in the proportion of two to three parts per thousand. It is fluid, and mingled intimately with the other ingredients of the blood. It occurs also, but in much smaller quantity, in the lymph. It is distinguished by what is called its "spontaneous" coagulation; that is, it coagulates on being withdrawn from the vessels, or on the occurrence of any stoppage to the circulation. It is rather more abundant in the blood of some of the lower animals than in that of the human subject. In general, it is found in larger quantity in the blood of the herbivora than in that of the carnivora. 2. Albumen. — Albumen occurs in the blood, the lymph, the- fluid of the pericardium, and in that of the serous cavities gene- rally. It is also present in the fluid which may be extracted by pressure from the muscular tissue. In the blood it occurs in the proportion of about seventy-five parts per thousand. The white of egg, which usually goes by the same name, is not identical with the albumen of the blood, though it resembles it in some respects; it is properly a secretion from the mucous membrane of the fowl's ovi- duct, and should be considered as a distinct organic substance. Albumen coagulates on being raised to the temperature of 160° F.; and the coagulum, like that of all the other proximate principles, is soluble hi caustic potass. It coagulates also by contact with alco- hol, the mineral acids, ferrocyanide of potassium in an acidulated solution, tannin, and the metallic salts. The alcoholic coagulum, if separated from the alcohol by washing, does not redissolve in water. A very small quantity of albumen has been sometimes found in the saliva. 3. Casein. — This substance exists in milk, in the proportion of about forty parts per thousand. It coagulates by contact with all the acids, mineral and organic ; but is not affected by a boiling temperature. It is coagulated also by the juices of the stomach. It is important as an article of food, being the principal organic in- gredient in all the preparations of milk. In a coagulated form, it constitutes the different varieties of cheese, which are more or less highly flavored with various oily matters remaining entangled in the coagulated casein. GLOBULINE. — MUCOSINE. 69 What is called vegetable casein or "legumine," is different from the casein of milk, and constitutes the organic substance present in various kinds of peas and beans. 4. Globuline. — This is the organic substance forming the prin- cipal mass of the red globules of the blood. It is nearly fluid in its natural condition, and readily dissolves in water. It does not dissolve, however, in the serum of the blood; and the globules, therefore, retain their natural form and consistency, unless the serum be diluted with an excess of water, Globuline resembles albumen in coagulating at the temperature of boiling water. It is said to differ from it, however, in not being coagulated by contact with alcohol. 5. Pepsine. — This substance occurs as an ingredient in the gastric juice. It is not the same with the substance which Schwann extracted by maceration from the mucous membrane of the stomach, and which is regarded by Kobin, Bernard, &c., as only an artificial product of the alteration of the gastric tissues. There seems no good reason, furthermore, why we should not designate by this name the organic substance which really does exist in the gastric juice. It occurs in this fluid in very small quantity, not over fifteen parts per thousand. It is coagulable by heat, and also by contact with alcohol. But if the alcoholic coagulum be well washed, it is again soluble in a watery acidulated fluid. 6. Panceeatine. — This is the organic substance of the pancreatic juice, where it occurs in great abundance. It coagulates by heat, and by contact with sulphate of magnesia in excess. In its natural condition it is fluid, but has a considerable degree of viscidity. 7. MucosiNE is the organic substance which is found in the dif- ferent varieties of mucus, and which imparts to them their viscidity and other physical characters. Some of these mucous secretions are so mixed with other fluids, that their consistency is more or less diminished ; others which remain pure, like that secreted by the mucous follicles of the cervix uteri, have nearly a semi-solid con- sistency. But little is known with regard to their other specific characters. The next three organic substances are solid or semi-solid in con- sistency. 70 PKOXIMATE PEINCIPLES OF THE THIRD CLASS. 8. OSTEINE is the organic substance of the bones, in which it is associated with a large proportion of phosphate of lime. It exists, in those bones which have been examined, in the proportion of about two hundred parts per thousand. It is this substance which by long boiling of the bones is transformed into gelatine or glue. In its natural condition, however, it is insoluble in water, even at the boiling temperature, and becomes soluble only after it has been permanently altered by ebullition. 9. Cartilagine. — This forms the organic ingredient of cartilage. Like that of the bones, it is altered by long boiling, and is converted into a peculiar kind of gelatine termed " chondrine." Chondrine differs from the gelatine of bones principally in being precipitated by acids and certain metallic salts which have no effect on the latter. Cartilagine, in its natural condition, is very solid, and is closely united with the calcareous salts. 10. MuscuLiNE. — This substance forms the principal mass of the muscular fibre. It is semi-solid, and insoluble in water, but soluble in dilute muriatic acid, from which it may be again precipitated by neutralizing with an alkali. It closely resembles albumen in its chemical composition, and like it, contains, according to Scherer, two equivalents of sulphur. The four remaining organic substances form a somewhat peculiar group, Tliey are the coloring matters of the body. They exist always in small quantity, compared with the other ingredients, but communicate to the tissues and fluids a very distinct coloration. They all contain iron as one of their ultimate elements. 11. H^matine is the coloring matter of the red globules of the blood. It is nearly fluid like the globuline, and is united with it in a kind of mutual solution. It is much less abundant than the globuline, and exists in the proportion of about one part of hsema- tine to seventeen parts of globuline. The following is the formula for its composition which is adopted by Lehmann : — Hsematine ^ C44H22N30gFe. When the blood-globules from any cause become disintegrated, the hsematine is readily imbibed after death by the walls of the bloodvessels and the neighboring parts, staining them of a deep red color. This coloration has sometimes been mistaken for an evidence MELANIJSTE. — UEOSACINE. 71 of arteritis ; but is really a simple effect of post-mortem imbibition, as above stated. 12. Melanine. — This is the blackish-brown coloring matter which is found in the choroid coat of the eye, the iris, the hair, and more or less abundantly in the epidermis. So far as can be ascer- tained, the coloring matter is the same in all these situations. It is very abundant in the black and brown races, less so in the yellow and white, but is present to a certain extent in all. Even where the tinges produced are entirely different, as, for example, in brown and blue eyes, the coloring matter appears to be the same in cha- racter, and to vary only in its quantity and the mode of its arrange- ment ; for the tinge of an animal tissue does not depend on its local pigment only, but also on the muscular fibres, fibres of areolar tissue, capillary bloodvessels, &c. All these ingredients of the* tissue are partially transparent, and by their mutual interlacement and superposition modify more or less the effect of the pigment which is deposited below or among them. Melanine is insoluble in water and the dilute acids, but dissolves slowly in caustic potass. Its ultimate composition resembles that of hsematine, but the proportion of iron is smaller. ^ 13. BiLiVERDiNE is the coloring matter of the bile. It is yellow by transmitted light, greenish by reflected light. On exposure to the air in its natural fluid condition, it absorbs oxygen and assumes a bright grass green color. The same effect is produced by treating it with nitric acid or other oxidizing substances. It occurs in very small quantity in the bile, from which it may be extracted by pre- cipitating it with milk of lime (Robin), from which it is afterward separated by dissolving out the lime with muriatic acid. Obtained in this form, however, it is insoluble in water, having been coagu- lated by contact with the calcareous matter; and is not, therefore, precisely in its original condition. 14. Ubosacine is the yellowish-red coloring matter of the urine. It consists of the same ultimate elements as the other coloring mat- ters, but occurs in the urine in such minute quantity, that the relative proportion of its elements has never been determined. It readily adheres to insoluble matters when they are precipitated from the urine, and is consequently found almost always, to a greater or less extent, as an ingredient in urinary calculi formed of the urates 72 PEOXIMATE PEINCIPLES OF THE THIKD CLASS. or of uric acid. When tlie urates are thrown down also in the form of a powder, as a urinary deposit, they are usually colored more or less deeply, according to the quantity of urosacine which is preci- pitated with them. The organic substances which exist in the body require for their production an abundant supply of similar substances in the food. All highly nutritious articles of diet, therefore, contain more or less of these substances. Still, though nitrogenous matters must be abundantly supplied, under some form, from without, yet the par- ticular kinds of organic substances, characteristic of the tissues, are formed in the body by a transformation of those which are intro- duced with the food. The organic matters derived from vegetables, though similar in their general characters to those existing in the animal body, are yet specifically different. The gluten of wheat, th/e legumine of peas and beans, are not the same with animal al- bumen and fibrin. The only organic substances taken with animal food, as a general rule, are the albumen of eggs, the casein of milk, and the musculine of flesh ; and even these, in the food of the human species, are so altered and coagulated by the process of cooking, as to lose their specific characters before being introduced into the alimentary canal. They are still further changed by the process of digestion, and are absorbed under another form into the blood. But from their, subsequent metamorphoses there are formed, in the different parts of the body, osteine, cartilagine, haematine, globuline, and all the other varieties of organic matter that cha- racterize the different tissues. These varieties, therefore, originate as such in the animal economy by the catalytic changes which the ingredients of the blood undergo in nutrition. Only a very small quantity of organic matter is discharged with the excretions. The coloring matters of the bile and urine, and the mucus of the urinary bladder, are almost the only ones that find an exit from the body in this way. There is a minute quantity of organic matter exhaled in a volatile form with the breath, and a little also, in all probability, from the cutaneous sur- face. But the entire quantity so discharged bears but a very small proportion to that which is daily introduced with the food. The organic substances, therefore, are decomposed in the interior of the body. They are transformed by the process of destructive assimi- lation, and their elements are finally eliminated and discharged under other forms of combination. OF FOOD. 73 CHAPTER V. OF FOOD. Under the term " food" are included all those substances, solid and liquid, which are necessary to sustain the process of nutrition. The first act of this process is the absorption from without of all those materials which enter into the composition of the living frame, or of others which may be converted into them in the in- terior of the body. The proximate principles of the first class, or the "inorganic substances," require to be supplied in sufficient quantity to keep up the natural proportion in which they exist in the various solids and fluids. As we have found it to be characteristic of these substances, except in a few instances, that they suffer no alteration in the in- terior of the body, but,- on the contrary, are absorbed, deposited in its tissue, and pass out of it afterward unchanged, nearly every one of them requires to be present under its own proper form, and in sufficient quantity in the food. The alkaline carbonates, which a,re formed, as we have seen, by a decomposition of the malates, citrates and tartrates, constitute almost the only exception to this rule. Since water enters so largely into the composition of nearly every part of the body, it is equally important as an ingredient of the food. In the case of the human subject, it is probably the most important substance to be supplied with constancy and regularity, and the system suffers more rapidly when entirely deprived of fluids, than when the supply of solid food only is withdrawn. A man may pass eight or ten hours, for example, without solid food, and suffer little or no inconvenience ; but if deprived of water for the same length of time, he becomes rapidly exhausted, and feels the deficiency in a very marked degree. Magendie found, in his experiments on dogs subjected to inanition,^ that if the animals ' Comptes Rendus, vol. xiii. p. 256. 74 OF FOOD. were supplied with water alone they lived six, eight, and even ten days longer than if they were deprived at the same time of both solid and liquid food. Chloride of sodium, also, is usually added to the food in considerable quantity, and requires to be supplied with tolerable regularity ; but the remaining inorganic materials, such as calcareous salts, the alkaline phosphates, &c., occur natu- rally in sufficient quantity in most of the articles which are used as food. The proximate principles of the second class, so far as they con- stitute ingredients of the food, are naturally divided into two groups : 1st, the sugar, and 2d, the oily matters. Since starch is always converted into sugar in the process of digestion, it may be included, as an alimentary substance, in the same group with the sugars. There is a natural desire in the human species for both saccharine and oleaginous food. In the purely carnivorous animals, however, though no starch or sugar be taken, yet the body is main- tained in a healthy condition. It has been supposed, therefore, that saccharine matters could not be absolutely necessary as food ; the more so since it has been found, by the experiments of CI. Bernard, that, in carnivorous animals kept exclusively on a diet of flesh, sugar is still formed in the liver, as well as in the mammary gland. The above conclusion, however, which has been drawn from these facts, does not apply practically to the human species. The car- nivorous animals have no desire for vegetable food, while in the human species there is a natural craving for it, which is almost universal. It may be dispensed with for a few days, but not with impunity for any great length of time. The experiment has often enough been tried, in the treatment of diabetes, of confining the patient to a strictly animal diet. It has been invariably found that, if this regimen be continued for some weeks, the desire for vegetable food on the part of the patient becomes so imperative that the plan of treatment is U-navoidably abandoned. A similar question has also arisen with regard to the oleaginous matters. Are these substances indispensable as ingredients of the food, or may they be replaced by other proximate principles, such as starch or sugar? It has already been seen, from the experiments of Boussingault and others, that a certain amount of fat is produced in the body over and above that which is taken with the food ; and it appears also that a regimen abounding in saccharine substances is favorable to the production of fat. It is altogether probable, therefore, that the materials for the production of fat may be OF FOOD. 70 derived, under these cii'cumstances, either directly or indirectly from saccharine matters. But saccharine matters alone are not entirely sufficient. M. Huber' thought he had demonstrated that bees fed on pure sugar would produce enough wax to show that the sugar could supply all that was necessary to the formation of the fatty matter of the wax. Dumas and Milne-Edwards, however, in repeating Huber's experiments,^ found that this was not the case. Bees, fed on pure sugar, soon cease to work, and sometimes perish in considerable numbers ; but if fed with honey, which contains some v/axy and other matters beside the sugar, they thrive upon it; and produce, in a given time, a much larger quantity of fat than was contained in the whole supply of food. The same thing was established by Boussingault with regard to starchy matters. He found that in fattening pigs, though the quantity of fat accumulated by the animal considerably exceeded that contained in the food, yet fat must enter to some extent into the composition of the food in order to maintain the animals in a good condition ; for pigs, fed on boiled potatoes alone (an article abounding in starch but nearly destitute of oily matter), fattened slowly and with great difficulty ; while those fed on potatoes mixed with a greasy flaid fattened readily, and accumulated, as mentioned above, much more fat than was contained in the food. The apparent discrepancy between these facts may be easily ex- plained, when we recollect that, in order that the animal may become fattened, it is necessary that he be supplied not only with the materials of the fat itself, but also with everything else which is necessary to maintain the body in a healthy condition. Oleaginous matter is one of these necessary substances. The fats which are taken in with the food are not destined to be simply transported into the body and deposited there unchanged. On the contrary, they are altered and used up in the processes of digestion and nutrition ; while the fats which appear in the body as constituents of the tissues are, in great part, of new formation, and are produced from materials derived, perhaps, from a variety of different sources. It is certain, then, that either one or the other of these two groups of substances, saccharine or oleaginous, must enter into the composition of the food; and furthermore, that, though the oily matters may sometimes be produced in the body from the sugars, ' Natural History of Bees, Edinboro', 1821, p. 330. ^ Aimales de Chim. et de Phys., 3d series, vol. xiv. p. 400. 76 OF FOOD. it is also necessary for the perfect nutrition of the body that fat be supplied, under its own form, with the food. For the human species, also, it is natural to have them both associated in the alimentary materials. They occur together in most vegetable sub- stances, and there is a natural desire for them both, as elements of the food. They are not, however, when alone, or even associated with each other, sufficient for the nutrition of the animal body. Magendie found that dogs, fed exclusively on starch or sugar, perished after a short time with symptoms of profound disturbance of the nutritive functions. An exclusive diet of butter or lard had a similar effect. The animal became exceedingly debilitated, though without much emaciation ; and after death, all the internal organs and tissues were found infiltrated with oil. Boussingault' performed a similar experiment, with a like result, upon a duck, which was kept upon an exclusive regimen of butter. " The duck received 1350 to 1500 grains of butter every day. At the end of three weeks it died of inanition. The butter oozed from every part of its body. The feathers looked as though they had been steeped in melted butter, and the body exhaled an unwholesome odor like that of butyric acid." Lehmann was also led to the same result by some experiments which he performed upon himself for the purpose of ascertaining the effect produced on the urine by different kinds of food.^ This observer confined himself first to a purely animal diet for three weeks, and afterwards to a purely vegetable one for sixteen days, without suffering any marked inconvenience. He then put himself upon a regimen consisting entirely of non-nitrogenous sub- stances, starch, sugar, gum, and oil, but was only able to continue this diet for two, or at most for three days, owing to the marked disturbance of the general health which rapidly supervened. The unpleasant symptoms, however, immediately disappeared on his return to an ordinary mixed diet. The same fact has been esta- blished more recently by Dr. Wm. A. Hammond, Assistant Surgeon U. S. Army,^ in a series of experiments which he performed upon himself. He was enabled to live for ten days on a diet composed exclusively of boiled starch and water. After the third day, how- ■ Chimie Agricole, p. 166. ^ Journal fiir praktische Chemie, vol. xxvii. p. 257. * Experimental Researches, &c., being the Prize Essay of the American Medical Association for 1857. OF FOOD. 77 ever, the general health began to deteriorate, and became verj much disturbed before the termination of the experiment. The prominent symptoms were debility, headache, pyrosis, and palpitation of the heart. After the starchy diet was abandoned, it required some days to restore the health to its usual condition. The proximate principles of the third class, or the organic sub- stances proper, enter so largely into the constitution of the animal tissues and fluids, that their importance, as elements of the food, is easily understood. No food can be long nutritious, unless a certain proportion of these substances be present in it. Since they are so abundant as ingredients of the body, their loss or absence from the food is felt more speedily and promptly than that of any other sub- stance except water. They have, therefore, sometimes received the name of " nutritious substances," in cbntradistinction to those of the second class, which contain no nitrogen, and which have been found by the experiments of Magendie and others to be insufficient for the support of life. The organic substances, however, when taken alone, are no more capable of supporting life indefinitely than the others. It was found in the experiments of the French " Gela- tine Commission"^ that animals fed on pure fibrin and albumen, as well as those fed on gelatine, become after a short time much en- feebled, refuse the food which is offered to them, or take it with reluctance, and finally die of inanition. This result has been ex- plained by supposing that these substances, when taken alone, excite after a time such disgust in the animal that they are either no longer taken, or if taken are not digested. But this diso-ust itself is simply an indication that the substances used are insufficient and finally useless as articles of food, and that the system demands instinctively other materials for its nourishment. The instinctive desire of animals for certain substances is the surest indication that they are in reality required for the nutritive process ; and on the other hand, the indifference or repugnance manifested for injurious or useless substances, is an equal evidence of their unfitness as articles of food. This repugnance is well de- scribed by Magendie, in the report of the commission above alluded to, while detailing the result of his investigations on the nutritive qualities of gelatine. " The result," he says, " of these first trials was that pure gelatine was not to the taste of the dogs experimented on. Some of them suffered the pangs of hunger with the gelatine ' Comptes Eendus, 1841, vol. xiii. p. 267. 78 OF FOOD. within their reach, and would not touch it ; others tasted of it, but wjould not eat: others still devoured a certain quantity of it once or twice, and then obstinately refused to make any farther use of it." In one instance, however, Magendie succeeded in inducing a dog to take a considerable quantity of pure fibrin daily throughout the whole course of the experiment; but notwithstanding this, the animal became emaciated like the others, and died at last with the same symptoms of inanition. The alimentary substances of the second class, however, viz., the sugars and the oils, have been sometimes thought less important than the albuminous matters, because they do not enter so largely or so permanently into the composition of the solid tissues. The saccharine matters, when taken as food, cannot be traced farther than the blood. They undergo already, in the circulating fluid, some change by which their essential character is lost, and they cannot be any longer recognized. The appearance of sugar in the mammary gland and the milk is only exceptional, and does not occur at all in the male subject. The fats are, it is true, very gene- rally distributed throughout the body, but it is only in the brain and nervous matter that they exist intimately united with thie re- maining ingredients of the tissues. Elsewhere, as already mentioned, it is deposited in distinct drops and granules, and so long as it re- mains in this condition must of course remain inactive, so far as reo-ards any chemical nutritive process. In this condition it seems to be held in reserve, ready to be absorbed by the blood, whenever it may be required for the purposes of nutrition. On being reab- sorbed, however, as soon as it again enters the blood or unites intimately with the substance of the tissues, it at once changes its condition and loses its former chemical constitution and properties. It is for these reasons that the albuminoid matters have been sometimes considered as the only " nutritious" substances, because they alone constitute under their own form a great part of the ingredients of the tissues, while the sugars and the oils rapidly dis- appear by decomposition. It has even been assumed that the pro- cess by which the sugar and the oils disappear is one of direct combustion or oxidation, and that they are destined solely to be consumed in this way, not to enter at all into the composition of tlie tissues, but only to maintain the heat of the body by an inces- sant process of combustion in the blood. They have been therefore termed the " combustible" or " heat-producing" elements, while the OF FOOD. 79 albuminoid substances were known as the nutritious or "plastic" elements. This distinction, however, has no real foundation. In the first place, it is not at all certain that the sugars and the oils which dis- appear in the body are destroyed by combustion. This is merely an inference which has been made without any direct proof. All we know positivel}^ in regard to the matter is that these substances soon become so altered in the blood that they can no longer be recognized by their ordinary chemical properties ; but we are still ignorant of the exact nature of the transformations which they undergo. Furthermore, the difference between the sugars and the oils on the one hand, and the albuminoid substances on the other, so far as regards their decomposition and disappearance in the body, is only a difference of time. The albuminoid substances become transformed more slowly, the sugars and the oils more rapidly. Even if it should be ascertained hereafter that the sugars and the oils really do not unite at all with the solid tissues, but are entirely decomposed in the blood, this would not make them any less im- portant as alimentary substances, since the blood is as essential a part of .the body as the solid tissues, and its nutrition must be pro- vided for equally with theirs. It is evident, therefore, that no single proximate principle, nor even any one class of them alone, can be sufficient for the nutrition of the body; but that the food, to be nourishing, must contain substances belonging to all the different groups of proximate prin- ciples. The albuminoid substances are first in importance because they constitute the largest part of the entire mass of the body ; and exhaustion therefore follows more rapidly when they are withheld than when the animal is deprived of other kinds of alimentary matter. But starchy and oleaginous substances are also requisite ; and the body feels the want of them sooner or later, though it may be plentifully supplied with albumen and fibrin. Finally, the in- organic saline matters, though in smaller quantity, are also neces- sary to the continuous maintenance of life. In order that the animal tissues and fluids remain in a healthy condition and take their proper part in the functions of life, they must be supplied with all the ingredients necessary to their constitution ; and a man may be starved to death at last by depriving him of chloride of sodium or phosphate of lime just as surely, though not so rapidly, as if he were deprived of albumen or oil. In the different kinds of food, accordingly, which have been 80 OF FOOL*. adopted by the universal and instinctive cTioice of man, the three different classes of proximate principles are all more or less abund- antly represented. In all of them there exists naturally a certain proportion of saline substances ; and water and chloride of sodium are gefnerally taken with them in addition. In milk, the first food supplied to the infant, we have casein which is an albuminoid substance, butter which represents the oily matters, and sugar of milk belonging to the saccharine group, together with water and saline matters, in the following proportions : — ^ Composition of Cow's Milk. Water 87.02 Casein ' . . . 4.48 Butter . . . . , 3.13 Sugar of milk 4.77 Soda 1 Chlorides of potassium and sodium ..... Phosphates of soda and potass Phosphate of lime • • .1-0.60 " magnesia . . Alkaline carbonates . . . . Iron, &c. . . . ■ . ■ • • • • • 100.00 In wheat flour, gluten is the albuminoid matter, sugar and starch the non-nitrogenous principles. Composition of Wheat Flouk. Gluten . . 10.2 Gum . 2.8 Starch . . 72.8 Water . 4.2 . 10.0 Sugar . 100.0 The other cereal grains mostly contain oil in addition to the above. Composition op Dkied Oatmeal. Starch 59.00 Bitter matter and sugar 8.25 Gray albuminous matter 4.30 Fatty oil 2.00 Gum • 2.50 Husk, mixture, and loss 23.95 100.00 Eggs contain albumen and salts in the white, with the addition of oily matter in the yolk. * The accompanying analyses of various kinds of food are taken from Pereira on Food and Diet, New York, 1843. OF FOOD. 81 Composition of Eggs. White of Egg. Yolk of Egg. Water .... 80.00 53.78 Albumen and mucus . 15.28 12.75 Yellow oil ... 28.75 Salts .... 4.72 4.72 100.00 100.00 In ordinary flesh or butcher's meat, we have the albuminoid matter of the muscular fibre and the fat of the adipose tissue. Composition of Ordinary Butcher's Meat. Meat devoid of fat . 85.7 | ^^^^^ .... 63.418 I Solid matter . . . 22.282 Fat, cellular tissue, &c 14.300 100.000 From what has been said above, it will easily be seen that the nutritious character of any substance, or its value as an article of food, does not depend simply upon its containing either one of the alimentary substances mentioned above in large quantity ; but upon its containing them mingled together in such proportion as is requisite for the healthy nutrition of the body. What these pro- portions are cannot be determined from simple chemical analysis, nor from any other data than those derived from direct observation and experiment. The total quantity of food required by man has , been variously estimated. It will necessarily vary, indeed, not only with the con- stitution and habits of the individual, but also with the quality of the food employed ; since some articles, such as corn and meat, con- tain very much more alimentary material in the same bulk than fresh fruits or vegetables. Any estimate, therefore, of the total quantity should state also the kind of food used ; otherwise, it will be altogether without value. From experiments performed while living on an exclusive diet of bread, fresh meat, and butter, witb coffee and water for drink, we have found that the entire quantity of food required during twenty-four hours by a man in full health, and taking free exercise in the open air, is as follows : — Meat 16 ounces. Bread . . . . , 19 " Butter 3^ " Fluids 52 " That is to say, rather less than two and a half pounds of solid food, and rather over three pints of liquid food. 6 82 OF FOOD. Another necessary consideration, in estimating the value of any substance as an article of food, is its digestibility. A vegetable or animal tissue may contain an abundance of albuminoid or starchy matter, but may be at the same time of such an unyielding consist- ency as to be insoluble in the digestive fluids, and therefore useless as an article of food. Bones and cartilages, and the fibres of yel- low elastic tissue, are indigestible, and therefore not nutritious. The same remark may be made with regard to the substances con- tained in woody fibre, and the hard coverings and kernels of various fruits. Everything, accordingly, which softens or disintegrates a hard alimentary substance renders it more digestible, and so far increases its value as an article of food. The preparation of food by cooking has a twofold object : first, to soften or disintegrate it, and second, to give it an attractive flavor. Many vegetable substances are so hard as to be entirely indigestible in a raw state. Eipe peas and beans, the difierent kinds of grain, and many roots and fruits, require to be softened by boil- ing, or some other culinary process, before they are ready for use. With them, the principal change produced by cooking is an altera- tion in consistency. With most kinds of animal food, however, the effect is somewhat different. In the case of muscular flesh, for example, the muscular fibres themselves are almost always more or less hardened by boiling or roasting ; but, at the same time, the fibrous tissue by which they are held together is gelatinized and softened, so that the muscular fibres are more easily separated from each other, and more readily attacked by the digestive fluids. But beside this, the organic substances contained in meat, which are all of them very insipid in the raw state, acquire, by the action of heat in cooking, a peculiar and agreeable flavor. This flavor excites the appetite and stimulates the flow of the digestive fluids, and renders, in this way, the entire process of digestion more easy and expeditious. The changes which the food undergoes in the interior of the body may be included under three different heads : first, digestion^ or the preparation of the food in the alimentary canal; second, assimila- tion^ by which the elements of the food are converted into the ani- mal tissues ; and third, excretion, by which it is again decomposed, and finally discharged from the body. DIGESTION. 83 CHAPTER VI. DIGESTION. Digestion is that process by whicli the food is reduced to a form iu which it can be absorbed from the intestinal canal, and taken up by the bloodvessels. This process does not occur in vegetables. For vegetables are dependent for their nutrition, mostly, if not entirely, upon a supply of inorganic substances, as water, saline matters, carbonic acid, and ammonia. These materials constitute the food upon which plants subsist, and are converted in their inte- rior into other substances, by the nutritive process. These mate- rials, furthermore, are constantly supplied to the vegetable under such a form as to be readily absorbed. Carbonic acid and ammonia exist in a gaseous form in the atmosphere, and are also to be found in solution, together with the requisiie saline matters, in the water with which the soil is penetrated. All these substances, therefore, are at once ready for absorption, and do not require any preliminary modification. But with animals and man the case is different. They cannot subsist upon these inorganic substances alone, but require for their support materials which have already been organ- ized, and which have previously constituted a part of animal or vegetable bodies. Their food is almost nvariably solid or semi-solid at the time when it is taken, and insoluble in water. Meat, bread, fruits, vegetables, &g., are all taken into the stomach in a solid and insoluble condition ; and even those substances which are naturally fluid, such as milk, albumen, white of egg, are almost always, in the human species, coagulated and solidified by the process of cooking, before being taken into the stomach. In animals, accordingly, the food requires to undergo a process of digestion, or liquefaction, before it can be absorbed. In all cases, the general characters of this process are the same. It consists essentially in the food being received into a canal, running through the body from mouth to anus, called the " alimentary canal," in which it comes in contact with certain digestive fluids, which act 84. , DIGESTION. upon it in such a way as to liquefy and dissolve it. These fluids are secreted by the mucous membrane of the alimentary canal, and by certain glandular organs situated in its neighborhood. Since the food always consists, as we have already seen, of a mixture of vari- ous substances, having different physical and chemical properties, the several digestive fluids are also different from each other ; each one of them exerting a peculiar action, which is more or less con- fined to particular species of food. As the food passes through the intestine from above downward, those parts of it which become liquefied are successively removed by absorption, and taken up by the vessels ; while the remaining portions, consisting of the indiges- tible matter, together with the refuse of the intestinal secretions, gradually acquire a firmer consistency owing to the absorption of the fluids, and are finally discharged from the intestine under the form of feces. In different species of animals, however, the difference in their habits, in the constitution of their tissues, and in the character of their food, is accompanied with a corresponding variation in the anatomy of the digestive apparatus, and the character of the secreted fluids. As a general rule, the digestive apparatus of herbivorous animals is more complex than that of the carnivora ; since, in vege- table substances, the nutritious matters are often present in a very solid and unmanageable form, as, for example, in raw starch and the cereal grains, and are nearly always entangled among vegetable cells and fibres of an indigestible character. In those instances, where the food consists mostly of herbage, as grass, leaves, &c., the digestible matters bear only a small proportion to the entire quan- tity; and a large mass of food must therefore be taken, in order that the requisite amount of nutritious material may be extracted from it. In such cases, accordingly, the alimentary canal is large and long; and is divided into many compartments, in which different processes of disintegration, transformation, and solution are carried on. In the common fowl, for instance (Fig. 16), the food, which con- sists mostly of grains, and frequently of insects with hard, coria- ceous integument, first passes down the oesophagus (a) into a diverticulum or pouch (6) termed the crop. Here it remains for a time, mingled with a watery secretion in which the grains are macerated and softened. The food is then carried farther down until it reaches a second dilatation (c), the proventriculus, or secreting stomach. The mucous membrane here is thick and DIGESTION. 85 glandular, and is provided witli numerous se- Fig' 16. creting follicles or crypts. From them an acid fluid is poured out, by which the food is subjected to further changes. It next passes into the gizzard {d\ or triturating stomach, a cavity inclosed by thick, muscular walls, and lined with a remarkably tough and horny epithelium. Here it is subjected to the crush- ing and grinding action of the muscular parietes, assisted by grains of sand and gravel, which the animal instinctively swallows with the food, by which it is so triturated and dis- integrated, that it is reduced to a uniform pulp, upon which the digestive fluids can effectually operate. The mass then passes into the intes- tine (e), where it meets with the intestinal juices, which complete the process of solution; and from the intestinal cavity it is finally ab- sorbed in a liquid form, by the vessels of the mucous membrane. In the ox, again, the sheep, the camel, the deer, and all ruminating animals, there are four distinct stomachs through which the food passes in succession; each lined with mucous membrane of a different structure, and adapted to perform a different part in the digestive process (Fig. 17). When first swallowed, the food is received into the ru- men, or paunch (b), a large sac, itself par- tially divided by incomplete partitions, and lined by a mucous membrane thickly set with long prominences or villi. Here it ac- cumulates while the animal is feeding, and is retained and macerated in its own fluids. When the anim.al has finished browsing, and the process of rumination commences, the food is regurgitated into the mouth by an inverted action of -tfee muscular walls of the paunch and oesophagus, and slowly masticated. It then descends again along the oesophagus ; but instead of enter- ing the first stomach, as before, it is turned off by a muscular valve into the second stomach, or reticulum (c), which is distinguished by the intersecting folds of its mucous membrane, which give it Alimentary Canal of Fowl. — a. Oesophagus. 6. Crop, c. Proventriculus, or secret- ing stomach, d. Gizzard, or triturating stomach, e. In- testine. /. Two long cjecal tubes which open into the in- testine a short distance above its termination. 86 DIGESTION. CoMPOu>T) Stomach of Ox. — a. CEsopliagus. h. Rumen, or first stomach, e. Eeticulum, or second, d. Omasus, or third, e. Obomasus, or fourth. /. Duodenum. a honey-combed or reticulated appearance. Here the food, already tritura'ed in the mouth, and ^^' mixed with the saliva, is further macerated in the fluids swallowed by the animal, which always ac- cumulate in considerable quan- tity in the reticulum. The next cavity is the omasus^ or "psalte- rium" (cf), in which the mucous membrane is arranged in longi- tudinal folds, alternately broad and narrow, lying parallel with each other, like the leaves of a book, so that the extent of mucous surface, brought in contact with the food, is very much increased. The exit from this cavity leads directly into the obomasus^ or " rennet" (e), which is the true digestive stomach, in which the mucous membrane is softer, thicker, and more glandular than elsewhere, and in which an acid and highly solvent fluid is secreted. Then follows the intestinal canal with its various divisions and variations. In the carnivora, on the other hand, the alimentary canal is shorter and narrower than in the preceding, and presents fewer complexities. The food, upon which these animals subsist, is softer than that of the herbivora, and less encumbered with indigestible matter ; so that the process of its solution requires a less extensive apparatus. In the human species, the food is naturally of a mixed cha- racter, containing both animal and vegetable substances. But the digestive apparatus in man resembles almost exactly that of the carnivora. For the vegetable matters which we take as food are, in the first place, artificially separated, to a great extent, from indi- gestible impurities ; and secondly, they are so softened by the process of cooking as to become nearly or quite as easily digestible as animal substances. In the human species, however, the process of digestion, though simpler than in the herbivora, is still complicated. The alimentary canal is here, also, divided into different compartments or cavities, which communicate with each other by narrow orifices. At its DIGESTION. commencement (Fig. 18), we find the cavity of the mouth, which is guarded at its posterior extremity by the muscular valve of the isthmus of the fauces. Through the pharynx and oesophagus (a), it com- municates with the second compartment, or the sto- mach (5), a flask-shaped dilatation, which is guarded at the cardiac and pyloric orifices by circular bands of muscular fibres. Then comes the small intestine (e), different parts of which, owing to the varying struc- ture of their mucous mem- branes, have received the different names of duode- num, jejunum, and ileum. In the duodenum, we have the orifices of the biliary and pancreatic ducts (/, g). Finally, we have the large intestine (/z, i, j\ k), separated from the smaller by the ileo-C8ecal valve, and ter- minating, at its lower ex- tremity, by the anus, at which is situated a double sphincter, for the purpose of guarding its orifice. Everywhere the alimentary canal is composed of a mucous membrane and a muscular coat, with a layer of submucous areolar tissue between the two. The mus- cular coat is everywhere composed of a double layer of longitudinal and transverse fibres, by the alternate contraction and relaxation of which the food is carried through the canal from above downward. The mucous Human Alimentary Canal. — a. (Esophagus. 6. Sto- mach, c. Cardiac orifice, d. Pylorus, e. Small intestine. /. Biliary duct. g. Pancreatic duct. h. Ascending colon. i. Transverse colon. J. Descending colon, k. Rectum. 88 DIGESTION". membrane presents, also, a different structure, and has different properties in different parts. In the mouth and oesophagus, it is smooth, with a hard, whitish, and tessellated epithelium. This kind of epithelium terminates abruptly at the cardiac orifice of the stomach. The mucous membrane of the gastric cavity is soft and glandular, covered with a transparent, columnar epithelium, and thrown into minute folds or projections on its free surface, which are sometimes reticulated with each other. In the small intestine, we find large transverse folds of mucous membrane, the valvulce conniventes, the minute villosities which cover its surface, and the peculiar glandular structures which it contains. Finally, in the large intestine, the mucous membrane is again different. It is here smooth and shining, free from villosities, and provided with a different glandular apparatus. Furthermore, the digestive secretions, also, vary in these different regions. In its passage from above downwards, the food meets with no less than five different digestive fluids. First it meets with the saliva in the cavity of the mouth ; second, with the gastric juice^ in the stomach; third, with the hile ; fourth, with the pancreatic fluid; and fifth, with the intestinal juice. It is the most important characteristic of the process of digestion, as established by modern researches, that different elements of the food are digested in different parts of the alimentary canal by the agency of different digestive fluids. By their action, the various ingredients of the alimentary mass are successively reduced to a fluid condition, and are taken up by the vessels of the intestinal mucous membrane. The action which is exerted upon the food by the digestive fluids is not that of a simple chemical solution. It is a transforma- tion, by which the ingredients of the food are altered in character at the same time that they undergo the process of liquefaction. The active agent in producing this change is in every instance an organic matter, which enters as an ingredient into the digestive fluid; and which, by coming in contact with the food, exerts upon it a catalytic action, and transforms its ingredients into other sub- stances. It is these newly formed substances which are finally absorbed by the vessels, and mingled with the general current of the circulation. In our study of the process of digestion, the different digestive fluids will be examined separately, and their action on the aliment- ary substances in the different regions of the digestive apparatus successively investigated. MASTICATION. 89 MASTICATION. In the first division of the alimentary canal, viz., the mouth, the food undergoes simultaneously two different operations, viz., mas- tication and insalivation. Mastication consists in the cutting and trituration of the food by the teeth, by the action of which it is reduced to a state of minute subdivision. This process is entirely a mechanical one. It is necessary, in order to prepare the food for the subsequent action of the digestive fluids. As this action is chemical in its nature, it will be exerted more promptly and effi- ciently if the food be finely divided than if it be brought in con- tact with the digestive fluids in a solid mass. This is always the case when a solid body is subjected to the chemical action of a solvent fluid ; since, by being broken up into minute particles, it offers a larger surface to the contact of the fluid, and is more readily attacked and dissolved or decomposed by it. In the structure of the teeth, and their physiological action, there are certain marked differences, corresponding with the habits of the animal, and the kind of food upon which it subsists. In fish and serpents, in which the food is swallowed entire, and in which the process of digestion, accordingly, is comparatively slow, the teeth are simply organs of prehension. They have generally the form of sharp, curved spines, with their points set backward (Fig. 19), and arranged in a double or triple row about the edges of the jaws, and sometimes ^^?!^ covering the mucous surfaces of the mouth, tongue, and palate. They serve merely to retain the prey, and prevent its escape, after it has been seized by the animal. In .1 • n 1 J.1 J? Skull op Rattlesnake. (After the carnivorous quadrupeds, as those of Achiiie-Eichard.) the dog and cat kind, and other similar families, there are three different kinds of teeth adapted to different mechanical purposes. (Fig. 20.) First, the incisors, twelve in num- ber, situated at the anterior part of the jaw, six in the superior, and six in the inferior maxilla, of flattened form, and placed with their thin edges running from side to side. The incisors, as their name indicates, are adapted for dividing the food by a cutting motion, like that of a pair of shears. Behind them come the canine teeth, or tusks, one on each side of the upper and under jaw. These are long, curved, conical, and pointed ; and are used as 90 DIGESTION. weapons of offence, and for laying hold of and retaining tlie prey. Lastly, the molars, eight or more in number on each side, are larger and broader than the incis- ors, and provided with serrated edges, each presenting several sharp points, arranged generally in a di- rection parallel with the line of the jaw. In these animals, mastication is very imperfect, since the food is not ground up, but only pierced and mangled by the action of the teeth before being swallowed into the stomach. In the herbivora, on the other hand, the incisors are pre- sent only in the lower jaw in the ruminating animals, though in the horse they are found in both the upper and lower maxilla (Fig. Skull of Polar Bear. Anterior view showing incisors and canines. Fig. 21. Skull of the Horse. Fig. 22. 21). They are used merely for cutting off the bundles of grass or herbage, on which the animal feeds. The canines are either absent or only slightly developed, and the real process of mastication is performed altogether by the molars. These are large and thick (Fig. 22), and present a broad, flat surface, diversified by variously folded and projecting ridges of enamel, with shallow grooves, intervening between them. By the lateral rubbing motion of the roughened surfaces against each other, the food is effectually comminuted and reduced to a pulpy mass. In the human subject, the teeth combine the Molar Tooth of the Horse. Grinding sur- face. MASTICATION. 91 In- human Teeth — Upper Jaw. — a. Incisors. 6. Canines. Anterior molars, d. Posterior molars. characters of those of the carnivora and the herbivora. (Fig. 23.) The incisors (a), four in number in each jaw, have, ^ig- 23. as in other instances, a cuttingedge running from side to side. The canines (&), which are situated immediately behind the former, are much less prominent and pointed than in the carnivora, and differ less in form from the incisors on the one hand, and the first molars on the other. The molars, again (c, d\ are thick and strong, and have compa- ratively flat surfaces, like those of the herbivora ; but instead of presenting curvilinear ridges, are covered with more or less conical eminences, like those of the carnivora. In the human subject, therefore, the teeth are evidently adapted for a mixed diet, consist- ing of both animal and vegetable food. Mastication is here as perfect as it is in the herbivora, though less prolonged and labori- ous ; for the vegetable substances used by man, as already remarked, are previously separated to a great extent from their impurities, and softened by cooking; so that they do not require, for their mas- tication, so extensive and powerful a triturating apparatus. Finally, animal substances are more completely masticated in the human subject than they are in the carnivora, and their digestion is accord- ingly completed with greater rapidity. We can easily estimate, from the facts above stated, the great importance, to the digestive process, of a thorough preliminary mastication. If the food be hastily swallowed in undivided masses, it must remain a long time undissolved in the stomach, where it will become a source of irritation and disturbance ; but if reduced beforehand, by mastication, to a state of minute subdivision, it is readily attacked by the digestive fluids, and becomes speedily and completely liquefied. 92 DIGESTION". SALIVA. At the same time that the food is masticated, it is mixed in the cavity of the mouth with the first of the digestive fluids, viz., the saliva. Human saliva, as it is obtained directly from the buccal cavity, is a colorless, slightly viscid and alkaline fluid, with a spe- cific gravity of 1005. When first discharged, it is frothy and opaline, holding in suspension minute, whitish flocculi. On being allowed to stand for some hours in a cylindrical glass vessel, an opaque, whitish deposit collects at the bottom, while the supernatant fluid becomes clear. The deposit, when examined by the micro- scope (Fig. 24), is seen to consist of abundant epithe- lium scales from the internal surface of the mouth, de- tached by mechanical irrita- tion, minute, roundish, gra- nular, nucleated cells, appa- rently epithelium from the mucous follicles, a certain amount of granular matter, and a few oil-globules. The supernatant fluid has a faint bluish tinge, becomes slightly opalescent by boiling, and the addition of nitric acid. Alcohol in excess, causes the precipitation of abundant whitish flocculi. According to Bidder and Schmidt,^ the composi- tion of saliva is as follows : — Buccal asd Glandulab Epithelium, with Granular Matter and Oil-globules ; deposited as sediment from human saliva. CoMPOsiTiox OF Saliva. Water Organic matter ...... Sulpho-cyanide of potassium . . . . Phosphates of soda, lime, and magnesia . Chlorides of sodium and potassium . Mixture of epithelium . . . • . . 995.16 1.34 0.06 .98 .84 1.62 1000.00 The organic substance present in the saliva has been occasionally ' Verdaunngsssefte und Stoffwechsel. Leipzig, 1852. SALIVA. 93 known by the name of piyaline. It is coagulable by alcohol, but not by a boiling temperature. A very little albumen is also pre- sent, mingled with the ptyaline, and produces the opalescence which appears in the saliva when raised to a boiling temperature. The sulpho-cyanogen may be detected by a solution of chloride of iron, which produces the characteristic red color of sulpho-cyanide of iron. The alkaline reaction of the saliva varies in intensity during the day, but is nearly always sufficiently distinct. The saliva is not a simple secretion, but a mixture of four dis- tinct fluids, which differ from each other in the source from which they are derived, and in their physical and chemical properties. These secretions are, in the human subject, first, that of the parotid gland ; second, that of the submaxillary ; third, that of the sub- lingual; and fourth, that of the mucous follicles of the mouth. These different fluids have been comparatively studied, in the lower animals, by Bernard, Frerichs, and Bidder and Schmidt. The parotid saliva is obtained in a state of purity from the dog by exposing the duct of Steno where it crosses the masseter muscle, and introducing into it, through an artificial opening, a fine silver canula. The parotid saliva then runs directly from its external orifice, without being mixed with that of the other salivary glands. It is clear, limpid, and watery, without the slightest viscidity, and has a faintly alkaline reaction. The submaxillary saliva is ob- tained in a similar manner, by inserting a canula into Wharton's duct. It differs from the parotid secretion, so far as its physical properties are concerned, chiefly in possessing a well-marked vis- cidity. It is alkaline in reaction. The sublingual saliva is also alkaline, colorless, and transparent, and possesses a greater degree of viscidity than that from the submaxillary. The mucous secre- tion of the follicles of the mouth, which forms properly a part of the saliva, is obtained by placing a ligature simultaneously on Wharton's and Steno's ducts, and on that of the sublingual gland, so as to shut out from the mouth all the glandular salivary secre- tions, and then collecting the fluid secreted by the buccal mucous membrane. This fluid is very scanty, and much more viscid than either of the other secretions ; so much so, that it cannot be poured out in drops when received in a glass vessel, but adheres strongly to the surface of the glass. According to Bernard,^ the principal distinction between these ' Le(;ons de Physiologie Experimentale, Paris, 1856, p. 93. ^4 DIGESTION. different salivary fluids resides in the character of the organic matter peculiar to each one. The organic ingredient of the parotid saliva is small in quantity, perfectly fluid, and analogous in some respects to albumen, since it coagulates by a boiling temperature. That of the submaxillary is moderately viscid, and has a tendency to solidify or gelatinize on cooling ; while that of the sublingual and mucous secretions is excessively viscid, but does not gelatinize at a low temperature. Tlie saliva proper consists, therefore, of a nearly homogeneous mixture of all these different secretions ; of which that from the parotid is the most abundant, tbat of the sublingual and of the mucous follicles of the mouth the least so. Bidder and Schmidt obtained, from one of the parotid glands of the dog, one hundred and thirty-six grains of fluid in an hour; from the submaxillary, eighty-seven grains ; and from the mucous follicles of the mouth, after ligature of both Wharton's and Steno's ducts, thirty-one grains. The saliva, as a whole, is not secreted with uniform rapidity at all times. While fasting, and while the tongue and jaws are at rest, it is supplied in but small quantity, just sufficient to keep the mucous membrane of the mouth moist and pliable. - Any movement of the jaws, however, increases the rapidity of its flow. It is still more powerfully stimulated by the introduction of food, particularly that which has a decided taste, or which requires an active movement of the jaws for its mastication. The saliva is then poured out in abundance, and continues to be rapidly secreted until the food is masticated and swallowed. A very curious fact has been observed by M. Colin, Professor of Anatomy and Physiology at the Veterinary School of Alfort,' viz., that in the horse and ass, as well as in the cow and other ruminat- ing animals, the parotid glands of the two opposite sides, during mastication, are never in active secretion at the same time; but that they alternate with each other, one remaining quiescent while the other is active, and vice versa. In these animals, mastication is said to be unilateral, that is, when the animal commences feeding or ruminating the food is triturated, for fifteen minutes or more, by the molars of one side only. It is then changed to the opposite side; and for the next fifteen minutes mastication is performed by the molars of that side only. It is then changed back again, and so on alternately, so that the direction of the lateral movements of ' Traite de Physiologie Comparee, Paris, 1854, p. 468. SALIVA. 95 the jaw may be reversed many times during the course of a meal. By establishing a salivary fistula simultaneously on each side, it is found that the flow of saliva corresponds with the direction of the masticatory movement. When the animal masticates on the right side, it is the right parotid which secretes actively, while but little saliva is supplied by the left ; when mastication is on the left side, the left parotid pours out an abundance of fluid, while the right is nearly inactive. It is probable, however, that this alternation of function does not exist, to the same extent at least, in man and the carnivora, in whom mastication is performed very nearly on both sides at once. Owing to the variations in the rapidity of its secretion, and also to the fact that it is not so readily excited by artificial means as by the presence of food, it becomes somewhat difiicult to estimate the total quantity of saliva secreted daily. The first attempt to do so was made by Mitscherlich,' who collected from two to three ounces in twenty-four hours from an accidental salivary fistula of Steno's duct in the human subject ; from which it was supposed that the total amount secreted by all the glands was from ten to twelve ounces daily. As this man was a hospital patient, however, and suffering from constitutional debility, the above calculation cannot be re-, garded as an accurate one, and accordingly Bidder and Schmidt^ make a higher estimate. One of these observers, in experimenting upon himself, collected from the mouth in one hour, without using any artificial stimulus to the secretion, 1500 grains of saliva ; and calculates, therefore, the amount secreted daily, making an allow- ance of seven hours for sleep, as not far from 25,000 grains, or about three and a half pounds avoirdupois. On repeating this experiment, however, we have not been able to collect from the mouth, without artificial stimulus, more than 556 grains of saliva per hour. This quantity, however, may be greatly increased by the introduction into the mouth of any smooth unirri- tating substance, as glass beads or the like ; and during the masti- cation of food, the saliva is poured out in very much greater abund- ance. The very sight and odor of nutritious food, when the appetite is excited, will stimulate to a remarkable degree the flow of saliva; and, as it is often expressed, "bring the water into the mouth." Any estimate, therefore, of the total quantity of saliva, based on the amount secreted in the intervals of mastication, would be a very « Simon's Chemistry of Man. Phila. ed., 1846, p. 295. ^ Op. cit., p. 14. 96 DIGESTION. imperfect one. We may make a tolerably accurate calculation, however, by ascertaining how much, is really secreted during a meal, over and above that which is produced at other times. We have found, for example, by experiments performed for this pur- pose, that wheaten bread gains during complete mastication 55 per cent, of its weight of saliva ; and that fresh cooked meat gains, under the same circumstances, 48 per cent, of its weight. We have already seen that the daily allowance of these two substances, for a man in full health, is 19 ounces of bread, and 16 ounces of meat. The quantity of saliva, then, required for the mastication of these two substances, is, for the bread 4,572 grains, and for the meat 3,360 grains. If we now calculate the quantity secreted between meals as continuing for 22 hours at 556 grains per hour, we have: — Saliva required for mastication of bread = 4572 grains. " " " " " meat = 3360 " " secreted in intervals of meals = 12232 " Total quantity in twenty-four hours = 20164 grains ; or rather less than 3 pounds avoirdupois. The most important question, connected with this subject, relates to ihQ function of the saliva in the digestive process. A very remarka- ble property of this fluid is that which was discovered by Leuchs in Germany, viz., that it possesses the power of converting boiled starch into sugar, if mixed with it in equal proportions, and kept for a short time at the temperature of 100° F. This phenomenon is one of catalysis, in which the starch is transformed into sugar by simple contact with the organic substance contained in the saliva. This organic substance, according to the experiments of Mialhe,^ may even be precipitated by alcohol, and kept in a dry state for an indefinite length of time without losing the power of converting starch into sugar, when again brought in contact with it in a state of solution. This action of ordinary human saliva on boiled starch takes place sometimes with great rapidity. Traces of glucose may occasionally be detected in the mixture in one minute after the two substances have been brought in contact; and we have even found that starch paste, introduced into the cavity of the mouth, if already at the temperature of 100° F., will yield traces of sugar at the end of half a minute. The rapidity, however, with which this action is mani- ' Chimie appliquee a la Physiologie et a la Therapeutique, Paris, 1856, p. 43. SALIVA. 97 fested, varies very mucli, as was formerly noticed by Lebraann, at different times ; owing, in all probability, to the varying constitution of the saliva itself. It is often impossible, for example, to find any evidences of sugar, in the mixture of starch and saliva, under five, ten, or fifteen minutes ; and it is frequently a longer time than this before the whole of the starch is completely transformed. Even when the conversion of the starch commences very promptly, it is often a long time before it is finished. If a thin starch paste, for example, which contains no traces of sugar, be taken into the mouth and thoroughly mixed with the buccal secretions, it will often, as already mentioned, begin to show the reaction of sugar in the course of half a minute ; but some of the starchy matter still remains, and will continue to manifest its characteristic reaction with iodine, for fifteen or twenty minutes, or even half an hour. The above action of the saliva on starch, according to the experi- ments of Magendie, Bernard, Bidder and Schmidt, &c., does not reside in either the parotid, submaxillary or mucous secretions taken separately ; but only in the mixed saliva, as it comes from the cavity of the mouth. The submaxillary and mucous secretions, however, taken together, produce the change ; though neither of them has any effect alone, nor even when mixed artificially with the saliva of the parotid. It was supposed, when this property of converting starch into sugar was first discovered to exist in the saliva, that it constituted the true physiological action of the secretion, and that the function of the saliva was, in reality, the digestion and liquefaction of starchy substances. It was very soon noticed, however, by the French observers, that this property of the saliva was rather an accidental than an essential one ; and that, although starchy substances are really converted into sugar, if mixed with saliva in a test-tube, yet they are not affected by it to the same degree in the natural process of digestion. We have already mentioned above the extremely variable activity of the saliva, in this respect, at different times ; and it must be recollected, also, that in digestion the food is not retained in the cavity of the mouth, but passes at once, after mas- tication, into the stomach. Several German observers, as Frerichs, Jacubowitsch, Bidder, and. Schmidt, maintained at first that the saccharine conversion of starch, after being commenced in the mouth, might be, and actually was, completed in the stomach. We have convinced ourselves, however, by frequent experiments, that this is not the case. If a dog, with a gastric fistula, be fed with a 7 98 DIGESTIOISr. mixture of meat and boiled starch, and portions of the fluid con- tents of the stomacli withdrawn afterward through the fistula, the starch is easily recognizable by its reaction with iodine for ten, fifteen, and twenty minutes afterward. In forty-five minutes, it is diminished in quantity, and in one hour has usually altogether dis- appeared ; but no sugar is to be detected at any time. Sometimes the starch disappears more rapidly than this ; but at no time, accord- ing to our observations, is there any indication of the presence of sugar in the gastric fluids. Bidder and Smith have also concluded, from subsequent investigations,^ that the first experiments performed Tinder their direction by Jacubowitsch were erroneous; and it is now acknowledged by them, as well as by the Frenchi observers, that sugar cannot be detected in the stomach, after the introduction of starch, in any form or by any method. In the ordinary process of digestion, in fact, starcby matters do not remain long enough in the mouth to be altered by the saliva, but pass at once into the sto- mach. Here they meet with, the gastric fluids, which become min- gled with them, and prevent the change which would otherwise be effected by the saliva. We have found that the gastric juice will interfere, in this manner, with the action of the saliva in the test- tube, as well as in the stomach. If two mixtures be made, one of starch and saliva, the other of starch, saliva, and gastric juice, and botb kept for fifteen minutes at the temperature of 100° F., in the first mixture the starch will be promptly converted into sugar, while in the second no such change will take place. The above action, therefore, of saliva on starch, though a curious and interesting pro- perty, has no significance as to its physiological function, since it does not take place in the natural digestive process. We shall see hereafter that there are other means provided for the digestion of starchy matters, altogether independent of the action of the saliva. The true function of the saliva is altogether a physical one. Its action is simply to moisten the food and facilitate its mastication, as well as to lubricate the triturated mass, and assist its passage down the oesophagus. Food which is hard and dry, like crusts, crackers, &c,, cannot be masticated and swallowed with readiness unless moistened by some fluid. If the saliva, therefore, be prevented from entering the cavity of the mouth, its loss does not interfere directly with the chemical changes of the food in digestion, but only with its mechanical preparation. This is the result of direct experi- " Op. cit., p. 26. SALIVA. 99 ments performed by various observers. Bidder and Schmidt/ after tying Steno's duct, together with the common duct of the sub- maxillary and sublingual glands on both sides in the dog, found that the immediate effect of such an operation was "a remarkable diminution of the fluids which exude upon the surfaces of the mouth; so that these surfaces retained their natural moisture only so long as the mouth was closed, and readily became dry on exposure to contact with the air. Accordingly, deglutition became evidently difficult and laborious, not only for dry food, like bread, but even for that of a tolerably moist consistency, like fresh meat. The ani- mals also became very thirsty, and were constantly ready to drink," Bernard^ also found that the only marked effect of cutting off" the flow of saliva from the mouth was a difficulty in the mechani- cal processes of mastication and deglutition. He first administered to a horse one pound of oats, in order to ascertain the rapidity with which mastication would naturally be accomplished. The above quantity of grain was thoroughly masticated and swallowed at the end of nine minutes. An opening had been previously made in the oesophagus at the lower part of the neck, so that none of the food reached the stomach; but each mouthful, as it passed down the oesophagus, was received at the oesophageal opening and examined by the experimenter. The parotid duct on each side of the face was then divided, and another pound of oats given to the animal. Mastication and deglutition were both found to be immediately retarded. The alimentary masses passed down the oesophagus at longer intervals, and their interior was no longer moist and pasty, as before, but dry and brittle. Finally, at the end of twenty-five minutes, the animal had succeeded in masticating and swallowing only about three-quarters of the quantity which he had previously disposed of in nine minutes. It appears, also, from the experiments of Magendie, Bernard, and Lassaigne, on horses and cows, that the quantity of saliva absorbed by the food during mastication is in direct proportion to its hard- ness and dryness, but has no particular relation to its chemical qualities. These experiments were performed as follows : The oeso- phagus was opened at the lower part of the neck, and a ligature placed upon it, between the wound and the stomach. The animal was then supplied with a previously weighed quantity of food, and this, as it passed out by the oesophageal opening, was received into ' Op. cit., p. 3. ^ Lemons de Physiologie Experimeutale, Paris, 1856, p. 146. 100 DIGESTION. appropriate vessels and again weighed. The difference in weight, before and after swallowing, indicated the quantity of saliva absorbed by the food. The following table gives the results of some of Las- saigne's experiments,' performed upon a horse : — Kind of Food employed. Quantity of Saline absokbed. For 100 parts of liay there were absorbed 400 parts saliva. " barley meal " 186 " " oats " 113 " " green stalks and leaves " 49 " It is evident, from the above facts, that the quantity of saliva produced has not so much to do with the chemical character of the food as with its physical condition. When the food is dry and hard, and requires much mastication, the saliva is secreted in abundance ; when it is soft and moist, a smaller quantity of the secretion is poured out; and finally, when the food is taken in a fluid form, as soup or milk, or reduced to powder and moistened artificially with a very large quantity of water, it is not mixed at all with the saliva, but passes at once into the cavity of the stomach. The abundant and watery fluid of the parotid gland is most useful in assisting mastication; while the glairy and mucous secretion of the submaxillary gland and the muciparous follicles serve to lubri- cate the exterior of the triturated mass, and facilitate its passage through the oesophagus. By the combined operation of the two processes which the food undergoes in the cavity of the mouth, its preliminary preparation is accomplished. It is triturated and disintegrated by the teeth, and, at the same time, by the movements of the jaws, tongue, and cheeks, it is intimately mixed with the salivary fluids, until the whole is reduced to a soft, pasty mass, of the same consistency throughout. It is then carried backward by the semi-involuntary movements of the tongue into the pharynx, and conducted by the muscular contractions of the oesophagus into the stomach. THE GASTEIO JUICE, AND STOMACH DIGESTION. The mucous membrane of the stomach is distinguished by its great vascularity and the abundant glandular apparatus with which it is provided. Its entire thickness is occupied by certain glandular organs, the gastric tubules or follicles, which are so closely set as to leave almost no space between them except what is required for the ' Comptes Rendus, vol. xxi. p. 362. THE GASTRIC JUICE, AND STOMACH DIGESTION. 101 capillary bloodvessels. The free surface of the gastric mucous membrane is not smooth, but is raised in minute ridges and pro- jecting eminences. In the cardiac portion (Fig. 25), these ridges are reticulated with each other, so as to include between them Fig. 25. Fig. 26. Fig. 27. Fig. 25. Free surface of G.vstric Macous Me.mbrane, viewed from above ; from Pig's Stomach, Car- diac portion. Magnified 70 diameters. Fig. 26. Free surface of Gastric Mococts Me.mbrane, viewed in vertical section; from Pig's Stomacli, Pyloric portion. Magnified 420 diameters. polygonal interspaces, each of which is encircled by a capillary network. In the pyloric portion (Fig. 26), these eminences are more or less pointed and coni- cal in form, and generally flattened from side to side. They contain each a capillary bloodvessel, which returns upon itself in a loop at the extremity of the projection, and communicates freely with adjacent vessels. The gastric follicles are very different in different parts of the stomach. In the pyloric portion (Fig. 27), they are nearly straight, simple tubules, ^\^ of an inch in diameter, easily separated from each other, lined with glandular epithelium, and ter- minating in cul-de-sacs at the under surface of the mucous mem- Mncous Membrane of Pig's Stomach, from Pyloric portion; vertical section; showing gastric tubules, and, at a, a closed follicle. Magnified 70 diameters. 102 DIGESTION. Gastric Tdedles from Pig's Stomach, Pyloric por- tion, showing their C^ecal Extremities. At a, a cylin- drical cast of epithelium, pressed out from a tubule, showing the size of its cavity. brane. They are sometimes slightly branched, or provided with one or two rounded diverticula, a short distance above their ter- mination. They open on the free surface of the mucous membrane, in the interspaces between the projecting folds or villi. Among these tubular glandules there is also found, in the gastric mucous mem- brane, another kind of glandu- lar organ, consisting of closed follicles, similar to the solitary glands of the small intestine. These follicles, which are not very numerous, are seated in the lower part of the mucous membrane, and enveloped by the c£ecal extremities of the tubules. (Fig. 27, a.) In the cardiac portion of the stomach, the tubules are very wide in the superficial part of the mucous membrane, and lined with large, distinctly marked cylinder epithelium cells. (Fig. 29.) In the deeper parts of the membrane they become branched and considerably reduced in size. From the point where the branching takes place to where they terminate below in cul-de sacs, they are lined with small glandular epithelium cells, and closely bound together by intervening areolar tissue, so as to present somewhat the appearance of compound glandules. The bloodvessels which come up from the submucous layer of areolar tissue form a close plexus around all these glandules, and provide the mucous Gastric Tubules from Pig's Stomach ; Cardiac portion. At a, a large tubule dividing into two small ones. b. Portion of a tubule, seen endwise, c. Its central cavity. THE GASTEIC JUICE, AND STOMACH DIGESTION. 103 membrane with an abundant supply of blood, both for the purposes of secretion and absorption. That part of digestion which takes place in the stomach has always been regarded as nearly, if not quite, the most important part of the whole process. The first observers who made any approximation to a correct idea of gastric digestion were Reaumur and Spallanzani, who showed by various methods that the reduction and liquefaction of the food in the stomach could not be owing to a mere contact with the gastric mucous membrane, or to compression by the muscular walls of the organ ; but that it must be attributed to a digestive fluid secreted by the mucous membrane, which pene- trates the food and reduces it to a fluid form. They regarded this process as a simple chemical solution, and considered the gastric juice as a universal solvent for all alimentary substances. They succeeded even in obtaining some of this gastric juice, mingled probably with many impurities, by causing the animals upon which they experimented to swallow sponges attached to the ends of cords, by which they were afterwards withdrawn, and the fluids, which they had absorbed, expressed and examined. The first decisive experiments on this point, however, were those performed by Dr. Beaumont, of the U. S. Army, on the person of Alexis St. Martin, a Canadian boatman, who had a permanent gas- tric fistula, accidentally produced by a gunshot wound. The musket, which was loaded with buckshot at the time of the accident, was discharged at the distance of a few feet from St. Martin's body, in such a manner as to tear away the integument at the lower part of the left chest, open the pleural cavity, and penetrate, through the lateral portion of the diaphragm, into the great pouch of the stomach. After the integument and the pleural and peritoneal surfaces had united and cicatrized, there remained a permanent opening, of about four-fifths of an inch in diameter, leading into the left extremity of the stomach, which was usually closed by a circular valve of pro- truding mucous membrane. This valve could be readily depressed at any time, so as to open the fistula and allow the contents of the stomach to be extracted for examination. Dr. Beaumont experimented upon this person at various intervals from the year 1825 to 1832.^ He established during the course of his examinations the following important facts : First, that the ac. tive agent in digestion is an acid fluid, secreted by the walls of the ' Experiments and Observations on tlie Gastric Juice. Boston, 1834. 10^ DIGESTION", stomach ; secondly, that this fluid is poured out by the glandular walls of the organ only during digestion, and under the stimulus of the food ; and finally, that it will exert its solvent action upon the food outside the body as well as in the stomach, if kept in glass phials upon a sand bath, at the temperature of 100° F. He made also a variety of other interesting investigations as to the effect of various kinds of stimulus on the secretion of the stomach, the rapidity with which the process of digestion takes place, the com- parative digestibility of various kinds of food, &c. &c. Since Dr. Beaumont's time it has been ascertained that similar gastric fistulee may be produced at will on some of the lower animals by a simple operation ; and the gastric juice has in this way been obtained, usually from the dog, by Blondlot, Schwann, Bernard, Lehmann and others. The simplest and most expeditious mode of doing the operation is the best. An incision should be made through the abdominal parietes in the median line, over the great curvature of the stomach. The anterior wall of the organ is then to be seized with a pair of hooked forceps, drawn out at the external wound, and opened with the point of a bistoury. A short silver canula one-half to three-quarters of an inch in diameter, armed at each extremity with a narrow projecting rim or flange, is then in- serted into the wound in the stomach, the edges of which are fast- ened round the tube with a ligature in order to prevent the escape of the gastric fluids into the peritoneal cavity. The stomach is then returned to its place in the abdomen, and the canula allowed to re- main with its external flange resting upon the edges of the wound in the abdominal integuments, which are to be drawn together by sutures. The animal may be kept perfectly quiet, during the ope- ration, by the administration of ether or chloroform. In a few days the ligatures come awa}^, the wounded peritoneal surfaces are united with each other, and the canula is retained in a permanent gastric fistula ; being prevented by its flaring extremities both from falling out of the abdomen and from being accidentally pushed into the stomach. It is closed externally by a cork, which may be with- drawn at pleasure, and the contents of the stomach withdrawn for examination. Experiments conducted in this manner confirm, in the main, the results obtained by Dr. Beaumont. Observations of this kind are in ^ome respects, indeed, more satisfactory when made upon the lower animals, than upon the human subject; since animals are en- tirely under the control of the experimenter, and all sources of THE GASTRIC JUICE, AND STOMACH DIGESTION". 105 deception or mistake are avoided, while the investigation is, at the same time, greatly facilitated by the simple character of their food. The gastric juice, like the saliva, is secreted in considerable quantity only under the stimulus of recently ingested food. Dr. Beaumont states that it is entirely absent during the intervals of digestion ; and that the stomach at that time contains no acid fluid, but only a little neutral or alkaline mucus. He was able to obtain a sufficient quantity of gastric juice for examination, by gently irri- tating the mucous membrane with a gum-elastic catheter, or the end of a glass rod, and by collecting the secretion as it ran in drops from the fistula. On the introduction of food, he found that the mucous membrane became turgid and reddened, a clear acid fluid collected everywhere in drops underneath the layer of mucus lin- ing the walls of the stomach, and was soon poured out abundantly into its cavity. We have found, however, that the rule laid down by Dr. Beaumont in this respect, though correct in the main, is not invariable. The truth is, the irritability of the gastric mucous membrane, and the readiness with which the flow of gastric juice may be excited, varies considerably in different animals ; even in those belonging to the same species. In experimenting with gastric fistulae on different dogs, for example, we have found in one instance, like Dr. Beaumont, that the gastric juice was always entirely absent in the intervals of digestion ; the mucous membrane then present- ing invariably either a neutral or slightly alkaline reaction. In this animal, which was a perfectly healthy one, the secretion could not be excited by any artificial means, such as glass rods, metallic catheters, and the like ; but only by the natural stimulus of ingested food. We have even seen tough and indigestible pieces of tendon, introduced through the fistula, expelled again in a few minutes, one after the other, without exciting the flow of a single drop of acid fluid; while pieces of fresh meat, introduced in the same way, pro- duced at once an abundant supply. In other instances, on the con- trary, the introduction of metallic catheters, &c.j into the empty stomach has produced a scanty flow of gastric juice ; and in experi- menting upon dogs that have been kept without food during various periods of time and then killed by section of the medulla oblongata, we have usually, though not always, found the gastric mucous mem- brane to present a distinctly acid reaction, even after an abstinence of six, seven, and eight days. There is at no time, however, under these circumstances, any considerable amount of fluid present in 106 DIGESTION. the stomacli ; but onl}^ just sufficient to moisten the gastric mucous membrane, and give it an acid reaction. The gastric juice, which is obtained by irritating the stomach with a metallic catheter, is clear, perfectly colorless, and acid in reaction. A sufficient quantity of it cannot be obtained by this method for any extended experiments ; and for that purpose, the animal should be fed, after a fast of twenty-four hours, with fresh lean meat, a little hardened by short boiling, in order to coagulate the fluids of the muscular tissue, and prevent their mixing with the gastric secretion. No effect is usually apparent within the first five minutes after the introduction of the food. At the end of that time the gastric juice begins to flow ; at first slowly, and in drops. It is then perfectly colorless, but very soon acquires a slight amber tinge. It then begins to flow more freely, usually in drops, but often running for a few seconds in a continuous stream. In this way from lij to ^iiss may be collected in the course of fifteen minutes. After this it becomes somewhat turbid with the debris of the food, which has begun to be disintegrated ; but from this it may be readily separated by filtration. After three hours, it con- tinues to run freely, but has become very much thickened, and even grumous in ' consistency, from the abundant admixture of alimentary debris. In six hours after the commencement of diges- tion it runs less freely, and in eight hours has become very scanty, though it continues to preserve the same physical appearances as before. It ceases to flow altogether in from nine to twelve hours, according to the quantity of food taken. For purposes of examination, the fluid drawn during the first fifteen minutes after feeding should be collected, and separated by filtration from accidental impurities. Obtained in this way, the gastric juice is a clear, watery fluid, without any appreciable vis- cidity, very distinctly acid to test paper, of a faint amber color, and with a specific gravity of 1010. It becomes opalescent on boiling, owing to the coagulation of its organic ingredients. The following is the composition of the gastric juice of the dog, based on a comparison of various analyses by Lehmann, and Bidder and Schmidt: — THE GASTRIC JUICE, AND STOMACH DIGESTIOISr. 107 Composition of Gastric Juice. Water 975.00 Organic matter ........ 15.00 Lactic acid ......... 4.78 Chloride of sodium . . . . . . . . 1.70 " " potassium . . . . • . • 1.08 " " calcium 0.20 " " ammoniura ....... 0.65 Phosphate of lime ........ 1.48 " " magnesia 0.06 " " iron 0.05 1000.00 In place of lactic acid, Bidder and Sclimidt found, in most of their analyses, hydrochloric acid. Lehmann admits that a small quantity of hydrochloric acid is sometimes present, but regards lactic acid as much the most abundant and important of the two. Robin and Verdeil also regard the acid reaction of the gastric juice as due to lactic acid ; and, finally, Bernard has shown,^ by a series of well contrived experiments, that the free acid of the dog's gastric juice is undoubtedly the lactic ; and that the hydrochloric acid obtained by distillation, is really produced by a decomposition of the chlorides, which enter into the composition of the fresh juice. The free acid is an extremely important ingredient of the gastric secretion, and is, in fact, essential to its physiological properties; for the gastric juice will not exert its solvent action upon the food, after it has been neutralized by the addition of an alkali or an alkaline carbonate. The most important ingredient of the gastric juice, beside the free acid, is its organic matter or " ferment," which is sometimes known under the name of -pepsine. This name, "pepsine," was originally given by Schwann to a substance which he obtained from the mucous membrane of the pig's stomach, by macerating it in distilled water until a putrid odor began to be developed. The substance in question was precipitated from the watery infusion by the addition of alcohol, and dried ; and if dissolved afterward in acidulated water, it was found to exert a solvent action on boiled white of Qgg. This substance, however, did not represent precisely the natural ingredient of the gastric secretion, and was probably a mixture of various matters, some of them the products of com- mencing decomposition of the mucous membrane itself. The name pepsine, if it be used at all, should be applied to the organic matter ' Le9ons de Physiologie Experimentale, Paris, 1856, p. 396. 108 DIGESTION. wTaicTi naturally occurs in solution in the gastric juice. It is alto- gether unessential, in this respect, from what source it may be originally derived. It has been regarded by Bernard and others, o'n somewhat insufficient grounds, as a product of the alteration of the mucus of the stomach. But whatever be its source, since it is always present in the secretion of the stomach, and takes an active part in the performance of its function, it can be regarded in ifo other light than as a real anatomical ingredient of the gastric juice, and as essential to its constitution. Pepsine is precipitated from its solution in the gastric juice by absolute alcohol, and by various metallic salts, but is not affected by ferrocyanide of potassium. It is precipitated also, and coagu- lated by a boiling temperature; and the gastric juice, accordingly, after being boiled, becomes turbid, and loses altogether its power of dissolving alimentary substances. Gastric juice is also affected in a remarkable manner by being mingled with Ule. We have found that four to six drops of dog's bile precipitate completely with 5j of gastric juice from the same animal; so that the whole of the biliary coloring matter is thrown down as a deposit, and the filtered fluid is found to have lost entirely its digestive power, though it retains an acid reaction. .A very singular property of the gastric juice is its inaptitude for putrefaction. It may be kept for an indefinite length of time in a common glass- stoppered bottle without developing any putrescent odor. A light deposit generally c^ollects at the bottom, and a con- fervoid vegetable growth or "mould" often shows itself in the fluid after it has been kept for one or two weeks. This growth has the form of white, globular masses, each of which is com- posed of delicate radiating branched filaments (Fig. 30) ; each filament consisting of a row of elongated cells, like other vegetable growths of a similar nature. These growths, however, are not accompa- nied by any putrefactive changes, and the gastric juice retains its acid reaction and CONFERVOID Vegetable, growing iu the Gastric Juice itS digCStivC prOpCrticS for of the Dog. The fibres have an average diameter of fi^nT-i-y- months ] -7000th of an inch. -^ THE GASTRIC JUICE, AND STOMACH DIGESTION. 109 By experimenting artificially with gastric juice on various ali- mentary substances, such as meat, boiled white of egg, &c., it is found, as Dr. Beaumont formerly observed, to exert a solvent action on these substances outside the body, as well as in the cavity of the stomach. This action is most energetic at the temperature of 100° F. It gradually diminishes in intensity below that point, and ceases altogether near 32°. If the temperature be elevated above 100° the action also becomes enfeebled, and is entirely suspended about 160°, or the temperature of coagulating albumen. Contrary to what was supposed, however, by Br. Beaumont and his predeces- sors, the gastric juice is not a universal solvent for all alimentary substances, but on the contrary, affects only a single class of the proximate principles, viz., the albuminoid or organic substances. Neither starch nor oil, when digested in it at the temperature of the body, suffers the slightest chemical alteration. Fatty matters are simply melted by the heat, and starchy substances are only hydrated and gelatinized to a certain extent by the combined influence of the warmth and moisture. Solid and semi-solid albuminoid matters, however, are at once attacked and liquefied by the digestive fluid. Pieces of coagulated white of egg sus- pended in it, in a test-tube, become gradually softened on their exterior, and their edges become pale and rounded. They grow thin and transparent ; and their substance finally melts away, leaving a light, scanty deposit, which collects at the bottom of the test-tube. While the disintegrating process is going on, it may be almost always noticed that minute, opaque spots show themselves in the substance of the liquefying albumen, indicating that certain parts of it are less easily attacked than the rest ; and the deposit which remains at the bottom is probably also composed of some ingre- dient, not soluble in the gastric juice. If pieces of fresh meat be treated in the same manner, the areolar tissue entering into its com- position is first dissolved, so that the muscular bundles become more distinct, and separate from each other. They gradually fall apart, and a little brownish deposit is at last all that remains at the bottom of the tube. If the hard, adipose tissue of beef or mutton be sub- jected to the same process, the walls of the fat vesicles and the intervening areolar tissue, together with the capillary bloodvessels, &;c., are dissolved ; while the oily matters are set free from their envelops, and collect in a white, opaque layer on the surface. In cheese, the casein is dissolved, and the oil set free, as above. In bread, the gluten is digested, and the starch left unchanged. In 110 DIGESTIOISr. milk, the casein is first coagulated by contact with the acid gastric fluids, and afterward slowly liquefied, like other albuminoid sub- stances. The time required for the complete liquefaction of these sub- stances varies with the quantity of matter present, and with its state of cohesion. The process is hastened by occasionally shaking up the mixture, so as to separate the parts already disintegrated, and bring the gastric fluid into contact with fresh portions of the digestible substance. The liquefying process which the food undergoes in the gastric juice is not a sknple solution. It is a catalytic transformation, produced in the albuminoid substances by contact with the or- ganic matter of the digestive fluid. This organic matter acts in a similar manner to that of the catalytic bodies, or "ferments," generally. Its peculiarity is that, for its active operation, it re- quires to be dissolved in an acidulated fluid. In common with other ferments, it requires also a moderate degree of warmth ; its action being checked, both by a very low, and a very high tempe- rature. By its operation the albuminoid matters of the food, what- ever may have been their original character, are all, without dis- tinction, converted into a new substance, viz., albuminose. This substance has the general characters belonging to the class of organic matters. It is uncrystallizable, and contains nitrogen as an ultimate element. It is precipitated, like albumen, by an excess of alcohol, and by the metallic salts ; but unlike albumen, is not affected by nitric acid or a boiling temperature. It is freely soluble in water, and after it is once produced by the digestive process, remains in a fluid condition, and is ready to be absorbed by the vessels. In this way, casein, fibrin, musculine, gluten, &c., are all reduced to the condition of albuminose. By experimenting as above, with a mixture of food and gastric juice in test tubes, we have found that the casein of cheese is entirely converted into ^albuminose, and dissolved under that form. A very considerable portion of raw white of egg, however, dissolves in the gastric juice directly as albumen, and retains its property of coagulating by heat. Soft-boiled white of egg and raw meat are principally con- verted into albuminose; but at the same time, a small portion of albumen is also taken up unchanged. The above process is a true liquefaction of the albuminoid sub- stances, and not a simple disintegration. If fresh meat be cut into small pieces, and artificially digested in gastric juice in test tubes. THE GASTRIC JUICE, AND STOMACH DIGESTION. Ill at 100° F., and the process assisted by occasional gentle agitation, the fluid continues to take up more and more of the digestible material for from eight to ten hours. At the end of that time if it be separated and filtered, the filtered fluid has a distinct, brownish color, and is saturated with dissolved animal matter. Its specific gravity is found to have increased from 1010 to 1020 ; and on the addition of alcohol it becomes turbid, with a very abundant whitish precipitate (albuminose). There is also a peculiar odor developed during this process, which resembles that produced in the malting of barley. Albuminose, in solution in gastric juice, has several peculiar properties. One of the most remarkable of these is that it inter- feres with the operation of Trommer's test for grape sugar (see page 52). We first observed and described this peculiarity in 1854,' but could not determine, at that time, upon what particular ingredient of the gastric juice it depended. A short time subse- quently it was also noticed by M. Longet, in Paris, who published his observations in the Gazette Sebdomadaire for February 9th, 1855.^ He attributed the reaction not to the gastric juice itself, but to the albuminose held in solution by it. We have since found this explanation to be correct. Gastric juice obtained from the empty stomach of the fasting animal, by irritation with a metallic catheter, which is clear and perfectly colorless, does not interfere in any way with Trommer's test ; but if it be macerated for some hours in a test-tube with finely chopped meat, at a temperature of 100°, it will then be found to have acquired the property in a marked degree. The reaction therefore depends undoubtedly upon the presence of albuminose in solution. As the gastric juice, drawn from the dog's stomach half an hour or more after the introduction of food, already contains some albuminose in solution, it presents the same reaction. If such gastric juice be mixed with a small quantity of glucose, and Trommer's test applied, no peculiarity is observed on first dropping in the sulphate of copper ; but on adding afterward the solution of potass, the mixture takes a rich purple hue, instead of the clear blue tinge, which is presented under ordinary circumstances. On boiling, the color changes to claret, cherry red, and finally to a light yellow; but no oxide of copper is deposited, and the fluid remains clear. If the albuminose be present only in small ' American Journ. Med. Sci., Oct. 1854, p. 319. * Nouvelles reclierches relatives a Taction du sue gastrique sur les substances albuminoides. — Gaz. Hebd. 9 F^vrier, 1855, p. 103. 112 DIGESTION. , quantity, an incomplete reduction of the copper takes place, so that the mixture becomes opaline and cloudy, but still without any well marked deposit. This interference will take place when sugar is present in very large proportion. We have found that in a mix- ture of honey and gastric juice in equal volumes, no reduction of copper takes place on the application of Trommer's test. It is remarkable, however, that if such a mixture be previously diluted with an equal quantity of water, the interference does not take place, and the copper is deposited as usual. Usually this peculiar reaction, now that we are acquainted with its existence, will not practically prevent the detection of sugar, when present ; since the presence of the sugar is distinctly indi- cated by the change of color, as above mentioned, from purple to yellow, though the copper may not be thrown down as a precipi- tate. All possibility of error, furthermore, may be avoided by adopting the following precautions. The purple color, already men- tioned, will, in the first place, serve to indicate the presence of the albuminoid ingredient in the suspected fluid. The mixture should then be evaporated to dryness, and extracted with alcohol, in order to eliminate the animal matters. After that, a watery solution of the sugar contained in the alcoholic extract will react as usual with Trommer's test, and reduce the oxide of copper without difficulty. Another remarkable property of gastric juice containing albu- minose, which is not, however, peculiar to it, but common to many other animal fluids, is that of interfering with the m.utual reaction of starch and iodine. If 5j of such gastric juice be mixed with 3j of iodine water and boiled starch be subsequently added, no blue color is produced; though if a larger quantity of iodine water be added, or if the tincture be used instead of the aqueous solution, the superabundant iodine then combines with the starch, and pro- duces the ordinary blue color. This property, like that described above, is not possessed by pure, colorless, gastric juice, taken from the empty stomach, but is acquired by it on being digested with albuminoid substances. Another important action which takes place in the stomach, beside the secretion of the gastric juice, is the peristaltic movement of the organ. This movement is accomplished by the alternate contraction and relaxation of the longitudinal and circular fibres of its muscular coat. The motion is minutely described by Dr. Beaumont, who examined it both by watching the movements of the food through the gastric fistula, and also by introducing into THE GASTRIC JUICE, AND STOMACH DIGESTION. 113 the stomach the bulb and stem of a thermometer. According to his observations, when the food first passes into the stomach, and tlie secretion of the gastric juice commences, the muscular coat, which was before quiescent, is excited and begins to contract ac- tively. The contraction takes place in such a manner that the food, after entering the cardiac orifice of the stomach, is first car- ried to the left, into the great pouch of the organ, thence downward and along the great curvature to the pyloric portion. At a short distance from the p^dorus. Dr. B. often found a circular constric- tion of the parietes, by which the bulb of the thermometer was gently grasped and drawn toward the pylorus, at the same time giving a twisting motion to the stem of the instrument, by which it was rotated in his fingers. In a moment or two, however, this constriction was relaxed, and the bulb of the thermometer again released, and carried together with the food along the small curva- ture of the organ to its cardiac extremity. This circuit was re- peated so long as anyibod remained in the stomach; but as the liquefied portions were successively removed toward the end of digestion it became less active, and at last ceased altogether when the stomach had become completely empty, and the organ returned to its ordinary quiescent condition. It is easy to observe the muscular action of the stomach during digestion in the dog, by the assistance of a gastric fistula, artificially established. If a metallic catheter be introduced through the fistula when the stomach is empty, it must usually be held care- fully in place, or it will fall out by its own weight. But immedi- ately upon the introduction of food, it can be felt that the catheter is grasped and retained with some force, by the contraction of the muscular coat. A twisting or rotatory motion of its extremity may also be frequently observed, similar to that described by Dr. Beaumont. This peristaltic action of the stomach, however, is a gentle one, and not at all active or violent in character. We have never seen, in opening the abdomen, any such energetic or exten- sive contractions of the stomach, even when full of food, as may be easily excited in the small intestine by the mere contact of the atmosphere, or by pinching them with the blades of a forceps. This action of the stomach, nevertheless, though quite gentle, is sufficient to produce a constant churning movement of the masti- cated food, by which it is carried back and forward to every part of the stomach, and rapidly incorporated with the gastric juice which is at the same time poured out by the mucus membrane ; so 8 114 DIGESTION. that the digestive fluid is made to penetrate equally every part of the alimentary mass, and the digestion of all its albuminous ingre- dients goes on simultaneously. This gentle and continuous move- ment of the stomach is one which cannot be successfully imitated in experiments on artificial digestion with gastric juice in test-tubes; and consequently the process, under these circumstances, is never so rapid or so complete as when it takes place in the interior of the stomach. The length of time which is required for digestion varies in different species of animals. In the carnivora a moderate meal of fresh uncooked meat requires from nine to twelve hours for its complete solution and disappearance from the stomach. According to Dr. Beaumont, the average time required for digestion in the human subject is considerably less; varying from one hour to five hours and a half, according to the kind of food employed. This is probably owing to the more complete mastication of the food which takes place in man, than in the carnivorous animals. By examining the contents of the stomach at various intervals after feeding. Dr. Beaumont made out a list, showing the comparative digestibility of different articles of food, of which the following are the most important: — Time required for digestion, according to Dr. Beaumont: — Kind of Food. Hours. Minutes. Pig's feet 1 00 Tripe 1 00 Trout (broiled) 1 30 Venison steak ........ 1 35 Milk 2 00 Roasted turkey 2 ^ 30 " beef 3 00 " mutton ........ 3 15 Veal (broiled) 4 00 Salt beef (boiled) .4 15 Roasted pork 5 15 The comparative digestibility of different substances varies more or less in different individuals according to temperament ; but the above list undoubtedly gives a correct average estimate of the time required for stomach digestion under ordinary conditions. A very interesting question is that which relates to the total gwanfe'^y of gastric juice secreted daily. Whenever direct experi- ments have been performed with a view of ascertaining this point, their results have given a considerably larger quantity than was anticipated. Bidder and Schmidt found that, in a dog weighing THE GASTRIC JUICE, AND STOMACH DIGESTION. 115 34 pounds, tliey were able to obtain by separate experiments, con- suming in all 12 hours, one pound and three-quarters of gastric juice. The total quantity, therefore, for 24 hours, in the same animal, would be 3| pounds; and,. by applying the same calcula- tion to a man of medium size, the authors estimate the total daily quantity in the human subject as but little less than 14 pounds (av.). This estimate is probably not an exaggerated one. In order to determine the question, however, if possible, in a different way, we adopted the following plan of experiment with the gastric juice of the dog. It was first ascertained, by direct experiment, that the fresh lean meat of the bullock's heart loses, by complete desicca- tion, 78 per cent, of its weight. 800 grains of such meat, cut into small pieces, were then digested for ten hours, in |iss of gastric juice at 100° F.; the mixture being thoroughly agitated as often as every hour, in order to insure the digestion of as large a quan- tity of meat as possible. The meat remaining undissolved was then collected on a previously weighed filter, and evaporated to dryness. The dry residue weighed 55 grains. This represented, allowing for the loss by evaporation, 250 grains of the meat, in its natural moist condition ; 50 grains of meat were then dissolved by siss of gastric juice, or 33 J grains per ounce. From these data we can form some idea of the large quantity of gastric juice secreted in the dog during the process of digestion. One pound of meat is only a moderate meal for a medium-sized animal ; and yet, to dissolve this quantity, no less than thirteen pints of gastric juice will be necessary. This quantity, or any approxi- mation to it, would be altogether incredible if we did not recollect that the gastric juice, as soon as it has dissolved its quota of food, is immediately reabsorbed, and again enters the circulation, together with the alimentary substances jsvhich it holds in solution. The secretion and reabsorption of the gastric juice then go on simulta- neously ; and the fluids which the blood loses by one process are incessantly restored to it by the other. A very large quantity, therefore, of the secretion may be poured out during the digestion of a meal, at an expense to the blood, at any one time, of only two or three ounces of fluid. The simplest investigation shows that the gastric juice does not accumulate in the stomach in any con- siderable quantity during digestion ; but that it is gradually secreted so long as any food remains undissolved, each portion, as it is digested, being disposed of by reabsorption, together with its solvent fluid. There is accordingly, during digestion, a constant 116 DIGESTION. circulation of the digestive fluids from the bloodvessels to the alimentary canal, and from the alimentary canal back again to the bloodvessels. That this circulation really does take place is proved by the fol- lowiog facts : First, if a dog be killed some hours after feeding, there is never more than a very small quantity of fluid found in the stomach, just sufficient to smear over and penetrate the half digested pieces of meat ; and, secondly, in the living animal, gastric juice, drawn from the fistula five or six hours after digestion has been going on, contains little or no more organic matter in solution than that extracted fifteen to thirty minutes after the introduction of food. It has evidently been freshly secreted ; and, in order to obtain gastric juice saturated with alimentary matter, it must be artificially digested with food in test tubes, where this constant absorption and renovation cannot take place. An unnecessary difficulty has sometimes been felt in understand- ing how it is that the gastric juice, which digests so readily all albuminous substances, should not destroy the walls of tbe stomach itself, which are composed of similar materials. This, in fact, was brought forward at an early day, as an insuperable objection to the doctrine of Reaumur and Spallanzani, that digestion was a process of chemical solution performed by a digestive fluid. It was said to be impossible that a fluid capable of dissolving animal matters should be secreted by the walls of the stomach without attacking them also, and destroying the organ by which it was produced. Since that time, various complicated hypotheses have been framed, in order to reconcile these apparently contradictory facts. The true explanation, however, as we believe, lies in this — that the process of digestion is not a simple solution, but a catalytic transformation of the alimentary 'substances, produced by contact with the pep- sine of the gastric juice. We know that all the organic sub- stances in the living tissues are constantly undergoing, in the process of nutrition, a series of catalytic changes, which are charac- teristic of the vital operations, and which are determined by the organized materials with which they are in contact, and by all the other conditions present in the living organism. These changes, therefore, of nutrition, secretion, &c., necessarily exclude for the time all other catalyses, and take precedence of them. In the same way, any dead organic matter, exposed to warmth, air, and moist- ure, putrefies; but if immersed in gastric juice, at the same temperature, the putrefactive changes are stopped or altogether THE GASTRIC JUICE, AND STOMACH DIGESTION. 117 prevented, because the catalytic actions, excited by the gastric juice, take precedence of those which constitute putrefaction. For a similar reason, the organic ingredient of the gastric juice, which acts readily on dead animal matter, has no effect on the living tissues of the stomach, because they are already subject to other catalytic influences, which exclude those of digestion, as well as those of putrefaction. As soon as life departs, however, and the peculiar actions taking place in the living tissues come to an end with the stoppage of the circulation, the walls of the stomach are really attacked by the gastric juice remaining in its cavity, and are more or less completely digested and liquefied. In the human subject, it is rare to make an examination of the body twenty-four or thirty-six hours after death, without finding the mucous mem- brane of the great pouch of the stomach more or less softened and disintegrated from this cause. Sometimes the mucous membrane is altogether destroyed, and the submucous cellular layer exposed ; and occasionally, when death has taken place suddenly during active digestion, while the stomach contained an abundance of gastric juice, all the coats of the organ have been found destroyed, and a perforation produced leading into the peritoneal cavity. These post-mortem changes show that, after death, the gastric juice really dissolves the coats of the stomach without difficulty. But during life, the chemical changes of nutrition, which are going on in their tissues, protect them from its influence, and effectually preserve their integrity. The secretion of the gastric juice is much influenced by nervous conditions. It was noticed by Dr. Beaumont, in his experiments upon St. Martin, that irritation of the temper, and other moral causes, would frequently diminish or altogether suspend the supply of the gastric fluids. Any febrile action in the system, or any unusual fatigue, was liable to exert a similar effect. Every one is aware how readily any mental disturbance, such as anxiety, anger, or vexation, will take away the appetite and interfere with diges- tion. Any nervous impression of this kind, occurring at the com- ■mencement of digestion, seems moreover to produce some change which has a lasting effect upon the process ; for it is very often noticed that when any annoyance, hurry, or anxiety occurs soon after the food has been taken, though it may last only for a few moments, the digestive process is not only liable to be suspended for the time, but to be permanently disturbed during the entire, day. In order that digestion, therefore, may go on properly in the 118 . DIGESTION. « stomacli, food must be taken only when the appetite demands it ; it should also be thoroughly masticated at the outset ; and, finally, both mind and body, particularly during the commencement of the process, should be free from any unusual or disagreeable excite- ment. INTESTINAL JUICES, AND THE DIGESTION OF SUGAR AND STARCH. From the stomach, those portions of the food which have not already suffered digestion, pass into the third division of the ali- mentary canal, viz., the small intestine. As already mentioned, it is only the albuminous matters which are digested jn the stomach. Cane sugar, it is true, is slowly converted by the gastric juice, out- side the body, into glucose. We have found that ten grains of cane sugar, dissolved in §ss of gastric juice, give traces of glucose at the end of two hours; and in three hours, the quantity of this substance is considerable. It cannot be shown, however, that the gastric juice exerts this effect on sugar during ordinary digestion. If pure cane sugar be given to a dog with a gastric fistula, while digestion of meat is going on, it disappears in from two to three hours, without any glucose being detected in the fluids withdrawn from the stomach. It is, therefore, either directly absorbed under the form of cane sugar, or passes, little by little, into the duodenum, where the intestinal fluids at once convert it into glucose. "It is equally certain that starchy .matters are not digested in the stomach, but pass unchanged into the small intestine. Here they meet with the mixed intestinal fluids, which act at once upon the starch, and convert it rapidly into sugar. The intestinal fluids, taken from the duodenum of a recently killed dog, exert this transforming action upon starch with the greatest promptitude, if mixed with it in a test-tube and kept at the temperature of 100° F. Starch is converted into sugar by this, means much more rapidly and certainly than by the saliva; and experiment shows that the intestinal fluids are the active agents in its digestion during life. If a dog be fed with a mixture of meat and boiled starch, and killed a short time after the meal, the stomach is found to contain starch but no sugar; while in the small intestine there is an abundance of sugar, and but little or no starch. If some observers have failed to detect sugar in the intestine after feeding the animal with starch, it is because they have delayed the examination until too INTESTINAL JUICES, DIGESTION OF SUGAR, ETC. 119 late. For it is remarkable how rapidly starchy substances, if previously disintegrated by boiling, are disposed of in the digest- ive process. If a dog, for example, be fed as above with boiled starch and meat, while some of the meat remains in the stomach for eight, nine, or ten hours, the starch begins immediately to pass into the intestine, where it is at once converted into sugar, and then as rapidly absorbed. The whole of the starch may be converted into sugar, and completely absorbed, in an hour's time. We have even found, at the end of three-quarters of an hour, after a tolerably full meal of boiled starch and meat, that all trace of both starch and sugar had disappeared from both stomach and intestine. The rapidity with which this passage of the starch into the duodenum takes place varies, to some extent, in different animals, according to the general activity of the digestive appa- ratus; but it is always a comparatively rapid process, when the starch is already liquefied and is administered in a pure form. There can be no doubt that the natural place for the digestion of starchy matters is the small intestine, and that it is accomplished by 'the action of the intestinal juices. Our knowledge is not very complete with regard to the exact nature of the fluids by which this digestion of the starch is accom- plished. The juices taken from the duodenum are generally a mixture of three different secretions, viz., the bile, the pancreatic fluid, and the intestinal juice proper. Of these, the bile may be left out of the question; since it does not, when in a pure state, exert any digestive action on starch. The pancreatic juice, on the other hand, has the property p. gi of converting starch into su- gar; but it is not known whether this fluid be always present in the duodenum. The true intestinal juice is the product of two sets of glan- dular organs, seated in the substance of or beneath the mucous membrane, viz., the follicles of LieberkiJhn and the glands of Brunner. The first of these, or Lieberkiihn's follicles (Fig. 31), are the most numerous. They are simple, follicles op lieberkuhx, from smaii in- '' ^ testine of Pig. 120 DIGESTION. Dearly straight tubules, lined witli columnar epithelium, and some- what similar in their appearance to the follicles of the pyloric portion of the stomach. They occupy the whole thickness of the mucous membrane, and are found in great numbers throughout the entire length of the small and large intestine. The glands of Brunner (Fig. 32), or the duodenal glandulse, as they are sometimes called, are Fig. 32. ^ confined to the upper part of the duodenum, where they exist, as a closely set layer, in the deeper portion of the mucous membrane, extending downward a short distance from the pylorus. They are composed of a great number of rounded follicles, or culs-de- sac, clustered round a central excretory duct. Each follicle consists of a delicate mem- branous wall, lined with glan- dular epithelium, and covered on its surface with small, dis- tinctly marked nuclei. The follicles collected round each duct are bound together by a thin layer of areolar tissue, and covered with a plexus of capillary bloodvessels. The intestinal jaice, which is the secreted product of the above glandular organs, has been less successfully studied than the other digestive fluids, owing to the difficulty of obtaining it in a pure state. The method usually adopted has been to make an opening in the abdomen of the living animal, take out a loop of intestine, empty it by gentle pressure, and then to shut off a portion of it from the rest of the intestinal cavity by a couple of ligatures, situated six or eight inches apart; after which the loop is returned into the abdomen, and the external wound closed by sutures. After sis or eight hours the animal is killed, and the fluid, which has collected in the isolated portion of intestine, taken out and examined. The above was the method adopted by Frerichs. Bid- der and Schmidt, in order to obtain pure intestinal juice, first tied the biliary and pancreatic ducts, so that both the bile and the pan- creatic juice should be shut out from the intestine, and then estab- Portion of one of Brtsner's Duodexal Glands, from Human Intestine. PANCREATIC JUICE, AND THE DIGESTION OF FAT. 121 lished an intestinal fistula below, from which they extracted the fluids which accumulated in the cavity of the gut. From the great abundance of the follicles of Lieberkiihn, we should expect to find the intestinal juice secreted in large quantity. It appears, however, in point of fact, to be quite scanty, as the quantity collected in the above manner by experimenters has rarely been sufficient for a thorough examination of its properties. It seems to resemble very closely, in its physical characters, the secretion of the mucous fol- licles of the mouth. It is colorless and glassy in appearance, viscid and mucous in consistency, and has a distinct alkaline reaction. It has the property, when pure, as well as when mixed with other secretions, of rapidly converting starch into sugar, at the tempe- rature of the living body. PANCREATIC JUICE, AND THE DIGESTION OF FAT, The only remaining ingredients of the food that require digestion are the oily matters. These are not affected, as we have already stated, by contact with the gastric juice ; and examination shows, furthermore, that they are not digested in the stomach. So long as they remain in the cavity of this organ they are unchanged in their essential properties. They are merely melted by the warmth of the stomach, and set free by the solution of the vesicles, fibres, or capillary tubes in which they are contained, or among which they are entangled ; and are still readily discernible by the eye, floating in larger or smaller drops on the surface of the semi-fluid alimentary mass. Yery soon, however, after its entrance into the intestine, the oily portion of the food loses its characteristic ap- pearance, and is converted into a white, opaque emulsion, which is gradually absorbed. This emulsion is termed the chyle^ and is always found in the small intestine during the digestion of fat, entangled among the valvulse conniventes, and adhering to the surface of the villi. The digestion of the oil, however, and its con- version into chyle, does not take place at once upon its entrance into the duodenum, but only after it has passed the orifices of the pancreatic and biliary ducts. Since these ducts almost invariably open into the intestine at or near the same point, it was for a long time dif&cult to decide by which of the two secretions the digestion of the oil was accomplished. M. Bernard, however, first threw some light on this question by experimenting on some of the lower 122 DIGESTION. animals, in whidi the two ducts open separately. In the rabbit, for example, the biliary duct opens as usual just below the pylorus, while the pancreatic duct communicates with the intestine some eight or ten inches lower down. Bernard fed these animals "with substances containing oil, or injected melted butter into the stomach; and, on killing them afterward, found that there was no chyle in the intestine between the opening of the biliary and pancreatic ducts, but that it was abundant immediately below the orifice of the latter. Above this point, also, he fo'und the lacteals empty or transparent, while below it they were full of white and opaque chyle. The result of these experiments, which have since been confirmed by Prof. Samuel Jackson of Philadelphia,' lead to the conclusion that the pancreatic fluid is the active agent in the diges- tion of oily substances; and an examination of the properties of this secretion, when obtained in a pure state from the living animal, fully confirm the above opinion. In order to obtain pancreatic juice from the dog, the animal must be etherized soon after digestion has commenced, an incision made in the upper part of the abdomen, a little to the right of the median line, and a loop of the duodenum, together with the lower extremity of the pancreas which lies adjacent to it, drawn out at the external wound. The pancreatic duct is then to be exposed and opened, and a small silver canula inserted into it and secured by a ligature. The whole is then returned into the abdomen and the wound closed by sutures, leaving only the end of the canula projecting from it. In the dog there are two pancreatic ducts, situated from half an inch to an inch apart. The lower one of these, which is usually the larger of the two, is the one best adapted for the insertion of the canula. After the eSects of etherization have passed off, and the digestive process has recommenced, the pancreatic juice begins to run from the orifice of the canula, at first very slowly and in drops. Sometimes the drops follow each other with rapidity for a few moments, and then an interval occurs during which the secretion seems entirely suspended. After a time it recommences, and continues to exhibit similar fluctuations during the whole course of the experiment. Its flow, however, is at all times scanty, compared with that of the gastric juice ; and we have never been able to collect at most more than five or six drachms during a period of several hours. Bidder and Schmidt obtained ' American Journ. Med. Sci., Oct. 1854. PANCREATIC JUICE, AND THE DIGESTION OF FAT. 123 on an average, from a dog weighing 44 pounds, 14|- grains of pan- creatic juice per hour ; and they calculate from this that the average daily quantity in the human subject is rather less than 2500 grains, or a little over one-third of a pound avoirdupois. Pancreatic juice obtained by the above process is a clear, color- less, somewhat viscid fluid, with a distinct alkaline reaction. Its composition, according to the analysis of Bidder and Schmidt, is as follows : — Composition of Pancreatic Juice. Water 900.76 Organic matter (pancreatine) ....... 90.38 Chloride of sodium ......... 7.36 Free soda .......... 0.32 Phosphate of soda 0.45 Sulphate of soda . 0.10 . Sulphate of potass ■ . . 0.02 r Lime 0.54 Combinations of } Magnesia ....... 0.05 y Oxide of iron 0.02 1000.00 The most important ingredient of the pancreatic juice is its organic matter or pancreatine. It will be seen that this is much more abundant in proportion to the other ingredients of the secre- tion than the organic matter of any other digestive fluid. It is coagulable by heat; and the pancreatic juice often solidifies com- pletely on boiling, like white of egg, so that not a drop of fluid remains after its coagulation. It is precipitated, furthermore, by nitric acid and by alcohol, and also by sulphate of magnesia in excess. By this last property, it may be distinguished from albu- men, which is not affected by contact with sulphate of magnesia. Fresh pancreatic juice, brought into contact with oily matters at the temperature of the body, exerts upon them, as was first noticed by Bernard, a very peculiar effect. It disintegrates them, and re- duces them to a state of complete emulsion, so that the mixture is at once converted into a white, opaque, creamy-looking fluid. This effect is instantaneous and permanent, and only requires that the two substances be well mixed by gentle agitation. It is singular that some of the German observers should deny that the pancreatic juice possesses this property, of emulsioning fat, to a greater extent than the bile and some other digestive fluids; and should state that although, when shaken up with oil, outside the body, it reduces the oily particles to a state of extreme minuteness, the emulsion 124 DIGESTION. is not permanent, and the oily particles "soon separate again on the surface.'" We have frequently repeated this experiment with different specimens of pancreatic juice obtained from the dog, and have never failed to see that the emulsion produced by it is by far more prompt and complete than that which takes place with saliva, gastric juice, or bile. The effect produced by these fluids is in fact altogether insignificant, in comparison with the prompt and energetic action exerted by the pancreatic juice. The emulsion produced with the latter secretion may be kept, furthermore, for at least twenty-four hours, according to our observations, without any appreciable separation of the oily particles, or a return to their original condition. The pancreatic juice, therefore, is peculiar in its action on oily substances, and reduces them at once to the condition of an emul- sion. The oil, in this process, does not suffer any chemical altera- tion. It is not decomposed or saponified, to any appreciable extent. It is simply emulsioned; that is, it is broken up into a state of minute subdivision, and retained in suspension by contact with the organic matter of the pancreatic juice. That its chemical condition is not altered is shown by the fact that it is still soluble in ether, which will withdraw the greater part of the fat from a mixture of oil and pancreatic juice, as well as from the chyle in the interior of the intestine. In a state of emulsion the fat, furthermore, is capable of being absorbed, and its digestion may be then said to be accom- plished. We find, then, that the digestion of the food is not a simple operation, but is made up of several different processes, which commence successively in different portions of the alimentary canal. In the first place, the food is subjected in the mouth to the physical operations of mastication and insalivation. Eeduced to a soft pulp and mixed abundantly with the saliva, it passes, secondly, into the stomach. Here it excites the secretion of the gastric juice, by the influence of which its chemical transformation and solution are commenced. If the meal consist wholly or partially of mus- cular flesh, the first effect of the gastric juice is to dissolve the intervening cellular substance, by which the tissue is disintegrated and the muscular fibres separated from each other. Afterward the muscular fibres themselves become swollen and softened by the imbibition of the gastric fluid, and are finally disintegrated and liquefied. In the small intestine, the pancreatic and intestinal ' Lehmann's Physiological Chemistry. Philada. ed., vol. i. p. 507. PHENOMENA OF INTESTINAL DIGESTION. 125 juices convert the starchy ingredients of the food into sugar, and break up the fatty matters into a fine emulsion, by which they are converted into chyle. Although the separate actions of these digestive fluids, however, commence at different points of the alimentary canal, they after- ward go on simultaneously in the small intestine ; and the changes Avhich take place here, and which constitute the process of intestinal digestion, form at the same time one of the most complicated and one of the most important parts of the whole digestive function. The phenomena of intestinal digestion may be studied, in the dog, by killing the animal at various periods after feeding, and examining the contents of the intestine. We have also succeeded, by establishing, in the same animal, an artificial intestinal fistula, in gaining still more satisfactory information on this point. The fistula may be established, for this purpose, by an operation precisely similar to that already described as employed for the production of a permanent fistula in the stomach. The silver tube having been introduced into the lower part of the duodenum, the wound is allowed to heal, and the intestinal secretions may then be with- drawn at will, and subjected to examination at different periods during digestion. By examining in this way, from time to time, the intestinal fluids, it at once becomes ^manifest that the action of the gastric juice, in the digestion of albuminoid substances, is not confined to the stomach, but continues after the food has passed into the intes- tine. About half an hour after the injection of a meal, the gastric juice begins to pass into the duodenum, where it may be recognized by its strongly-marked acidity, and by its peculiar action, already described, in interfering with Trommer's test for grape sugar. It has accordingly already dissolved some of the ingredients of the food while still in the stomach, and contains a certain quantity of albuminose in solution. It soon afterward, as it continues to pass into the duodenum, becomes mingled with the debris of muscular fibres, fat vesicles, and oil drops; substances which are easily recognizable under the microscope, and which produce a grayish turbidity in the fluid drawn from the fistula. This turbid admix- ture becomes thicker and thicker from the second to the tenth or twelfth hour; after which the intestinal fluids become rapidly less abundant, and finally disappear almost entirely, as the process of digestion comes to an end. The passage of disintegrated muscular tissue into the intestine 126 DIGESTION. CosTENTS OP Stomach dueixg Digestion OF Meat, from the Dog. — a. Fat Vesicle, filled with opaque, solid, granular fat. b, h. Bits of partially dis- integrated muscular fibre, c. Oil globules. may also be shown, as already mentioned, by killing the animal and examining the contents of the alimentary canal. During the digestion of muscular flesh ^^s- 33. 2,xi(\ adipose tissue, the sto- mach contains masses of soft- ened meat, smeared over with gastric juice, and also a mo- derate quantity of grayish grumous fluid, with an acid reaction. This fluid contains muscular fibres, isolated from each other, and more or less disintegrated by the action of the gastric juice. (Fig. 33.) The fat vesicles are but little or not at all altered in the stomach, and there are only a few free oil globules to be seen floating in the mixed fluids, contained in the cavity of the organ. In the duodenum the muscular fibres are further disintegrated. (Fig. 3J:.) They become very much broken up, pale and transparent, but can still ^^^' be recognized by the granu- lar markings and striations which are characteristic of them. The fat vesicles also begin to become altered in the duodenum.. The solid granular fat of beef, and simi- lar kinds of meat, becomes liquefied and emulsioned; and appears under the form of free oil drops and fatty mole- cules; while the fat vesicle itself is partially emptied, and From Dtodexum of Dog, ditrixg Diges- bcCOmeS morC Or IcSS Col- TiON OF Meat. — a. Fat vesicle, with its. contents , j i i • n l T„ j-U^ diminishing. The vesicle is beginning to shrivel and lapSCd and shriVClled. In the the fat breaking up. 6, 6. Disintegrated muscular middle and loWCr partS of the fibre, e, c. Oil globules. . . ,.p,. n~ j o/j\ intestine (Figs. 3o and 36) these changes continue. The muscular fibres become constantly PHENOMENA OF INTESTINAL DIGESTION. 127 Fig. 35. more and more disintegrated, and a large quantity of granular debris is produced, wliich is at last also dissolved. The fat also progressively disappears, and the vesicles may be seen in the lower part of the intes- tine, entirely collapsed and empty. In this way the digestion of tlie different ingredients of the food goes on in a con- tinuous manner, from the sto- mach throughout the entire length of the small intestine. At the same time, it results in the production of three diflerent substances, viz : 1st. Albuminose, produced by the ^ ,, o . ' '■ ... From Middle of Small Intestine. — a, a. action of the gastric juice Fat vesicles, nearly emptied of their contents. on the albuminoid matters; 2d. An oily emulsion, pro- duced by the action of the pancreatic juice on fat; and, 3d. Sugar, produced from the transformation of starch by the mixed intestinal fluids. These substances are then ready to be taken up into the circulation ; and the next change which they undergo, in the regular course of the vital processes, is that of ah- sorption. This process will form the subject of the next chapter. Fig. 36. From last quarter of Small Intestine. -a, a. Fat vesicles, quite empty and shrivelled. 128 ABSORPTION". CHAPTEK VII. ABSORPTION. Beside the glands of Brunner and the follicles of Lieberkiihii, already described, there are, in the inner part of the walls of the intestine, certain glandular- Fig. 37. One of the closed Follicles of Peter's Patches, from Small Intestine of Pig. Magnified 50 diameters. Fig. 38. Glanduls; Agminat;e, from Small Intestine of Pig. Magnified 20 diameters. looking bodies which are termed " glandulse solitariae," and " glandulae agminatse." The glandulse solitariae are globular or ovoid bodies, about one-thirtieth of an inch in diameter, situated partly in, and partly beneath, the intestinal mucous membrane. Each glandule (Fig. 37) is formed of an investing cap- sule, closed on all sides, and containing in its interior a soft pulpy mass, which con- sists of minute cellular bodies, imbedded in a homogeneous substance. The inclosed mass is penetrated by capillary bloodvessels, which penetrate through the investing cap- sule, inosculate freely with each other, and return upon themselves in loops, near the centre of the glandular body. There is no external opening or duct ; in fact, the contents of the vesicle, being pulpy and vascular, as already de- scribed, are not to be regarded ABSORPTION. 129 as a secretion, but as constituting a kind of solid glandular tissue. The gland ul^e agminate (Fig. 38), or " Payer's patches," as they are sometimes called, consist of aggregations of similar globular or ovoid bodies, found most abundantly toward the lower extremity of the small intestine. Both the solitary and agminated glandules are evidently connected with the lacteals and the system of the mesen- teric glands, which latter organs they resemble very much in their minute structure. They are probably to be regarded as the first row of mesenteric glands, situated in the walls of the intestinal canal. Another set of organs, intimately connected with the process of absorption, are the villi of the small intestine. These are conical vascular eminences of the mucous membrane, thickly set over the whole internal surface of the small intestine. In the upper portion of the intestine, they are flattened and triangular in form, resembling somewhat the conical projections of the pyloric portion of the sto- mach. In the lower part, they are long and filiform, and often slightly enlarged, or club-shaped, at their free extremity (Fig. 89), and frequently attaining the length of one-thirty-fifth of an inch. They are covered externally with a layer of columnar epithelium, similar to that which lines the rest of the intestinal mucous membrane, and contain in their interior two sets of vessels. The most superficial of these are the capillary bloodvessels, which are supplied in each villus by a twig of the mesenteric artery, and which form, by their fre- quent inosculation, an exceedingly close and abundant network, almost imme- diately beneath the epithelial layer. The}^ unite at the base of the villus, and form a minute vein, which is one of the commencing rootlets of the por- tal vein. In the central part of the vil- lus, and lying nearly in its axis, there is another vessel, with thinner and more transparent walls, which is the commencement of a lacteal. The precise manner in which the lacteal originates in the extremity of the villus is not known. It commences near the apex, either by a 9 Extremity of I x testis a i. Villus; from the Dog. — u. Layer of epithelium, b. Bloodvessel, c. Lacteal vessel. 130 ABSORPTION. blind extremity or by an irregular plexus, passes, in a straight or somewhat wavy line, toward the base of the villus, and then becomes continuous with a small twig of the mesenteric lacteals. The villi are the active agents in the process of absorption. By their projecting form, and their great abundance, they increase enor- mously the extent of surface over which the digested fluids come in contact with the intestinal mucous membrane, and increase, also, to a corresponding degree, the energy with which absorption takes place. They hang out into the nutritious, semi-fluid mass contained in the intestinal cavity, as the roots of a tree penetrate the soil; and they imbibe the liquefied portions of the food, with a rapidity which is in direct proportion to their extent of surface, and the activity of their circulation. The process of absorption is also hastened by the peristaltic movements of the intestine. The muscular layer here, as in other parts of the alimentary canal, is double, consisting of both circular and longitudinal fibres. The action of these fibres may be readily seen by pinching the exposed intestine with the blades of a forceps. A contraction then takes place at the spot irritated, by which the intestine is reduced in diameter, its cavity obliterated, and its con- tents forced onward into the succeeding portion of the alimentary canal. The local contraction then propagates itself to the neighbor- ing parts, while the portion originally contracted becomes relaxed ; so that a slow, continuous, creeping motion of the intestine is pro- duced, by successive waves of contraction and relaxation, which follow each other from above downward. At the same time, the longitudinal fibres have a similar alternating action, drawing the narrowed portions of intestine up and down, as they successively enter into contraction, or become relaxed in the intervals. The effect of the whole is to produce a peculiar, writhing, worm-like, or "vermicular" motion, among the different coils of intestine. During life, the vermicular or peristaltic motion of the intestine is excited by the presence of food undergoing digestion. By its action, the substances which pass from the stomach into the intestine are steadily carried from above downward, so as to traverse the entire length of the small intestine, and to come in contact successively with the whole extent of its mucous membrane. During this pas- sage, the absorption of the digested food is constantly going on. Its liquefied portions are taken up by the villi of the mucous mem- brane, and successively disappear; so that, at the termination of the small intestine, there remains only the undigestible portion of the ABSORPTION. 131 food, together, with the refuse of the intestinal secretions. These pass through the ileo-ca?cal orifice into the large intestine, and there become reduced to the condition of feces. The absorption of the digested fluids is accomplished both by the bloodvessels and the lacteals. It was formerly supposed that the lacteals were the only agents in this process; but it has now been long known that this opinion was erroneous, and that the bloodvessels take at least an equal part in absorption, and are in some respects the most active and important agents of the two. Abundant experiments have demonstrated not only that soluble substances introduced into the intestine may be soon afterward detected in tlie blood of the portal vein, but that absorption takes place more rapidly and abundantly by the bloodvessels than it does by the lacteals. The most decisive of these experiments were those performed by Panizza on the abdominal circulation.' This observer opened the abdomen of a horse and drew out a fold of the small intestine, eight or nine inches in length (Fig. 40, a, a), ¥m. 40. P A>-izz a"s ExPEK IME XT. — art. Intestine, b. Point of ligature of iiie.«Rnteric vein, n Opening in intestine for introduction of poison, d. Opening in mesenteric vein behind the ligature. which he included between two ligatures. A ligature was then placed (at b) upon the mesenteric vein receiving the blood from this portion of intestine; and, in order that the circulation might not be interrupted, an opening was made (at d) in the vein behind ' In Matteucci's Lectures on the Physical Phenomena of Living Beings, Pereira's edition, p. 83. 132 ABSORPTION. the ligature, so that the blood brought by the mesenteric artery, after circulating in the intestinal capillaries, passed out at the opening, and was collected in a vessel for examination. Hydro- cyanic acid was then introduced into the intestine by an opening at c, and almost immediately afterward its presence was detected in the venous blood flowing from the orifice at d. The animal, how- ever, was not poisoned, since the acid was prevented from gaining an entrance into the general circulation by the ligature at h. Panizza afterward varied this experiment in the following manner: Instead of tying the mesenteric vein, he simply com- pressed it. Then, hydrocyanic acid being introduced into the intestine as above, no effect was produced on the animal, so long as compression was maintained upon the vein. But as soon as the blood was allowed to pass again through the vessels, symptoms of general poisoning at once became manifest. Lastly, in a third experiment, the same observer removed all the nerves and lacteal vessels supplying the intestinal fold, leaving the bloodvessels alone untouched. Hydrocyanic acid now being introduced into the intes- tine, found an entrance at once into the general circulation, and the animal was immediately poisoned The bloodvessels, therefore, are not only capable of absorbing fluids from the intestine, but may even take them up more rapidly and abundantly than the lacteals. These two sets of vessels, however, do not absorb all the alimen- tary matters indiscriminately. It is one of the most important of the facts which have been established by modern researches on digestion that tbe different substances, produced by the operation of the digestive fluids on the food, pass into the circulation by different routes. The fatty matters are taken up by the lacteals under the form of chyle, while the saccharine and albuminous matters pass by ab- sorption into the portal vein. Accordingly, after the digestion of a meal containing starchy and animal matters mixed, albuminose and sugar are botb found in the blood of the portal vein, while they can- not be detected, in any large quantity, in the contents of the lacteals. These substances, bowever, do not mingle at once with the general mass of the circulation, but owing to the anatomical distribution of the portal vein, pass first through the capillary circulation of the liver. Soon after being introduced into the blood, and coming in contact with its organic ingredients, they become altered and con- verted, by catalytic transformation, into other substances. The albuminose passes into the condition of ordinary albumen, and probably also partly into that of fibrin ; while the sugar rapidly ABSORPTION, 133 Fig. 41. becomes decomposed, and loses its characteristic properties; so that, on arriving at the entrance of the general circulation, both these newly absorbed ingredients have become already assimilated to those which previously existed in the blood. The chyle in the intestine consists, as we have already mentioned, of oily matters which have not been chemically altered, but simply reduced to a state of emulsion. In chyle drawn from the lacteals or the thoracic duct (Fig. 41), it still presents itself in the same condition and retains all the chemical properties of oil. Examined by the microscope, it is seen to exist under the form of fine granules and molecules, which present the ordinary appearances of oil in a state of minute subdivi- sion. The chyle, therefore, does not represent the entire product of the digestive pro- cess, but contains only the fatty substances, suspended by emulsion in a serous fluid. During the time that intes- tinal absorption is going on, after a meal containing fatty ingredients, the lacteals may be seen as white, opaque vessels, dis- tended with milky chyle, passing through the mesentery, and con- verging from its intestinal border toward the receptaculum chyli, near the spinal column. During their course, they pass through several successive rows of mesenteric glands, which also become turgid with chyle, while the process of digestion is going on. The lacteals then conduct the chyle to the receptaculum chyli, whence it passes upward through the thoracic duct, and is finally dis- charged, at the termination of this canal, into the left subclavian vein. (Fig. 42.) It is then mingled with the returning current of venous blood, and passes into the right cavities of the heart. The lacteals, however, are not a special system of vessels by them- selves, but are simply a part of the great system of " absorbent" or " lymphatic" vessels, which are to be found everywhere in the integu- ments of the head, the parietes of the trunk, the upper and lower extremities, and in the muscular tissues and mucous membranes Chyle from c o m m e x c e .m e x t o f Thoracic PtJCT, from the Dog. — The molecules viiiy iu size from 1-1 0,000th of au inch dowu-ward. 134 ABSORPTION. throughout the body. The walls of these vessels are thinner and more transparent than those of the arteries and veins, and they 'are consequently less easily de- tected by ordinary dissection. They originate in the tissues of the above-mentioned parts by an irregular plexus. They pass from the extremities to- ward the trunk, converging and uniting with each other like the veins, their principal branches taking usually the same direc- tion with the nerves and blood- vessels, and passing, at various points in their course, through certain glandular bodies, the "lymphatic" or "absorbent" glands. The lymphatic glands, among which are included the mesenteric glands, consist of an external layer of fibrous tissue and a contained pulp or paren- chyma. The investing layer of fibrous tissue sends off thin septa or laminae from its inter- nal surface, which penetrate the substance of the gland in every direction and unite with each other at various points. In this way they form an interlacing laminated framework, which divides the substance of the gland into numerous roianded spaces or alveoli. These alveoli are not completely isolated, but communi- cate with each other by narrow openings, where the intervening septa are incomplete. These cavities are filled with a soft, reddish pulp, which is penetrated, according to Kolliker, like the solitary and agminated glands of the intestine, by a fine network of capil- lary bloodvessels. The solitary and agminated glands of the intestine are, therefore, closely analogous in their structure to the lymphatics. The former are to be regarded as simple, the latter as compound vascular glands. The arrangement of the lymphatic vessels in the interior of the Lacteals, Thoracic Duct, &c. — a. Intes- tine, h. Vena cava inferior, c, c. Kight and left subclavian veins, d. Point of opening of thoracic duct into left subclavian. ABSOEPTION. 135 glands is not precisely understood. Each lymphatic vessel, as it enters the gland, breaks up into a number of minute ramifications, the vasa afferentia ; and other similar twigs, forming the vasa effer- entia^ pass off in the opposite direction, from the further side of the gland ; but the exact mode of communication between the two has not been definitely ascertained. All the fluids, however, arriving by the vasa afferentia, must pass in some way through the tissue of the gland, before they are carried away again by the vasa efferentia. From the lower extremities the lymphatic vessels enter the abdomen at the groin and converge toward the receptaculum chyli, into which their fluid is discharged, and afterward conveyed, by the thoracic duct, to the left subclavian vein. The fluid which these vessels contain is called the lymph. It is a colorless or slightly yellowish transparent fluid, which is absorbed by the lymphatic vessels from the tissues in which they originate. But little is known regarding its composition, except that it con- tains, beside water and saline matters, a small quantity of fibrin and albumen. Its ingredients are evidently derived from the meta- morphosis of the tissues, and are returned to the centre of the circulation in order to be eliminated by excretion, or in order to undergo some new transforming or renovating process. We are ignorant, however, with regard to the precise nature of their charac- ter and destination. The lacteals are simply that portion of the absorbents which originate in the mucous membrane of the small intestine. During the intervals of digestion, these vessels contain a colorless and transparent lymph, entirely similar to that which is found in other parts of the absorbent system. After a meal containing only starchy or albuminoid substances, there is no apparent change in the character of their contents. But after a meal containing fatty matters, these substances are taken up by the absorbents of the intestine, which then become filled with the white chylous emul- sion, and assume the appearance of lacteals, (Fig. 48.) It is for this reason that lacteal vessels do not show themselves upon the stomach nor upon the first few inches of the duodenum ; because oleaginous matters, as we have seen, are not digested in the stomach, but only after they have entered the intestine and passed the orifice of the pancreatic duct. The presence of chyle in the lacteals is, therefore, not a con- stant, but only a periodical phenomenon. The fatty substances constituting the chyle begin to be absorbed during the process of 136 ABSORPTION. digestion, as soon as they have been disintegrated and emulsioned by the action of the intestinal fluids. As digestion proceeds, they accumulate in larger quantity, and gradually fill the whole lacteal Fig. 43. LACTEALS AXD Lr-MPHATICS. system and the thoracic duct. As they are discharged into the subclavian vein, and mingled with the blood, they can still be dis- tinguished in the circulating fluid, as a mixture of oily molecules and granules, between the orifice of the thoracic duct and the right side of the heart. While passing through the pulmonary circula- ABSORPTIOlSr. 137 tion, however, they disappear. Precisely what becomes of them, or what particular chemical changes they undergo, is not certainly known. They are, at all events, so altered in the blood, while passing through the lungs, that they lose the form of a fatty emul- sion, and are no longer to be recognized by the usual tests for oleaginous substances. The absorption of fat from the intestine is not, however, exclu- sively performed by the lacteals. Some of it is also taken up, under the same form, by the bloodvessels. It has been ascertained by the experiments of Bernard' that the blood of the mesenteric veins, in the carnivorous animals, contains, during intestinal diges- tion, a considerable amount of fatty matter in a state of minute subdivision. Other observers, also (Lehmann, Schultz, Simon), have found the blood of the portal vein to be considerably richer in fat than that of other veins, particularly while intestinal digestion is going on with activity. In birds, reptiles, and fish, furthermore, according to Bernard, the intestinal lymphatics are never filled with opaque chyle, but only with a transparent lymph; so that these animals may be said to be destitute of lacteals, and in them the fatty substances, like other alimentary materials, are taken up altogether by the bloodvessels. In quadrupeds, on the other hand, and in the human subject, the absorption of fat is accomplished both by the bloodvessels and the lacteals. A certain portion is taken up by the former, while the superabundance of the fatty emulsion is absorbed by the latter. A difficulty has long been experienced in accounting for the absorp- tion of fat from the intestine, owing to its being considered as a non- endosmotic substance; that is, as incapable, in its free or undissolved condition, of penetrating and passing through an animal membrane by endosmosis. It is stated, indeed, that if a fine oily emulsion be placed on one side of an animal membrane in an endosmometer, and pure water on the other, the water will readily penetrate the substance of the membrane, while the oily particles cannot be made to pass, even under a high pressure. Though this be true, how- ever, for pure water, it is not true for slightly alkaline fluids, like the serum of the blood and the lymph. This has been demon- strated by the experiments af Matteucci, in which he made an emulsion with an alkaline fluid containing 43 parts per thou- sand of caustic potass. Such a solution has no perceptible alkaline ' Lemons de Physiologic Experiinentale. Paris, 1'56, p. 325. 138 ABSORPTION. taste, and its action on reddened litmus paper is about equal to that of the lymph and chyle. If this emulsion were placed in an endosmometer, together with a watery alkaline solution of similar strength, it was found that the oily particles penetrated through the animal membrane without much difficulty, and mingled with the fluid on the opposite side. Although, therefore, we cannot explain the exact mechanism of absorption in the case of fat, still we know that it is not in opposition to ^^' * the ordinary phenomena of endosmosis; for endosmosis will take place with a fatty emulsion, provided the fluids used in the experiment be slightly alkaline in reaction. It is, accordingly, by a pro- cess of endosmosis, or imbi- bition, that the villi take up the digested fatty substances. There are no open orifices or canals, into which the oil penetrates; but it passes di- rectly into and through the substance of the villi. The epithelial cells covering the external surface of the villus are the first active agents in this absorption. In the intervals of digestion (Fig. 44) these cells are but slightly Intestinal Epithelium; from the Dog, -while fasting. Fig. 45. Intestinal Epithelium; from the Dog, dur- ing the digestion of fat. granular and nearly trans- parent in appearance. But if examined during the diges- tion and absorption of fat (Fig, 45), their substance is seen to be crowded with oily particles, which they have taken up from the intestinal cavity by absorption. The oily matter then passes on- ward, penetrating deeper and deeper into the substance of the villus, until it is at last received by the capillary ves- sels and lacteals in its centre. ABSOEPTION. 139 The fatty substances taken up by the portal vein, like those ab- sorbed by the lacteals, do not at once enter the general circulation, but pass first through the capillary system of the liver. Thence they are carried, with the blood of the hepatic vein, to the right side of the heart, and subsequently through the capillary system of the lungs. During this passage they become altered in character, as above described, and lose for the most part the distinguishing characteristics of oily matter, before they have passed beyond the pulmonary circulation. But as digestion proceeds, an increasing quantity of fatty matter finds its way, by these two passages, into the blood; and a time at last arrives when the whole of the fat so introduced is not destroyed during its passage through the lungs. Its absorption taking place at this time more rapidly than its decomposition, it begins to ap- pear,, in moderate quantity, in the blood of the general circulation ; and, lastly, when the intestinal absorption is at its point of greatest activity, it is found in considerable abundance throughout the entire vascular system. At this period, some hours after the inges- tion of food rich in oleaginous matters, the blood of the general circulation everywhere contains a superabundance of fat, derived from the digestive process. If blood be then drawn from the veins or arteries in any part of the body, it will present the pecu- liar appearance known as that of " chylous" or " milky" blood. After the separation of the clot, the serum presents a turbid ap- pearance; and the fatty substances, which it contains, rise to the top after a few hours, and cover its surface with a partially opaque and creamy-looking pellicle. This appearance has been occasion- ally observed in the human subject, particularly in bleeding for apoplectic attacks occurring after a full meal, and has been mis- taken, in some instances, for a morbid phenomenon. It is, however, a perfectly natural one, and depends simply on the rapid absorp- tion, at certain periods of digestion, of oleaginous substances from the intestine. It can be produced at will, at any time, in the dog, by feeding him with fat meat, and drawing blood, seven or eight hours afterward, from the carotid artery or the jugular vein. This state of things continues for a varying length of time, according to the amount of oleaginous matters contained in the food. When digestion is terminated, and the fat ceases to be intro- duced in unusual quantity into the circulation, its transformation and decomposition continuing to take place in the blood, it dis- appears gradually from the veins, arteries, and capillaries of the 140 ABSORPTION. general system; and, finally, when the whole of the fat has been disposed of by the nutritive processes, the serum again becomes transparent, and the blood returns to its ordinary condition. In this manner the nutritive elements of the food, prepared for absorption by the digestive process, are taken up into the circulation under the different forms of albuminose, sugar, and chyle, and accu- mulate as such, at certain times, in the blood. But these conditions are only temporary, or transitional. The nutritive materials soon pass, by catalytic transformation, into other forms, and become assimilated to the pre-existing elements of the circulating fluid. They thus accomplish finally the whole object of digestion ; which is to replenish the blood by a supply of new materials from without. There are, however, two other intermediate processes, taking place partly in the liver and partly in the intestine, at about the same time, and having for their object the final preparation and perfec- tion of the circulating fluid. These two processes require to be studied, before we can pass on to the particular description of the blood itself. They are : 1st, the secretion and reabsorption of the bile ; and 2d, the production of sugar in the liver, and its subse- quent decomposition in the blood. THE BILE. 141 CHAPTER VIII. THE BILE. The bile is more easily obtained in a state of purity than any other of the secretions which find their way into the intestinal canal, owing to the existence of a gall-bladder in which it accu- mulates, and from which it may be readily obtained without any other admixture than the mucus of the gall-bladder itself. Not- withstanding this, its study has proved an unusually difficult one. This difficulty has resulted from the peculiar nature of the biliary ingredients, and the readiness with which they become altered by chemical manipulation; and it is, accordingly, only quite recently that we have arrived at a correct idea of its real constitution. The bile, as it comes from the gall-bladder, is a somewhat viscid and glutinous fluid, varying in color and specific gravity according to the species of animal from which it is obtained. Human bile is of a dark golden brown color, ox bile of a greenish yellow, pig's bile of a nearly clear yellow, and dog's bile of a deep brown. We have found the specific gravity of human bile to be 1018, that of ox bile 1024, that of pig's bile 1030 to 1036. The reaction of the bile with test-paper cannot easily be determined ; since it has only a bleaching or decolorizing effect on litmus, and does not turn it either blue or red. It is probably either neutral or very slightly alkaline. A very characteristic physical property of the bile is that of frothing up into a soap-like foam when shaken in a test- tube, or when air is forcibly blown into it through a small glass tube or blowpipe. The bubbles of foam, thus produced, remain for a long time without breaking, and adhere closely to each other and to the sides of the glass vessel. The following is an analysis of the bile of the ox, based on the calculations of Berzelius, Frerichs, and Lehmann : — 142 THE BILE. Composition of Ox Bile. Water 880.00 Gljko-cholate of soda ........ \ Tauro-cholate " " } ^^'^^ Biliverdine ..........") Fats Oleates, margarates, and stearates of soda and potass . Cholesterin ......... Chloride of sodium ........ ^ Phosphate of soda ....... " " lime " " magnesia ...... Carbonates of soda and potass . . . • . . Mucus of the gall-bladder . . . . . . . .1.34 1- 13.42 15.24 1000.00 Biliverdine. — Of the above mentioned ingredients, hiliverdine is peculiar to the bile, and therefore important, though not pre- sent in large quantity. This is the coloring matter of the bile. It is, like the other coloring matters, an uncrystallizable organic substance, containing nitrogen, and yielding to ultimate analysis a small quantity of iron. It exists in such small quantity in the bile that its exact proportion has never been determined. It is formed, so far as can be ascertained, in the substance of the liver, and does not pre-exist in the blood. It may, however, be reabsorbed in cases of biliary obstruction, when it circulates with the blood and stains nearly all the tissues and fluids of the body, of a peculiar lemon yellow color. This is the symptom which is characteristic of jaundice. Cholesterin (CjjHgjO). — This is a crystallizable substance which resembles the fats in many respects ; since it is destitute of nitrogen, readily inflammable, soluble in alcohol and ether, and entirely in- soluble in water. It is not saponifiable, however, by contact with the alkalies, and is distinguished on this account from the ordinary fatty substances. It occurs, in a crystalline form, mixed with color- ing matter, as an abundant ingredient in most biliary calculi ; and is found also in different regions of the body, forming a part of various morbid deposits. We have met with it in the fluid of hydrocele, and in the interior of many encysted tumors. The crystals of cholesterin (Fig. 46) have the form of very thin, color- less, transparent, rhomboidal plates, portions of which are often cut out by lines of cleavage parallel to the sides of the crystal. They frequently occur deposited in layers, in which the outlines of THE BILE. 143 Cholesterin is not formed in the Fig. 46. the subjacent crystals show very distinctly through the substance of those which are placed above liver, but originates in the substance of the brain and nervous tissue, from which it may be extracted in large quantity by the action of alcohol. From these tissues it is absorbed by the blood, then conveyed to the liver, and discharged with the bile. The fatty substances and inorganic saline ingredients of the bile require no special description. C HO LESTER IX from an Encysted Tumor. Biliary Salts. — By far the most important and characteristic ingredients of this secretion are the two saline substances mentioned above as the glyko-cholate and tauro-cholate of soda. These substances were first discovered by Strecker, in 1848, in the bile of the ox. They are both freely soluble in water and in alcohol, but insoluble in ether. One of them, the tauro-cholate, has the property, when itself in solution in water, of dissolving a certain quantity of fat ; and it is probably owing to this circumstance that some free fat is present in the bile. The two biliary substances are obtained from ox-bile in the follow- ing manner : — The bile is first evaporated to dryness by the water-bath. The dry residue is then pulverized and treated with absolute alcohol, in the proportion of at least 5j of alcohol to every five grains of dry residue. The filtered alcoholic solution has a clear yellowish color. It contains, beside the glyko-cholate and tauro-cholate of soda, the coloring matter and more or less of the fats originally present in the bile. On the addition of a small quantity of ether, a dense, whitish precipitate is formed, which disappears again on agitating and thoroughly mixing the fluids. On the repeated addition of ether, the precipitate again falls down, and when the ether has been added in considerable excess, six to twelve times the volume of the alcoholic solution, the precipitate remains permanent, and the whole mixture is filled with a dense, whitish, opaque deposit, consisting of 144 THE BILE. the glyko-cholate and tauro-cholate of soda, thrown down under the form of heavy flakes and granules, part of which subside to the bottom of the test-tube, while part remain for a time in suspen- sion. Gradually these flakes and granules unite with each other and fuse together into clear, brownish-yellow, oily, or resinous- looking drops. At the bottom of the test-tube, after two or three hours, there is usually collected a nearly homogeneous layer of this deposit, while the remainder continues to adhere to the sides of the glass in small, circular, transparent drops. The deposit is semi-fluid in consistency, and sticky, like Canada balsam or half- rnelted resin ; and it is on this account that the ingredients compos- ing it have been called the " resinous matters" of the bile. They have, however, no real chemical relation with true resinous bodies, since they both contain nitrogen, and differ from resins also in other important particulars. At the end of twelve to twenty-four hours, the glyko-cholate of soda begins to crystallize. The crystals radiate from various points in the resinous deposit, and shoot upward into the supernatant fluid, in white, silky bundles. (Fig, 47.) If some of these crystals Fig. 47 Fig. 48. Ox -BILE, extracted with absolute alcohol and precipitated with ether. Gt.tko-cholate of Soda from Ox-bile, after two days' crystallization. At the lower part of the figure the crystals are melting into drops, from the evaporation of the ether and absorption of moisture. be removed and examined by the microscope, they are found to be of a very delicate acicular form, running to a finely pointed extremity, and radiating, as already mentioned, from a central THE BILE. 145 point. (Fig. 48.) As the ether evaporates, the crystals absorb moisture from the air, and melt up rapidly into clear resinous drops; so that it is difficult to keep them under the microscope long enough for a correct drawing and measurement. The crystal- lization in the test-tube goes on after the first day, and the crystals increase in quantity for three or four, or even five or six days, until the whole of the glyko-cholate of soda present has assumed the solid form. The tauro-cholate, however, is uncrystallizable, and remains in an amorphous condition. If a portion of the deposit be now removed and examined by the microscope, it is seen that the crystals of gl3^ko-cholate of soda have increased conside- ^^' rably in thickness (Fig. 49), so that their transverse dia- meter may be readily esti- mated. The uncrystallizable tauro-cholate appears under the form of circular drops, varying considerably in size, clear, transparent, strongly refractive, and bounded by a dark, well-defined outline. These drops are not to he distin- guished, by any of their ojyiical properties, from oil-glohides, as they usually appear under the microscope. They have the same refractive power, the same dark outline and bright centre, and the same degree of consistency. They would consequently be liable at all times to be mistaken for oil-globules, were it not for the complete dissimilarity of their chemical properties. Both the glyko-cholate and tauro-cholate of soda are very freely soluble in water. If the mixture of alcohol and ether be poured ofl' and distilled water added, the deposit dissolves rapidly and completely, with a more or less distinct yellowish color, accord- ing to the proportion of coloring matter originally present in the bile. The two biliary substances present in the watery solution may be separated from each other by the following means. On tlie addition of acetate of lead, the glyko-cholate of soda is decom- posed, and precipitates as a glyko-cholate of lead. The precipitate, 10 Glyko-cholate and Tauro-cholate of Soda, from Os-bile, after six days' crystalliza- tion. The glyko-cholate is crystallized ; the tauro- cliolate is in fluid drops. 146 THE BILE. separated bj filtration from the remaining fluid, is then decomposed in turn by carbonate of soda, and the original glyko-cholate of soda reproduced. The filtered fluid which remains, and which contains the tauro-cholate of soda, is then treated with subacetaie of lead, which precipitates a tauro-cholate of lead. This is separated by filtration, washed, and decomposed again by carbonate of soda, as in the former case. The two biliary substances in ox bile may, therefore, be dis- tinguished by their- reactions with the salts of lead. Both are precipitable by the subacetate; but the glyko-cholate of soda is precipitable also by the acetate, while the tauro-cholate is not so. If subacetate of lead, therefore, be added to the mixed watery solu- tion of the two substances, and the whole filtered, the subsequent addition of acetate of lead to the filtered fluid will produce no pre- cipitate, because both the biliary matters have been entirely thrown down with the deposit; but if the acetate of lead be first added, it will precipitate the glyko-cholate alone, and the tauro-cholate may afterward be thrown down separately by the subacetate. These two substances, examined separately, have been found to possess the following properties : — Glylw-cholate of soda (^diOfi^^^^O^^ crystallizes, when precipi- tated by ether from its alcoholic solution, in radiating bundles of fine white silky needles, as above described. It is composed of soda, united with a peculiar acid of organic origin, viz., glyko-cholic acid (C52li42-^0,i,HO). This acid is crystallizable and contains nitro- gen, as shown by the above formula, which is that given by Leh- man n. If boiled for a long time with a dilute solution of potass, glyko-cholic acid is decomposed with the production of two new substances; the first a non-nitrogenous acid body, cholic acid (C48H3g09,IIO) ; the second a nitrogenous neutral body, glycine (C4H5NOJ. Hence the name, glyko-cholic acid, given to the original substance, as if it were a combination of cholic acid with glycine. In reality, however, these two substances do not exist originally in the glyko-cholic acid, but are rather new combinations of its elements, produced by long boiling, in contact with potass and water. They are not, therefore, to be regarded as, in any way, natural ingredients of the bile, and do not throw any light on the real constitution of glyko-cholic acid. Tauro-cholate of soda (^diO^G^^^^^^O^^ is also a very abundant ingredient of the bile. It is said by Robin and YerdeiP that it is ' Chimie Anatomique et Physiologique, vol. ii. p. 473. THE BILE. 147 not crystallizable, owing probably to its not having been separated as yet in a perfectly pure condition. Lehmann states, on the con- trary, that it may crystallize,' when kept for a long time in contact with ether. We have not been able to obtain this substance, how- ever, in a crystalline form. Its acid constituent, tauro-cliolic acid^ is a nitrogenous body, like glyko-cholic acid, but differs from the latter by containing in addition two equivalents of sulphur. By long boiling in a dilute solution of potass, it is decomposed with the production of two other substances; the first of them the same acid body mentioned above as derived from the glyko-cholic, viz., cholic acid; and the second a new nitrogenous neutral body, viz., taurine {Q^^'^'^f)^. The same remark holds good with regard to these two bodies, that we have already made in respect to the sup- posed constituents of glyko-cholic acid. Neither cholic acid nor taurine can be properly regarded as really ingredients of tauro- cholic acid, but only as artificial products resulting from its altera- tion and decomposition. The glyko-cholates and tauro-cholates are formed, so far as we know, exclusively in the liver; since they have not been found in the blood, nor in any other part of the body, in healthy animals; nor even, in the experiments of Kunde, Moleschott, and Lehmann on frogs,^ after the entire extirpation of the liver, and consequent suppression of the bile. These substances are, therefore, produced in the glandular cells of the liver, by transformation of some other of their ingredients. They are then exuded in a soluble form, as part of the bile, and finally discharged by the excretory hepatic ducts. The two substances described above as the tauro-cholate and glyko-cholate of soda exist, properly speaking, only in the bile of the ox, where they were first discovered by Strecker. In examin- ing the biliary secretions of different species of animals, Strecker found so great a resemblance between them, that he was disposed to regard their ingredients as essentially the same. Having estab- lished the existence in ox bile of two peculiar substances, one crystallizable and non-sulphurous (glyko-cholate), the other uncrys- tallizable and sulphurous (tauro-cholate), he was led to consider the bile in' all species of animals as containing the same substances, and as differing only in the relative quantity in which the two ' Physiological Chemistry, Phil, ed., vol. i. p. 209. 2 Lehmann's Physiological Chemistry, Phil, ed., vol. i. p. 476. 148 THE BILE. Fig. 50. were present. The only exception to this was supposed to be pig's bile, in which Strecker found a peculiar organic acid, the " hyo- cbolic" or " hjo-cholinic" acid, in combination with soda as a base. The above conclusion of his, however, was not entirely correct. It is true that the bile of all animals, so far as examined, contains peculiar substances, which resemble each other in being freely soluble in water, soluble in absolute alcohol, and insoluble in ether; and in giving also a peculiar reaction with Pettenkofer's test, to be described presently. But, at the same time, these substances pre- sent certain minor differences in different animals, which show them not to be identical. In dog's bile, for example, there are, as in ox bile, two substances precipitable by ether from their alcoholic solution; one crystalliz- able, the other not so. But the former of these substances crystallizes much more readily than the glyko-cholate of soda from ox-bile. Dog's bile will not unfrequently begin to crystallize freely in five to six hours after precipitation by ether (Fig. 50); while in ox-bile it is usually twelve, and often twenty- four or even forty-eight hours before crystalliza- tion is fully established. But it is more particu- larly in their reaction with the salts of lead that the difference between these substances becomes manifest. For while the crystallizable substance of ox-bile is precipitated by acetate of lead, that of dog's bile is not affected by it. If dog's bile be evaporated to dryness, extracted with absolute alcohol, the alcoholic solution precipitated by ether, and the ether precipitate then dissolved in water, the addition of acetate of lead to the watery solution produces not the slightest tur- bidity. If subacetate of lead be then added in excess, a copious precipitate falls, composed of both the crystallizable and uncrystallizable sub- stances. If the lead precipitate be then separated by filtration, washed, and decomposed, as above described, by carbonate of soda, the watery solu- tion will contain the re-formed soda salts of the bile. The watery solution may then be evaporated to dryness, extracted with absolute alcohol, and the alcoholic solution precipi- tated by ether ; when the ether precipitate crystallizes partially after a time, as in fresh bile. Both the biliary matters of dog's bile Dog's Bile, extract- ed with absolute alcohol and precipitated with ether. THE BILE, 149 Fig. 51. are therefore precipitable by sabacetateof lead, but neither of them by the acetate. Instead of calling them, consequently, glykocholate and tauro-cholate of soda, we shall speak of them simply as the "crystalline" and "resinous" biliary substances. In cat's bile, the biliary substances act very much as in dog's bile. The ether- precipitate of the alcoholic solution contains here also a crystalline and a resinous substance; both of which are precipitable from their watery solution by subacetate of lead, but neither of them by the acetate. In pig's bile, on the other hand, there is no crystallizable sub- stance, but the ether-precipitate is altogether resinous in appearance. Notwithstanding this, its watery solution. precipitates abundantly by both the acetate and subacetate of lead. In human bile, again, there is no crystallizable substance. We have found that the dried bile, extracted with absolute alcohol, makes a clear, brandy red solution, which precipitates abundantly with ether in excess ; but the ether-precipitate, if allowed to stand, shows no sign of crystallization, even at the end of three weeks (Fig. 51). If the resinous pre- cipitate be separated by decantation and dissolved in water, it precipitates, as in the case of pig's bile, by both the acetate and subacetate of lead. This might, perhaps, be attributed to the pre- sence of two different substances, as in ox-bile, one precipitated by the acetate, the other by the subacetate of lead. Such, however, is not the case. For if the watery solution be precipitated by the acetate of lead and then filtered, the filtered fluid gives no precipitate afterward by the subacetate ; and if first precipitated by the sub- acetate, it gives no precipitate after filtration by the acetate. The entire biliary ingredients, therefore, of human bile are precipitated by both or either of the salts of lead. Dift'erent kinds of bile vary also in other re- spects; as, for example, their specific gravity, the depth and tinge of their color, the quantity of fat which they contain, &c. &c. We have already mentioned the variations in color and specific gravity. The alcoholic solution of dried ox-bile, furthermore, does not pre- cipitate at all on the addition of water ; while that of human bile, Human Bile, ex- tracted with absolute alcohol and precipitat- ed by ether. 150 THE BILE. of pig's bile, and of dog's bile precipitate abundantly with distilled water, owing to the quantity of fat which they hold in solution. These variations, however, are of secondary importance compared with those which we have already mentioned, and which show that the crystalline and resinous substances in different kinds of bile, though resembling each other in very many respects, are yet in reality far from being identical. TESTS FOE BILE. In investigating the physiology of any animal fluid it is, of course, of the first importance to have a convenient and reliable test by which its presence may be detected. For a long time the only test employed in the case of bile, was that which depended on a change of color produced hy oxidizing substances. If the bile, for example, or a mixture containing bile, be exposed in an open glass vessel for a few hours, the upper layers of the fluid, which are in contact with the atmosphere, gradually assume a greenish tinge, which becomes deeper with the length of time which elapses, and the quantity of bile existing in the fluid. Nitric acid, added to a mixture of bile and shaken up, produces a dense precipitate which takes a bright grass-green hue. Tincture of iodine produces the same change of color, when added in small quantity ; and pro- bably there are various other substances which would have the same effect. It is by this test that the bile has so often been recog- nized in the urine, serous effusions, the solid tissues, kc, in cases of jaundice. But it is very insufficient for anything like accurate investigation, since the appearances are produced simply by the action of an oxidizing agent on the coloring matter of the bile. A green color produced by nitric acid does not, therefore, indicate the presence of the biliary substances proper, but only of the biliver- dine. On the other hand, if the coloring matter be absent, the biliary substances themselves cannot be detected by it. For if the biliary substances of dog's bile be precipitated by ether from an alcoholic solution, dissolved in w^ater and decolorized by animal charcoal, the colorless watery solution will then give no green color on the addition of nitric acid or tincture of iodine, though it may precipitate abundantly by subacetate of lead, and give the other reactions of the crystalline and resinous biliary matters in a perfectly distinct manner. TESTS FOR BILE. 151 Petienkofer'' s Test. — This is undoubtedly the best test yet pro- posed for the detection of the biliary substances. It consists in mixing with a watery solution of the bile, or of the biliary sub- stances, a little cane sugar, and then adding sulphuric acid to the mixture until a red, lake or purple color is produced. A solution may be made of cane sugar, in the proportion of one part of sugar to four parts of water, and kept for use. One drop of this solution is mixed with the suspected fluid, and the sulphuric acid then imme- diately added. On first dropping in the sulphuric acid, a whitish precipitate falls, which is abundant in the case of ox-bile, less so in that of the dog. This precipitate redissolves in a slight excess of sulphuric acid, which should then continue to be added until the mixture assumes a somewhat syrupy consistency and an opalescent look, owing to the development of minute bubbles of air. A red color then begins to show itself at the bottom of the test-tube, and afterward spreads through the mixture, until the whole fluid is of a clear, bright, cherry red. This color gradually changes to a lake, and finally to a deep, rich, opaque purple. If three or four vol- umes of water be then added to the mixture, a copious precipitate falls down, and the color is destroyed. Various circumstances modify, to some extent, the rapidity and distinctness with which the above changes are produced. If the biliary substances be present in large quantity, and nearly pure, the red color shows itself at once, after adding an equal volume of sulphuric acid, and almost immediately passed into a strong purple. If they be scanty, on the other hand, the red color may not show itself for seven or eight minutes, nor the purple under twenty or twenty-five minutes. If foreign matters, again, not of a biliary nature, be also present, they are apt to be acted on by the sulphuric acid, and, by becoming discolored, interfere with the clearness and brilliancy of the tinges produced. On this account it is indispen- sable, in delicate examinations, to evaporate the suspected fluid to dryness, extract the dry residue with absolute alcohol, precipitate the alcoholic solution with ether, and dissolve the ether precipitate in water before applying the test. In this manner, all foreign sub- stances which might do harm will be eliminated, and the test will succeed without difficulty. It must not be forgotten, furthermore, that the sugar itself is liable to be acted on and discolored by sulphuric acid when added in excess, and may therefore by itself, give rise to confusion. A little care and practice, however, will enable the experimenter to avoid 152 THE BILE. all chance of deception from this source. AVhen sulphuric acid is mixed with a watery solution containing cane sugar, after it has been added in considerable excess, a yellowish color begins to show itself, owing to the commencing decomposition of the sugar. This color gradually deepens until it has become a dark, dingy, muddy brown ; but there is never at any time any clear red or purple color unless biliary matters be present. If the bile be present in but small quantity, the colors produced by it may be modified and obscured by the dingy yellow and brown of the sugar; but even this difficulty may be avoided by paying attention to the following precautions. In the first place, only very little sugar should be added to the suspected fluid. In the second place, the sulphuric acid should be added very gradually, and the mixture closely watched to detect the first changes of color. If bile be present, the red color peculiar to it is always produced before the yellowish tinge which indicates the decomposition of the sugar. When the biliary matters, therefore, are present in small quantity, the addi- tion of sulphuric acid should be stopped at that point, and the colors, though faint, will then remain clear, and give unmistakable evidence of the presence of bile. The red color alone is not sufficient as an indication of bile It is in fact only the commencement of the change which indicates the biliary matters. If these matters be present, the color passes, as we have already mentioned, first into a lake, then into a purple ; and it is this lake and purple color alone which can be regarded as really characteristic of the biliary reaction. It is important to observe that Pettenkofer's reaction is produced by the presence of either or both of the biliary substances proper ; and is not at all dependent on the coloring matter of the bile. For if the two biliary substances, crystalline and resinous, be extracted by the process above described, and, after being dissolved in water, decolorized with animal charcoal, the watery solution will still give Pettenkofer's reaction perfectly, though no coloring matter be pre- sent, and though no green tinge can be produced by the addition of nitric acid or tincture of iodine. If the two biliary substances be then separated from each other, and tested in distinct solutions, each solution will give the same reaction promptly and completely. Various objections have been urged against this test. It has been stated to be uncertain and variable in its action. Eobin and Verdeil' say that its reactions "do not belong exclusively to the » Op cit., vol. ii. p. 468. VARIATIONS AND FUNCTIONS OF BILE. 153 bile, and may therefore give rise to mistakes." Some fatty sub- stances and volatile oils (olein, oleic acid, oil of turpentine, oil of caraway) have been stated to produce similar red and violet colors, when treated with sugar and sulphuric acid. These objections, however, have not much, if any, practical weight. The test no doubt requires some care and practice in its application, as we have already pointed out ; but this is the case also, to a greater or less extent, with nearly all chemical tests, and particularly with those for sub- stances of organic origin. No other substance is, in point of fact, liable to be met with in the intestinal fluids or the blood, which would simulate the reactions of the biliary matters. We have found that the fatty matters of the chyle, taken from the thoracic duct, do not give any coloration which would be mistaken for that of the bile. When the volatile oils (caraway and turpentine) are acted on by sulphuric acid, a red color is produced which after- ward becomes brown and blackish, and a peculiar, tarry, empyreu- raatic odor is developed at the same time ; but we do not get the lake and purple colors spoken of above. Finally, if the precaution be observed — first of extracting the suspected matters with absolute alcohol, then precipitating with ether and dissolving the precipitate in water, no ambiguity could result from the presence of any of the above substances. Pettenkofer's test, then, if used with care, is extremely useful, and may lead to many valuable results. Indeed, no other test than this can be at all relied on to determine the presence or absence of the biliary substances proper. VARIATIONS AND FUNCTIONS OF BILE. With regard to the entire quantity of bile secreted daily, we have had no very positive knowledge, until the experiments of Bidder and Schmidt, published in 1852.* These experiments were per- formed on cats, dogs, sheep, and rabbits, in the following manner. The abdomen was opened, and a ligature placed upon the ductus communis choledochus, so as to prevent the bile finding its way into the intestine. An opening was then made in the fundus of the gall-bladder, by which the bile was discharged externally. The bile, so discharged, was received into previously weighed vessels, ' Verdaungssaefte und StoffwechseL Leipzig, 1852. 154: THE BILE. and its quantity accurately determined. Each observation usually occupied about two hours, during which period the temporary fluctuations occasionally observable in the quantity of bile dis- charged were mutually corrected, so fai; as the entire result was concerned. The animal was then killed, weighed and carefully examined, in order to make sure that the biliary duct had been securely tied, and that no inflammatory alteration had taken place in the abdominal organs. The observations were made at very different periods after the last meal, so as to determine the influence exerted by the digestive process upon the rapidity of the secretion. The average quantity of bile for twenty-four hours was then calcu- lated from a comparison of the above results ; and the quantity of its solid ingredients was also ascertained in each instance by eva- porating a portion of the bile in the water-bath, and weighing the dry residue. Bidder and Schmidt found in this way that the daily quantity of bile varied considerably in different species of animals. It was very much greater in the herbivorous animals used for experiment than in the carnivora. The results obtained by these observers are as follows .• — For every pound weight of the entire body there is secreted during 24 hours Feesh Bile. Dry Residue. In the cat 102 grains. 5.712 grains. "dog 140 " 6.916 " " sheep 178 " 9.408 " " rabbit 958 " 17.290 " Since, in the human subject, the digestive processes and the nutritive actions generally resemble those of the carnivora, rather than those of the herbivora, it is probable that the daily quantity of bile in man is very similar to that in the carnivorous animals. If we apply to the human subject the average results obtained by Bidder and Schmidt from the cat and dog, we find that, in an adult man, weighing 140 pounds, the daily quantity of the bile will be certainly not less than 16,940 grains, or very nearly 2|- pounds avoirdupois. It is a matter of great importance, in regard to the bile, as well as the other intestinal fluids, to ascertain whether it be a constant secretion, like the urine and perspiration, or whether it be intermit- tent, like the gastric juice, and discharged only during the digestive process. In order to determine this point, we have performed the VARIATIONS AND FUNCTIONS OF BILE. 155 following series of experiments on dogs. The animals were kept confined, and killed at various periods after feeding, sometimes by the inoculation of woorara, sometimes by hydrocyanic acid, but most frequently by section of the medulla oblongata. The contents of the intestine were then collected and examined. In all instances, the bile was also taken from the gall-bladder, and treated in the same way, for purposes of comparison. The intes- tinal contents always presented some peculiarities of appearance when treated with alcohol and ether, owing probably to the pre- sence of other substances than the bile ; but they always gave evidence of the presence of biliary matters as well. The biliary substances could almost always be recognized by the microscope in the ether-precipitate of the alcoholic solution ; the resinous substance, under the form of rounded, oily-looking drops (Fig. 52), and the other, under the form of crystalline groups, generally presenting the appearance of double bundles of slender, radiat- ing, slightly curved or wavy, needle-shaped crys- tals. These substances, dis- solved in water, gave a pur- ple color with sugar and sulphuric acid. These ex- periments were tried after the animals had been kept for one, two, three, five, six, seven, eight, and twelve days "without food. The result showed that, in all these instances, bile was present in the small intes- tine. It is, therefore, plainly not an intermittent secre- tion, -nor one which is con- cerned exclusively in the digestive process ; but its secretion is con- stant, and it continues to be discharged into the intestine for many days after the animal has been deprived of food. Crystalline and Eesinohs Biliary Sub- STANOEs; from Small Intestine of Dog, after two days' fasting. The next point of importance to be examined relates to the time after feeding at which the hile passes into the intestine in the greatest abundance. Bidder and Schmidt have already investigated this point in the following manner. They operated, as above described, 156 THE BILE. by tying the common bile-duct, and tlien opening the fundus of the gall-bladder, so as to produce a biliary fistula, by which the whole of the bile was drawn off. By doing this operation, and collecting and weighing the fluid discharged at different periods, they came to the conclusion that the flow of bile begins to increase within two and a half hours after the introduction of food into the stomach, but that it does not reach its maximum of activity till the end of twelve or fifteen hours. Other observers, however, have obtained different results. Arnold,' for example, found the quantity to be largest soon after meals, decreasing again after the fourth hour. Kolliker and Miiller,^- again, found it largest between the sixth and eighth hours. Bidder and Schmidt's experiments, indeed, strictly speaking, show only the time at which the bile is most actively secreted by the liver, but* not when it is actually discharged into the intestine. Our own experiments, bearing on this point, were performed on dogs, by making a permanent ^^^' ^^' duodenal fistulse, on the same plan that gastric fistulee have so often been established for the examination of the gastric juice. (Fig. 53.) An incision was made through the abdominal walls, a short distance to the right of the median line, the floating portion of the duodenum drawn up toward the external wound, opened by a longitudinal inci- sion, and a silver tube, armed at each end with a narrow projecting collar or flange, in- serted into it by one extremity, five and a half inches below the pylorus, and two and a half inches below the orifice of the lower pancreatic duct. The other extremity of the tube was left projecting from the external opening in the abdominal pa- rietes, the parts secured by sutures, and the wound allowed to heal. Duodenal Fistula. — a. Stomach. 6. Duo- denum, c, c, c. Pancreas ; its two ducts are seen opening into the duodenum, one near the orifice of the biliary duct, d, the other a short distance lower down. e. Silver tube passing through the abdominal walls and opening into the duodenum. ' In Am. Journ. Med. Sci., April, 1856. Ibid., April, 1857. VARIATIONS AND FUNCTIONS OF BILE. 157 After cicatrization was complete, and the animal had entirely recovered his healthy condition and appetite, the intestinal fluids were drawn oflf at various intervals after feeding, and their contents examined. This operation, which is rather more difficult than that of making a permanent gastric fistula, is nevertheless exceedingly useful when it succeeds, since it enables us to study, not only the time and rate of the biliary discharge, but also, as mentioned in a previous chapter (Chap. VI.), many other extremely interesting matters connected with intestinal digestion. In order to ascertain the absolute quantity of bile discharged into the intestine, and its variations during digestion, the duodenal fluids were drawn off, for fifteen minutes at a time, at various periods after feeding, collected, weighed, and examined separately, as follows: each separate quantity was evaporated to dryness, its dry residue extracted with absolute alcohol, the alcoholic solution precipitated with ether, and the ether-precipitate, regarded as repre- senting the amount of biliary matters present, dried, weighed, and then treated with Pettenkofer's test, in order to determine, as nearly as possible, their degree of purity or admixture. The result of these experiments is given in the following table. At the eigh- teenth hour so small a quantity of fluid was obtained, that the amount of its biliary ingredients was not ascertained. It reacted perfectly, however, with Pettenkofer's test, showing that bile was really present. Time after Quautity of fluid Dry residue Quantity of Proportion of feeding. iu l.i minutes. of same. biliary matters. biliary matters to dry residue. Immediately 640 grains 33 grains 10 grains .30 1 hour 1,990 " 105 " 4 " .03 3 hours 780 " 60 " 4 " .07 6 " 750 " 73 " H " .05 9 " 860 " 78 " 4i " .06 12 " 325 " 23 " 3f " .16 15 " 347 " 18 " 4 " .22 18 " . 21 « 384 " 11 " 1 '• .09 24 " 163 " H " 3i " .34 25 " 151 " 5 " 3 " .60 From this it appears that the bile passes into the intestine in by far the largest quantity immediately after feeding, and within the first hour. After that time its discharge remains pretty constant ; not varying much from four grains of solid biliary matters every fifteen minutes, or sixteen grains per hour. The animal used for the above observations weighed thirty-six and a half pounds. 158 THE BILE. The next point to be ascertained with regard to this question is the following, viz : What hecomes of the bile in its passage through the intestine? Our experiments, performed with a view of settling this point, were tried on dogs. The animals were fed with fresh meat, and then killed at various intervals after the meals, the abdo- men opened, ligatures placed upon the intestine at various points, and the contents of its upper, middle, and lower portions collected and examined separately. The results thus obtained show that, under ordinary circumstances, the bile, which is quite abundant in the duodenum and upper part of the small intestine, diminishes in quantity from above downward, and is not to be found in the large intestine. The entire quantity of the intestinal contents also dimi- nishes, and their consistency increases, as we approach the ileo- caecal valve; and at the same time their color changes from a light yellow to a dark bronze or blackish-green, which is always strongly pronounced in the last quarter of the small intestine. The contents of the small and large intestine were furthermore evaporated to dryness, extracted with absolute alcohol, and the alcoholic solutions precipitated with ether ; the quantity of ether precipitate being regarded as representing approximatively that of the biliary substances proper. The result showed that the quantity of this ether precipitate is, both positively and relatively, very much less in the large intestine than in the small. Its proportion to the entire solid contents, is only one-fifth or one-sixth as great in the large intestine as it is in the small. But even this inconsiderable quantity, found in the contents of the large intestine, does not con- sist of biliary matters ; for the watery solutions being treated with sugar and sulphuric acid, those from both the upper and lower portions of the small intestine always gave Pettenkofer's reaction promptly and perfectly in less than a minute and a half; while in that from the large intestine no red or purple color was produced, even at the end of three hours. The small intestine consequently contains, at all times, substances giving all the reactions of the biliary ingredients; while in the contents of the large intestine no such substances can be recognized by Pettenkofer's test. The biliary matters, therefore, disappear in their passage through the intestine. In endeavoring to ascertain what is the precise /M?-/c^?■o?^ of the Ule in the intestine, our first object must be to determine what part, if VAKIATIONS AND FUNCTIONS OF BILE. 159 any, it takes in tlie digestive process. As the liver is situated, like the salivary glands and the pancreas, in the immediate vicinity of the alimentary canal, and like them, discharges its secretion into the cavity of the intestine, it seems at first natural to regard the bile as one of the digestive fluids. We have previously shown, however, that the digestion of all the different elements of the food is provided for by other secretions ; and furthermore, if we examine experimentally the digestive power of bile on alimentary substances, we obtain only a negative result. Bile exerts no special action upon either albuminoid, starchy, or oleaginous matters, when mixed with them in test-tubes and kept at the temperature of 100° F. It has therefore, apparently, no direct influence in the digestion of these substances. Furthermore it appears, from the experiments detailed above, that the secretion of the bile and its discharge into the intestine are not confined to the periods of digestion, but take place con- stantly, and continue even after the animal has been kept for many days without food. These facts would lead us to regard the bile as simply an excrementitious fluid ; containing only ingredients resulting from the waste and disintegration of the animal tissues, and not intended to perform any particular function, digestive or otherwise, but merely to be eliminated from the blood, and discharged from the system. The same view is more or less supported, also, by the following facts, viz: — 1st. The bile is produced, unlike all the other animal secretions, from venous blood ; that is, the blood of the portal vein, which has already become contaminated by circulation through the abdominal organs, and may be supposed to contain disorganized and effete ingredients ; and 2d. Its complete suppression produces, in the human subject, symptoms of poisoning of the nervous system, analogous to those which follow the suppression of the urine, or the stoppage of respi- ration, and the patient dies, usually in a comatose condition, at the end of ten or twelve days. The above circumstances, taken together, would combine to make it appear that the bile is simply an excrementitious fluid, not necessary or useful as a secretion, but only destined, like the urine, to be eliminated and discharged. Nevertheless experiment has shown that such is not the case ; and that, in point of fact, it is necessary for the life of the animal, not only that the bile be secreted and discharged, but furthermore that it be discharged into the 160 THE JBILE. intestine, and pass through the tract of the alimentary canal. The most satisfactory experiments of this kind are those of Bidder and Schmidt,^ in which they tied the common biliary duct in dogs, and then established a permanent fistula in the fundus of the gall-bladder through which the bile was allowed to flow by a free external orifice. In this manner the bile was effectually excluded from the. intestine, but at the same time was freely and wholly discharged from the body, by the artificial fistula. If the bile therefore were simply an excrementitious fluid, its deleterious ingredients being all eliminated as usual, the animals would not suffer any serious injury from this operation. If, on the contrary, they were found to suffer or die in consequence of it, it would show that the bile has really some im- portant function to perform in the intestinal canal, and is not simply excrementitious in its nature. The result showed that the effects of such an experiment were fatal to the animal. Four dogs only survived the immediate effects of the operation, and were afterward frequently used for purposes of experiment. One of them was an animal from which the spleen had been previously removed, and whose appetite, as usual after this operation, was morbidly ravenous; his sj^stem, accordingly, being placed under such unnatural conditions as to make him an unfit subject for further experiment. In the second animal that survived, the communication of the biliary duct with the intestine became re-established after eighteen days, and the experiment con- sequently had no result. In the remaining two animals, however, everything was successful. The fistula in the gall-bladder became permanently established ; and the bile-duct, as was proved subse- quently by post-mortem examination, remained completely closed, so that no bile found its way into the intestine. Both these ani- mals died; one of them at the end of twenty-seven days, the other at the end of thirty-six days. In both, the symptoms were nearly the same, viz., constant and progressive emaciation, which proceeded to such a degree that nearly every trace of fat disappeared from the body. The loss of flesh amounted, in one case to more than two- fifths, and in the other to nearly one-half the entire weight of the animal. There was also a falling off of the hair, and an unusually disagreeable, putrescent odor in the feces and in the breath. Not- withstanding this, the appetite, remained good. Digestion was not essentially interfered with, and none of the food was discharged ' Op. cit., p. 103. VAEIATIONS AND FUNCTIONS OF BILE. 161 with the feces ; but there was much rumbling and gurgling in the intestines, and abundant discharge of flatus, more strongly marked in one instance than in the other. There was no pain ; and death took place, at last, without any violent symptoms, but by a simple and gradual failure of the vital powers. How is it, then, that although the bile be not an active agent in digestion, its presence in the alimentary canal is still essential to life ? What office does it perform there, and how is it finally dis- posed of? We have already shown that the bile disappears in its passage through the intestine. This disappearance may be explained in two different ways. First, the biliary matters may be actually re- absorbed from the intestine, and taken up by the bloodvessels ; or secondly, they may be so altered and decomposed by the intestinal fluids as to lose the power of giving Pettenkofer's reaction with sugar and sulphuric acid, and so pass off with the feces in an insoluble form. Bidder and Schmidt' have finally determined this point in a satisfactory manner ; and have demonstrated that the biliary substances are actually reabsorbed, by showing that the quantity of sulphur present in the feces is far inferior to that contained in the biliary ingredients as they are discharged into the intestine. These observers collected and analyzed all the feces passed, dur- ing five days, by a healthy'dog, weighing 17.7 pounds. The entire fecal mass during this period weighed 1508.15 grains, ( Water 874.20 grains. Containing I g^^.^^,^^.^^^ g33_g5 , 1508.15 The solid residue was composed as follows: — Neutral fat, soluble in ether . . 43.710 grains. Fat, witli traces of biliary matter . 77.035 " Alcohol extract with biliary matter 58.900 containing 1.085 grs. of sulphur. Substances not of a biliary nature extracted by muriatic acid and hot alcohol .... 148.800 containing 1.302 grs. of sulphur. 2.387 Fatty acids with oxide of iron . 98.425 Residue consisting of hair, sand, &c., 207.080 633.950 ' Op. cit., p. 217. 11 162 THE BILE. Now, as it lias already been shown that the dog secretes, during 24 hours, 6.916 grains of solid biliary matter for every pound weight of the whole body, the entire quantity of biliary matter secreted in five days by the above animal, weighing 17.7 pounds, must have been 612.5 grains, or nearly as much as the whole weight of the dried feces. But furthermore, the natural proportion of sulphur in dog's bile (derived from the uncrystallizable biliary matter), is six per cent, of the dry residue. The 612.5 grains of dry bile, secreted during five days, contained therefore 86.75 grains of sulphur. But the entire quantity of sulphur, existing in any form in the feces, was 5.952 grains ; and of this only 2.387 grains were derived from substances which could have been the products of biliary matters — the remainder being derived from the hairs which are always contained in abundance in the feces of the dog. That is, not more than one-fifteenth part of the sulphur, originally present in the bile, could be detected in the feces. As this is a simple chemical element, not decomposable by any known means, it must, accordingly, have been reabsorbed from the intestine. We have endeavored to complete the evidence thus furnished by Bidder and Schmidt, and to demonstrate directly the reabsorption of the biliary matters, by searching for them in the ingredients of the portal blood. AVe have examined, for this purpose, the portal blood of dogs, killed at various periods after feeding. The animals were killed by section of the medulla oblongata, a ligature imme- diately placed on the portal vein, while the circulation was still active, and the requisite quantity of blood collected by opening the vein. The blood was sometimes immediately evaporated to dryness by the water bath. Sometimes it was coagulated by boil- ing in a porcelain capsule, over a spirit lamp, with water and an excess of sulphate of soda, and the filtered watery solution after- ward examined. But most frequently the blood, after being col- lected from the vein, was coagulated by the gradual addition of three times its volume of alcohol at ninety-five per cent., stirring the mixture constantly, so as to make the coagulation gradual and uniform. It was then filtered, the moist mass remaining on the filter subjected to strong pressure in a linen bag, by a porcelain press, and the fluid thus obtained added to that previously filtered. The entire spirituous solution was then evaporated to dryness, the dry residue extracted with absolute alcohol, and the alcoholic solution treated as usual, with ether, &c., to discover the presence of biliary matters. In every instance, blood was taken at the same time from VARIATIONS AND FUNCTIONS OF BILE. 163 the jugular, or tlie abdominal vena cava, and treated in the same way for purposes of comparison. "We have examined the blood, in this way, one, four, six, nine, eleven and a half, twelve, and twenty hours after feeding. As the result of these examinations, we have found that in the venous blood, both of the portal vein and of the general circulation, there exists a substance soluble in water and absolute alcohol, and precipitable by ether from its alcoholic solution. This substance is often consi- derably more abundant in the portal blood than in that taken from the general venous system. It adheres closely to the sides of the glass after precipitation, so that it is always difficult, and often im- possible, to obtain enough of it, mixed with ether, for microscopic examination. It dissolves, also, like the biliary substances, with great readiness in water ; but in no instance have we ever been able to obtain from it such a satisfactory reaction with Pettenkofer's test, as would indicate the presence of bile. This is not because the reaction is masked, as might be suspected, by some of the other ingredients of the blood ; for if at the same time, two drops of bile be added to half an ounce of blood taken from the abdominal vena cava, and the two specimens treated alike, the ether precipitate may be considerably more abundant in the case of the portal blood ; and yet that from the blood of the vena cava, dissolved in water, will give Pettenkofer's reaction for bile perfectly, while that of the por- tal blood will give no such reaction. Notwithstanding, then, the irresistible evidence afforded by the experiments of Bidder and Schmidt, that the biliary matters are really taken up by the portal blood, we have failed to recognize them there by Pettenkofer's test. They must accordingly undergo certain alterations in the intestine, previously to their absorption, so that they no longer give the ordinary reaction of the biliary sub- stances. We cannot say, at present, precisely what these alterations are ; but they are evidently transformations of a catalytic nature, produced by the contact of the bile with the intestinal juices. The bile, therefore, is a secretion which has not yet accomplished its function when it is discharged from the liver and poured into the intestine. On the contrary, during its passage through the intestine it is still in the interior of the body, in contact with glandular sur- faces, and mingled with various organic substances, the ingredients of the intestinal fluids, which act upon it as catalytic bodies, and produce in it new transformations. This may account for the fact stated above, that the bile, though a constant and uninterrupted 164 THE BILE. secretion, is nevertheless poured into the intestine in the greatest abundance immediately after a hearty meal. This is not because it is to take any direct part in the digestion of the food ; but because the intestinal fluids, being themselves present at that time in the greatest abundance, can then act upon and decompose the greatest quantity of bile. At all events, the biliary ingredients, after being altered and transformed in the intestine, as they might be in the interior of a glandular organ, re-enter the blood under some new form, and are carried away by the circulation, to complete their function in some other part of the body. FOKMATION OF SUGAR IN THE LIVER. 165 CHAPTER IX. FORMATION OF SUGAR IN THE LIVER. Beside the secretion of bile, the liver performs also another exceedingly important function, viz., the -produclion of sugar by a metamorphosis of some of its organic ingredients. Under ordinary circumstances a considerable quantity of sac- charine matter is introduced with the food, or produced, from starchy substances, by the digestive process in the intestinal canal. In man and the herbivorous animals, accordingly, an abundant * supply of sugar is derived from these sources ; and, as we have already shown, the sugar thus introduced is necessary for the proper support of the vital functions. For though the saccharine matter absorbed from the intestine is destroyed by decomposition soon after entering the circulation, yet the chemical changes by which its decomposition is eifected are themselves necessary for the proper constitution of the blood, and the healthy nutrition of the tissues. Experiment shows, however, that the system does not depend, for • its supply of sugar, entirely upon external sources ; but that sac- charine matter is also produced independently, in the tissue of the liver, whatever may be the nature of the food upon which the animal subsists. This important function was first discovered by M. Claude Bernard' in 1848, and described by him under the name of the glycogenic function of the liver. It has long been known that sugar may be abundantly secreted, under some circumstances, when no vegetable matters have been taken with the food. The milk, for example, of all animals, car- nivorous as Avell as herbivorous, contains a notable proportion of sugar ; and the quantity thus secreted, during lactation, is in some instances very great. In the human subject, also, when suffering from diabetes, the amount of saccharine matter discharged with the ' Nouvelle Fonction du Foie. Paris, 1853. 166 FORMATION OF SUGAR IN THE LIVER. urine has often appeared to be altogether out of proportion to that which could be accounted for by the vegetable substances taken as food. The experiments of Bernard, the most important of which we have repeatedly confirmed, in common with other investigators, show that in these instances most of the sugar has an internal origin, and that it first makes its appearance in the tissue of the liver. If a carnivorous animal, as, for example, a dog or a cat, be fed for several days exclusively upon meat, and then killed, the liver alone of all the internal organs is found to contain sugar among its other ingredients. For this purpose, a portion of the organ should be cut into small pieces, reduced to a pulp by grinding in a mortar with a little water, and the mixture coagulated by boiling with an excess of sulphate of soda, in order to precipitate the albuminous and coloring matters. The filtered fluid will then reduce the oxide of copper, with great readiness, on the application of Trommer's test. A decoction of the same tissue, mixed with a little yeast, will also give rise to fermentation, producing alcohol and carbonic acid, as is usual with saccharine solutions. On the contrary, the tissues of the spleen, the kidneys, the lungs, the muscles, &c,, treated in the same way, give no indication of sugar, and do not reduce the salts of copper. Every otlier organ in the body may be entirely desti- tute of sugar, but the liver always contains it in considerable quan- tity, provided the animal be healthy. Even the blood of the portal vein, examined by a similar process, contains no saccharine element, and yet the tissue of the organ supplied by it shows an abundance of saccharine ingredients. It is remarkable for how long a time the liver will continue to exhibit the presence of sugar, after all external supplies of this substance have been cut off. Bernard kept two dogs under his own observation, one for a period of three, the other of eight months,^ during which period they were confined strictly to a diet of animal food (boiled calves' heads and tripe), and then killed. Upon ex- amination, the liver was found, in each instance, to contain a proportion of sugar fully equal to that present in the organ under ordinary circumstances. The sugar, therefore, which is found in the liver after death, is a normal ingredient of the hepatic tissue. It is not formed in other parts of the body, nor absorbed from the intestinal canal, but takes ' Nouvelle Fonction du Foie, p. 50. FORMATION OF SUGAR IN THE LIVER. 167 its origin in the liver itself; it is produced, as a new formation, by a secreting process in the tissue of the organ. The presence of sugar in the liver is common to all species of animals, so far as is yet known. Bernard found it invariably in monkeys, dogs, cats, rabbits, the horse, the ox, the goat, the sheep, in birds, in reptiles, and in most kinds of fish. It was only in two species of fish, viz., the eel and the ray (Murasna angnilla and Raia batis), that he sometimes failed to discover it; but the failure in these instances was apparently owing to the commencing putres- cence of the tissue, by which the sugar had probably been destroyed. In the fresh liver of the human subject, examined after death from accidental violence, sugar was found to be present in the proportion of 1.10 to 2.14 per cent, of the entire weight of the organ. The following list shows the average percentage of sugar present in the healthy liver of man and different species of animals, accord- ing to the examinations of Bernard : — Pekcentage of Sugar in the Liver. « In man .... 1.68 In ox . . . . 2.30 " monkey " dog . " cat " rabbit " sheep 2.15 " horse .... 4.08 1.69 " goat .... 3.89 1.94 " birds .... 1.49 1.94 " reptiles . . . 1.04 2.00 " fish . . . . 1.45 With regard to the nature and properties of the liver sugar, it resembles very closely glucose, or the sugar of starch, the sugar of honey, and the sugar of milk, though it is not absolutely identical with either one of them. Its solution reduces, as we have seen, the salts of copper in Trommer's test, and becomes colored brown when boiled with caustic potass. It ferments very readily, also, when mixed witb yeast and kept at the temperature of 70° to 100° F. It is distinguished from all the other sugars, according to Bernard,^ by the readiness with which it becomes decomposed in the blood — since cane sugar and beet root sugar, if injected into the circulation of a living animal, pass through the system without sensible decom- position, and are discharged unchanged with the urine ; sugar of milk and glucose, if injected in moderate quantity, are decomposed in the blood, but if introduced in greater abundance make their appearance also in the urine ; while a solution of liver sugar, though injected in much larger quantity than either of the others, may dis- ' Lemons de Physiologic Experimentale. Paris, 1855, p. 213. 168 rORMATIOX OF SUGAR IN THE LIVER. appear altogether in the circulation, without passing off' by the kidneys. This substance is therefore a sugar of animal origin, similar in its properties to other varieties of saccharine matter, derived from different sources. The sugar of the liver is not produced in the blood by a direct decomposition of the elements of the circulating fluid in the vessels of the organ, but takes its origin in the solid substance of the hepatic tissue^ as a natural ingredient of its organic texture. The blood which may be pressed out from a liver recently extracted from the body, it is true, contains sugar ; but this sugar it has absorbed from the tissues of the organ in which it circulates. This is demonstrated by the singular fact that the fresh liver of a recently killed animal, though it may be entirely drained of blood and of the sugar which it contained at the moment of death, will still continue for a certain time to produce a saccharine substance. If such a liver be injected with water by the portal vein, and all the blood contained in its vessels washed out by the stream, the water which escapes by the hepatic vein will still be found to contain sugar. M. Bernard has found' that if all the sugar contained in a fresh liver be extracted in this manner by a prolonged watery injection, so that neither the water which escapes by the hepatic vein, nor the substance of the liver itself, contain any further traces of sugar, and if the organ be then laid aside for twenty-four hours, both the tissue of the liver and the fluid which exudes from it will be found at the end of that time to have again become highly saccharine. The sugar, therefore, is evidently not produced in the blood circulating through the liver, but in the substance of the organ itself. Once having originated in the hepatic tissue, it is absorbed thence by the blood, and trans- ported by the circulation, as we shall hereafter show, to other parts of the body. The sugar which thus originates in the tissue of the liver, is pro- duced by a mutual decomposition and transformation of various other ingredients of the hepatic substance ; these chemical changes being a part of the nutritive processes by which the tissue of the organ is constantly sustained and nourished. There is probably a series of several different transformations which take place in this manner, the details of which are not yet known to us. It has been discovered, however, that one change at least precedes the final ' Gazette Hebdomadaire, Paris, Oct. 5, 1855. FORMATION OF SUGAR IN THE LIVER. 169 production of saccharine matter ; and that the sugar itself is pro- duced by the transformation of another peculiar substance, of ante- rior formation. This substance, which precedes the formation of sugar, and which is itself produced in the tissue of the liver, is known by the name of the glycogenic matter^ or glycogene. This glj^cogenic matter may be extracted from the liver in the following manner. The organ is taken immediately from the body of the recently killed animal, cut into small pieces, and coagulated by being placed for a few minutes in boiling water. This is in order to prevent the albuminous liquids of the organ from acting upon the glj'cogenic matter and decomposing it at a medium temperature. The coagulated tissue is then drained, placed in a mortar, reduced to a pulp by bruising and grinding, and afterward boiled in dis- tilled water for a quarter of an hour or more, by which the glyco- genic matter is extracted and held in solution by the boiling water. The liquid of decoction, which should be as concentrated as pos- sible, must then be expressed, strained, and filtered, after which it appears as a strongly opalescent fluid, of a slightly yellowish tinge. The glycogenic matter which is held in solution may be precipi- tated by the addition to the filtered fluid of five times its volume of alcohol. The precipitate, after being repeatedly washed with alcohol in order to remove sugar and biliary matters, may then be redissolved in distilled water. It may be precipitated from its watery solution either by alcohol in excess or by crystallizable acetic acid, in both of which it is entirely insoluble, and mav be afterward kept in the dry state for an indefinite time without losing its properties. The glycogenic matter, obtained in this way, is regarded as intermediate in its nature and properties between hydrated starch and dextrine. Its ultimate composition, according to M. Pelouze," is as follows : — When brought into contact with iodine, it produces a coloration varying from violet to a deep, clear, maroon red. It does not reduce the salts of copper in Troramer's test, nor does it ferment when placed in contact with yeast at the proper temperature. It does not, therefore, of itself contain sugar. It may easily be con- verted into sugar, however, by contact with any of the animal ferments, as, for example, those contained in the saliva or in the ' Journal de Physiologie, Paris, 1858, p. 552. 170 FORMATION OF SUGAR IN" THE LIVER. blood. If a solution of glycogenic matter be mixed witb fresli human saliva, and kept for a few minutes at the temperature of 100° F., the mixture will then be found to have acquired the power of reducing the salts of copper and of entering into fermentation by contact with yeast. The glycogenic matter has therefore been converted into sugar by a process of catalysis, in the same manner as vegetable starch would be transformed under similar conditions. The glycogenic matter which is thus destined to be converted into sugar, is formed in the liver by the processes of nutrition. It may be extracted, as we have seen above, from the hepatic tissue of carnivorous animals, and is equally present when they have been exclusively confined for many days to a meat diet. It is not in- troduced with the food ; for the fleshy meat of the herbivora does not contain it in appreciable quantity, though these animals so constantly take starchy substances with their food. In them, the starchy matters are transformed into sugar by digestion, and the sugar so produced is rapidly destroyed after entering the circula- tion ; so that usually neither saccharine nor starchy substances are to be discovered in the muscular tissue. M. Poggiale' found that in very many experiments, performed by a commission of the French Academy for the purpose of examining this subject, glyco- genic matter was detected in ordinary butcher's meat only once. We have also found it to be absent from the fresh meat of the bullock's heart, when examined in the manner described above, l^evertheless, in dogs fed exclusively upon this food for eight days, glycogenic matter may be found in abundance in the liver, while it does not exist in other parts of the body, as the spleen, kidney, lungs, &c. Furthermore, in a dog fed exclusively for eight days upon the fresh meat of the bullock's heart, and then killed four hours after a meal of the same food, at which time intestinal absorption is going on in full vigor, the liver contains, as above mentioned, both glycogenic matter and sugar; but neither sugar nor glycogenic mat- ter can be found in the blood of the portal vein, when subjected to a similar examination. The glycogenic matter, accordingly, does not originate from any external source, but is formed in the tissue of the liver ; where it is soon afterward transformed into, sugar, while still forming a part of the substance of the organ. ' Journal de Pbysiologie, Paris, 1858, p. 558. FOEMATION OF SUGAE IN THE LIVER. 171 The formation of sugar in the liver is therefore a function com- posed of two distinct and successive processes, viz : first, the forma- tion, in the hepatic tissue, of a glycogenic matter, having some resemblance to dextrine ; and secondly, the conversion of this gly- cogenic matter into sugar, by a process of catalysis and transforma- tion. The sugar thus produced in the substance of the liver is absorbed from it by the blood circulating in its vessels. The mechanism of this absorption is probably the same with that which goes on in other parts of the circulation. It is a process of transudation and endosmosis, by which the blood in the vessels takes up the saccha- rine fluids of the liver, during its passage through the organ. While the blood of the portal vein, .therefore, in an animal fed exclusively upon meat, contains no sugar, the blood of the hepatic vein, as it passes upward to the heart, is always rich in saccharine ingredients. This difference can easily be demonstrated by exa- mining comparatively the two kinds of blood, portal and heptic, from the recently killed animal. The blood in its passage through the liver is found to have acquired a new ingredient, and shows, upon examination, all the properties of a saccharine liquid. The sugar produced in the liver is accordingly to be regarded as a true secretion, formed by the glandular tissue of the organ, by a similar process to that of other glandular secretions. It differs from the latter, not in tbe manner of its production, but only in the mode of its discharge. For while the biliary matters produced in the liver are absorbed by the hepatic ducts and conducted down- ward to the gall-bladder and the intestine, the sugar is absorbed by the bloodvessels of the organ and carried upward, by the hepatic veins, toward the heart and the general circulation. The production of sugar in the liver during health is a constant process, continuing, in many cases, for several days after the animal has been altogether deprived of food. Its activity, howev&r, like that of most other secretions, is subject to periodical augmentation and diminution. Under ordinary circumstances, the sugar, which is absorbed by the blood from the tissue of the liver, disappears very soon after entering the circulation. As the bile is transformed in the intestine, so the sugar is decomposed in the blood. We are not yet acquainted, however, with the precise nature of the changes which it undergoes after entering the vascular system. It is very probable, according to the views of Lehmann and Robin, that it is at first converted into lactic acid (CgHgOg), which decomposes in 172 FORMATION OF SUGAR IN THE LIVER. turn the alkaline carbonates, setting free carbonic acid, and form ing lactates of soda and potass. But whatever be the exact mode of its transformation, it is certain that the sugar disappears rapidly; and while it exists in considerable quantity in the liver and in the blood of the hepatic veins and the right side of the heart, it is not usually to be found in the pulmonary veins nor in the blood of the general circulation. About two and a half or three hours, however, after the inges- tion of food, according to the investigations of Bernard, the circu- lation of blood through the portal system and the liver becomes considerably accelerated. A larger quantity of sugar is then pro- duced in the liver and carried away from the organ by the hepatic veins ; so that a portion of it then escapes decomposition while passing through the lungs, and begins to appear in the blood of the arterial system. Soon afterward it appears also in the blood of the capillaries ; and from four to six hours after the commence- ment of digestion it is produced in the liver so much more rapidly than it is destroyed in the blood, that the surplus quantity circulates throughout the body, and the blood everywhere has a slightly saccharine character. It does not, however, in the healthy condi- tion, make its appearance in any of the secretions. After the sixth hour, this unusual activity of the sugar producing function begins again to diminish ; and the transformation of the sugar in the circulation going on as before, it gradually disappears as an ingredient of the blood. Finally, the ordinary equilibrium between its production and its decomposition is re-established, and it can no longer be found except in the liver and in that part of the circulatory system which is between the liver and the lungs. There is, therefore, a periodical increase in the amount of unde- composed sugar in the blood, as we have already shown to be the case with the fatty matter absorbed during digestion ; but this in- crease is soon followed by a corresponding diminution, and during the greater portion of the time its decomposition keeps pace with its production, and it is consequently prevented from appearing in the blood of the general circulation. There are produced, accordingly, in the liver, two different secre- tions, viz., bile and sugar. Both of them originate by transforma- tion of the ingredients of the hepatic tissue, from which they are absorbed by two different sets of vessels. The bile is taken up by the biliary ducts, and by them discharged into the intestine; while the sugar is carried off by the hepatic veins, to be decomposed in the circulation, and become subservient to the nutrition of the blood. THE SPLEEN. 173 CHAPTER X. THE SPLEEN. The spleen is an exceedingly vascular organ, situated in the vicinity of the great pouch of the stomach and supplied abund- antly by branches of the coeliac axis. Its veins, like those of the digestive abdominal organs, form a part of the great portal system, and conduct the blood which has passed through it to the liver, before it mingles again with the general current of the circulation. The spleen is covered on its exterior by an investing membrane or capsule, which forms a protective sac, containing the soft pulp of Avhich the greater part of the organ is composed. This capsule, in the spleen of the ox, is thick, whitish and opaque, and is com- posed to a great extent of yellow elastic tissue. It accordingly possesses, in a high degree, the physical property of elasticity, and may be widely stretched without laceration ; returning readily to its original size as soon as the extending force is relaxed. In the carnivorous animals, on the other hand, the capsule of the spleen is thinner, and more colorless and transparent. It con- tains here but very little elastic tissue, being composed mostly of smooth, involuntary muscular fibres, connected in layers by a little intervening areolar tissue. In the herbivorous animals, accordingly, the capsule of the spleen is simply elastic, while in the carnivora it is contractile. In both instances, however, the elastic and contractile properties of the capsule subserve a nearly similar purpose. There is every reason to believe that the spleen is subject to occasional and per- haps regular variations in size, owing to the varying condition of the abdominal circulation. Dr. William Dobson^ found that the size of the organ increased, from the third hour after feeding up to the fifth ; when it arrived at its maximum, gradually, decreasing after that period. When these periodical congestions take place, ' In Gray, on tlie Structure and Uses of the Spleen. London, 1854, p. 40. 174. THE SPLEEN. the organ becoming turgid with blood, the capsule is distended ; and limits, by its resisting power, the degree of tumefaction to which the spleen is liable. When the disturbing cause has again passed away, and the circulation is about to return to its ordinary condition, the elasticity of the capsule in the herbivora and its con- tractility in the carnivora, compress the soft vascular tissue within, and reduce the organ to its original dimensions. This contractile action of the investing capsule can be readily seen in the dog or the cat, by opening the abdomen while digestion is going on, ex- posing the spleen and removing it, after ligature of its vessels. When first exposed, the organ is plump and rounded, and presents externally a smooth and shining surface. But as soon as it has been removed from the abdomen and its vessels divided, it begins to contract sensibly, becomes reduced in size, stiffj and resisting to the touch ; while its surface, at the same time, becomes uniformly wrinkled, by the contraction of its muscular fibres. In its interior, the substance of the spleen is traversed everywhere by slender and ribbon-like cords of fibrous tissue, which radiate from the sheath of its principal arterial trunks, and are finally attached to the internal surface of its investing capsule. These fibrous cords, or irabeculce, as they are called, by their frequent branching and mutual interlacement, form a kind of skeleton or framework by which the soft splenic pulp is embraced, and the shape and integrity of the organ maintained. They are composed of similar elements to those of the investing capsule, viz., elastic tissue and involuntary muscular fibres, united with each other by a varying quantity of the fibres of areolar tissue. The interstices between the trabeculae of the spleen are occupied by the splenic pulp; a soft, reddish substance, which contains, beside a few nerves and lymphatics, capillary bloodvessels in great profusion, and certain whitish globular bodies, which may be re- garded as the distinguishing anatomical elements of the organ, and which are termed the Malpighian bodies of the spleen. The Malpighian bodies are very abundant, and are scattered throughout the splenic pulp, being most frequently attached to the sides, or at the point of bifurcation of some small artery. They are readily visible to the naked eye in the spleen of the ox, upon a fresh section^of the organ, as minute, whitish, rounded bodies, which may be separated, by careful manipulation, from the surrounding parts. In the carnivorous animals, on the other hand, and in the human subject, it is more difficult to distinguish them by the un- THE SPLEEN. 176 aided eye, though they always exist in the spleen in a healthy condition. Their average diameter, according to Kcilliker, is y^\ of an inch. They consist of a closed sac, or capsule, containing in its interior a viscid, semi-solid mass of cells, cell-nuclei, and homo- geneous substance. Each Malpighian body is covered, on its exte- rior, by a network of fine capillary bloodvessels; and it is now perfectly well settled, by the observations of various anatomists (Kcilliker, Busk, Huxley, &c.), that bloodvessels also penetrate into the substance of the Malpighian body, and there form an internal capillary plexus. The spleen is accordingly a glandular organ, analogous in its minute structure to the solitary and agminated glands of the small intestine, and to the lymphatic glands throughout the body. Like them, it is a gland without an excretory duct ; and resembles also, in this respect, the thyroid and thymus glands and the supra-renal capsules. All these organs have a structure which is evidently glandular in its nature, and yet the name of glands has been some- times refused to them because they have, as above mentioned, no duct, and produce apparently no distinct secretion. We have already seen, however, that a secretion may be produced in the interior of a glandular organ, like the sugar in the substance of the liver, and yet not be discharged by its excretory duct. The veins of the gland, in this instance, perform the part of excretory ducts. They absorb the new materials, and convey them, through the medium of the blood, to other parts of the body, where they suffer subsequent alterations, and are finally decomposed in the circula- tion. The action of such organs is consequently to modify the consti- tution of the blood. As the blood passes through their tissue, it absorbs from the glandular substance certain materials which it did not previously contain, and which are necessary to the perfect con- stitution of the circulating fluid. The blood, as it passes out from the organ, has therefore a different composition from that which it possessed before its entrance; and on this account the name of vascular glands has been applied to all the glandular organs above mentioned, which are destitute of excretory ducts, and is eminently applicable to the spleen. The precise alteration, however, which is effected in the blood during its passage through the splenic tissue, has not yet been discovered. Various hypotheses have been advanced from time to time, as to the processes which go on in this organ ; many of them 176 THE SPLEEIST. vague and indefinite in character, and some of them directly con- tradictory of each other. None, however, have yet been offered which are entirely satisfactory in themselves, or which rest on suffi- ciently reliable evidence. A very remarkable fact with regard to the spleen is that it may be entirely removed, in many of the lower animals, without its loss producing any serious permanent injury. This experiment has been frequently performed by various observers, and we have our- selves repeated it several times with similar results. The organ may be easily removed, in the dog or the cat, by drawing it out of the abdomen, through an opening in the median line, placing a few ligatures upon the vessels of the gastro-splenic omentum, and then dividing the vessels between the ligatures and the spleen. The wound usually heals without difficulty ; and if the animal be killed some weeks afterward, the only remaining trace of the operation is an adhesion of the omentum to the inner surface of the abdominal parietes, at the situation of the original wound. The most constant and permanent effect of a removal of the spleen is an unusual increase of the appetite. This symptom we have observed in some instances to be excessively developed ; so that the animal would at all times throw himself, with an unnatural avidity, upon any kind of food offered him. We have seen a dog, subjected to this operation, afterward feed without hesitation upon the flesh of other dogs; and even devour greedily the entrails, taken warm from the abdomen of the recently killed animal. The food taken in this unusual quantity is, however, perfectly well digested ; and the animal will often gain very perceptibly in weight. In one instance, a cat, in whom the unnatural appetite was marked though not excessive, increased in weight from five to six pounds, in the course of a little less than two months; and at the same time the fur became sleek and glossy, and there was a considerable improvement in the general appearance of the animal. Another symptom, which usually follows removal of the spleen, is an unnatural ferocity of disposition. The animal will frequently attack others, of its own or a different species, without any appa- rent cause, and without any regard to the difference of size, strength, &c. This symptom is sometimes equally excessive with that of an unnatural appetite ; while in other instances it shows itself only in occasional outbursts of irritability and violence. Neither of the symptoms, however, which we have just de- scribed, appear to exert any permanently injurious effect upon the THE SPLEEN". 177 animal Avhich has been subjected to the operation; and life may be prolonged for an indefinite period, without any serious disturbance of the nutritive process, after the spleen has been completely extirpated. We must accordingly regard the spleen, not as a single organ, but as associated with others, which may completely, or to a great extent, perform its functions after its entire removal. We have already noticed the similarity in structure between the spleen and the mesenteric and lymphatic glands ; a similarity which has led some writers to regard them as more or less closely associated with each other in function, and to consider the spleen as an unusually developed lymphatic or mesenteric gland. It is true that this organ is provided with a comparatively scanty supply of lymphatic vessels ; and the chyle, which is absorbed from the intestine, does not pass through the spleen, as it passes through the remaining mesenteric glands. Still, the physiological action of the spleen may correspond with that of the other lymphatic glands, so far as regards its influence on the blood; and there can be little doubt that its function is shared, either by them or by some other glan- dular organs, which become unnaturally active, and more or less perfectly supply its place after its complete removal. 12 178 THE BLOOD. CHAPTER XI. THE BLOOD. The blood, as it exists in its natural condition, while circulating in the vessels, is a thick opaque fluid, varying in color in different parts of the body from a brilliant scarlet to a dark purple. It has a slightly alkaline reaction, and a specific gravity of 1055. It is not, however, an entirely homogeneous fluid, but is found on microscopic examination to consist, first, of a nearly colorless, transparent, alkaline fluid, termed the plasma, containing water, fibrin, albumen, salts, &c., in a state of mutual solution; and, secondly, of a large number of distinct cells, or corpuscles, the blood- globules, swimming freely in the liquid plasma. These glo- bules, which are so small as not to be distinguished by the naked eye, by being mixed thus abundantly with the fluid plasma, give to the entire mass of the blood an opaque appearance and a uni- form red color. BLOOD-GLOBULES. On microscopic examination it is found that the globules of the blood are of two kinds, viz., red and white ; of these the red are by far the most abundant. The o'ed globules of the blood present, under the microscope, a perfectly circular outline and a smooth exterior. (B'ig. 54.) Their size varies somewhat, in human blood, even in the same specimen. The greater number of them have a transverse diameter of 3500 ^^ an inch; but there are many smaller ones to be seen, which are not more than gJ^o or even 4o'oo of ^^ ii^^h in diameter. Their form is that of a spheroid, very much flattened on its opposite surfaces, somewhat like a round biscuit, or a thick piece of money with rounded edges. The blood-globule accordingly, when seen flatwise, presents a comparatively broad surface and a circular out- BLOOD-GLOBULES. 179 Human Blood-globules. — a. Red globules, seeu fiatwi-e. h. Red globules, seen edgewise, e. White globule. line (a); but if it be made to roll over, it will present itself edge- wise during its rotation and assume the flattened form indicated at h. The thickness of the globule, seen in this position, is about y 15^55 of an inch, or a little less than one-fifth of its transverse diameter. When the globules are exa- mined lying upon their broad surfaces, it can be seen that these surfaces are not exactly flat, but that there is on each side a slight central depres- sion, so that the rounded edges of the blood-globule are evidently thicker than its middle portion. This ine- quality produces a remark- able optical effect. The sub- stance of which the blood-globule is composed refracts light more strongly than the fluid plasma. Therefore, when examined with the microscope, by transmitted light, the thick edges of the globules act as double convex lenses, and concentrate the light above the level of the fluid. Conse- quently, if the object-glass be carried upward by the ad- justing screw of the micro- scope, and lifted away from tbe stage, so that the blood- globules fall beyond its fo- cus, their edges will appear brighter. But the central por- tion of each globule, being excavated on both sides, acts as a double concave lens, and disperses the light from a point below the level of the fluid. It, therefore, grows brighter as the object-glass is carried downward, and the object falls within its focus. Red Globules op the Blood, beyond the focus of the microscope. seen a little An alternating appearance of the 180 THE BLOOD. blood-globules may, therefore, be produced by viewing tliem first beyond and then within the focus of the instrument. When be- yond the focus, the globules will be seen with a bright rim and a darlv centre. (Fig. 55.) When within it, they will appear with a dark rim and a bright centre. (Fig. 56.) The blood-globules accord- ingly have the form of a thickened disk with rounded edges and a double central excavation. They have, con- sequently, been sometimes called "blood-disks," instead of blood-globules. The term " disk," however, does not in- dicate their exact shape, any more than the other ; and the term " blood-corpuscle," which is also sometimes used, does not indicate it at all. And although the term " blood-globule" may not be precisely a correct one, still it is the most convenient ; and need not give rise to any confusion, if we remember the real shape of the bodies designated by it. This term will, consequently, be employed whenever we have occasion to speak of the blood-globules in the following pages. Within a minute after being The same, seea a little withiu the focus. Fig. 57. Blood-globules of coin. adhering together, like rolls placed under the microscope, the blood-globules, after a fluctuating movement of short duration, very often arrano;e themselves in slip;ht- ly curved rows or chains, in which they adhere to each other by their flat surfaces, presenting an appearance which has been aptly com- pared with that of rolls of coin. This is probably owing merely to the coagulation of the blood, which takes place very rapidly when it is spread BLOOD-GLOBULES. 181 out in thin layers and in contact with glass surfaces; and which, by compressing the globules, forces them into such a position that they may occupy the least possible space. This position is evi- dently that in which they are applied to each other by their flat surfaces, as above described. The color of the blood-globules, when viewed by transmitted light and spread out in a thin layer, is a light amber or pale yellow. It is, on the contrary, deep red when they are seen by reflected light, or piled together in comparatively thick layers. When viewed singly, they are so transparent that the outlines of those lying under- neath can be easily seen, showing through the substance of the superjacent globules. Their consistency is peculiar. They are not solid bodies, as they have been sometimes inadvertently described ; but on the contrary have a consistency which is very nearly fluid. They are in consequence exceedingly flexible, and easily elongated, bent, or otherwise distorted by accidental pressure, or in passing through the narrow currents of fluid which often establish them- selves accidentally in a drop of blood under microscopic examina- tion. This distortion, however, is only temporary, and the globules regain their original shape, as soon as the accidental pressure is taken off. The peculiar flexibility and elasticity thus noticed are characteristic of the red globules of the blood, and may always serve to distinguish them from any other free cells which may be found in the animal tissues or fluids. In structure the blood-globules are homogeneous. They have been sometimes erroneously described as consisting of a closed vesicle or cell-wall, containing in its cavity some fluid or serai-fluid substance of a different character from that composing the wall of the vesicle itself. No such structure, however, is really to be seen in them. Each blood-globule consists of a mass of organized ani- mal substance, perfectly or nearly homogeneous in appearance, and of the same color, consistency and composition throughout. In some of the lower animals (birds, reptiles, fish) it contains also a granular nucleus, imbedded in the substance of the globule ; but in no instance is there any distinction to be made out between an external cell-wall and an internal cavity. The appearance of the blood-globules is altered by the addition of various foreign substances. If water be added, so as to dilute the plasma, the globules absorb it by imbibition, swell, lose their double central concavity and become paler. If a larger quantity of water be added, they finally dissolve and disappear altogether. 182 THE BLOOD. B L D - G L O B U L E S , water. swollen by the imbibitiou of When a moderate quantity of water is mixed with the blood, the edges of the globules, being thicker than the central portions, and absorbing water more ^ig- 5S- abundantly, become turgid, and encroach gradually upon the central part. (Fig. 58.) It is very common to see the central depression, under these circumstances, disap- pear on one side before it is lost on the other, so that the globule, as it swells up, curls over towards one side, and assumes a peculiar cup- shaped form (a). This form may often be seen in blood- globules that have been soaking for some time in the urine, or in any other animal fluid of a less density than the plasma of the blood. Dilute acetic acid dissolves the blood-globules more promptly than water, and solutions of the caustic alkalies more promptly still. If a drop of blood be allowed partially to evaporate while under the microscope, the globules Fig- 59. near the edges of the prepa- ration often diminish in size, and at the same time present a shrunken and crenated ap- pearance, as if minute gran- ules were projecting from their surfaces (Fig. 59); an effect apparently produced by the evaporation of part of their watery ingredients. For some unexplained rea- son, however, a similar dis- tortion is often produced in some of the globules by the addition of certain other ani- mal fluids, as for example the saliva; and a few can even be seen in this condition after the addition of pure water. Bloob-globdles crenated. shrunken, with their margins BLOOD-GLOBULES. 183 The entire mass of the blood-globules, in proportion to the rest of the circulating fluid, can only be approximately measured by the eve in a microscopic examination. In ordinary analyses the globules are usually estimated as amounting to about fifteen per cent., by weight, of the entire blood. This estimate, however, refers, properly speaking, not to the globules themselves, but only to their dry residue, after the water which they contain has been lost by evaporation. It is easily seen, by examination with the microscope, that the globules, in their natural semi-fluid condition, are really much more abundant than this, and constitute fully one-half the entire mass of the blood; that is, the intercellular fluid, or plasma, is not more abundant than the globules themselves which are sus- pended in it. When separated from the other ingredients of the blood and examined by themselves, the globules are found, ac- cording to Lehmann, to present the following composition : — CoMPosiTiox OF THE Blocd-Globules IN 1000 Parts. Water 688.00 Globnline ........... 282.22 Hsematine 16.75 Fatty substances ......... 2.31 Undetermined (extractive) matters . . . . . . 2.60 Chloride of sodium ....... " potassium ...... Phosphates of soda and potass ..... Sulphates " " ...... Phosphate of lime ....... " magnesia . . . . . . 1- 8.12 1000.00 The most important of these ingredients is the glohuline. This is an organic substance, nearly fluid in its natural condition by union with water, and constituting the greater part of the mass of the blood-globules. It is soluble in water, but insoluble in the plasma of the blood, owing to the presence in that fluid of albumen and saline matters. If the blood be largely diluted, however, the globuline is dissolved, as already mentioned, and the blood-globules are destroyed. Globuline coagulates by heat; but, according to Eobin and Yerdeil, only becomes opalescent at 160°, and requires for its complete coagulation a temperature of 200° F. The hcematme is the coloring matter of the globules. It is, like globuline, an organic substance, but is present in much smaller quan- tity than the latter. It is not contained in the form of a powder, mechanically deposited in the globuline, but the two substances are 184 THE BLOOD. intimately mingled throughout the mass of the blood-globule, just as the fibrin and albumen are mingled in the plasma. Heematine contains, like the other coloring matters, a small proportion of iron. This iron has been supposed to exist under the form of an oxide ; and to contribute directly in this way to the red color of the sub- stance in question. But it is now ascertained that although the iron is found in an oxidized form in the ashes of the blood-globules after they have been destroyed by heat, its oxidation probably takes place during the process of incineration. So far as we know, there- fore, the iron exists originally in the h^ematine as an ultimate element, directly combined with the other ingredients of this sub- stance, in the same manner as the carbon, the hydrogen, or the nitrogen. The blood-globules of all the warm blooded quadrupeds, with the exception of the family of the caraelidse, resemble those of the human species in shape and structure. They differ, however, some- what in size, being usually rather smaller than in man. There are but two species in which they are known to be larger than in man, viz., the Indian elephant, in which they are 27^00 of ^^^ inch, and the two-toed sloth {Bradypus didactylus), in which they are ogVo of an inch in diameter. In the musk deer of Java they are smaller than in any other known species, measuring rather less than yg^oo- of an inch. The following is a list showing the size of the red globules of the blood in the principal mammalian species, taken from the measurements of Mr. Gulliver,' Diameter of Red Globules in the Ape . • jj'do of an i nch. Cat . • 4iV7irOf an inch. Horse iHUff Fox . iiVff Ox . 4 A (J Wolf . 5sW Sheep ssr'oij Elephant . J705 Goat . B^'oij Red deer T(JO-ff_ Dog . FsVff Musk deer . • TSoUS In all these instances the form and general appearance of the globules are the same. The only exception to this rule among the mammalians is in the family of the camelidas (camel, dromedary, lama), in which the globules present an oval outline instead of a circular one. In other respects they resemble the foregoing. In the three remaining classes of vertebrate animals, viz., birds, reptiles and fish, the blood-globules differ so much from the above that they can be readily distinguished by microscopic examination. In works of William Hewson, Sydenham edition, London, 1846, p. 327. BLOOD-GLOBULES. 185 Blood-globules of Frog. seen edgewise, h. White globule. -a. Blood-globule They are oval in form, and contain a colorless granular nucleus imbedded in their substance. They are also considerably larger than the blood-globules of the mammalians, particularly in the class of reptiles. In the frog (Fig. 60) they measure joVu of an inch in their long diameter; and in Menohran- chiis, the great water lizard of the northern lakes, ^^^ of an inch. In Proteus angui- nus they attain the size, ac- cording to Dr. Carpenter,' of gi^y of an inch. Beside the corpuscles de- scribed above, there are glo- bules of another kind found in the blood, viz., the lokite globules. These globules are very much less numerous than the red ; the proportion between the two, in human blood, being one white to two or three hundred red globules. In reptiles, the relative quantity of the w^hite globules is greater, but they are always considerably less abundant than the red. They differ also from the latter in shape, size, color and consistency. They are globular in form, white or colorless, and instead of being homogeneous like the others, their substance is filled everywhere with minute dark molecules which give them a finely granular appearance. (Fig. 54, c.) In size they are considerably larger than the red globules, being about 05V0 of an inch in diameter. They are also more consistent than the others, and do not so easily glide along in the minute currents of a drop of blood under examination, but adhere readily to the surfaces of the glass. If treated with dilute acetic acid, they swell up and become smooth and circular in outline ; and at the same time a separation or partial coagulation seems to take place in the substance of which they are composed, so that an irregular collection of granular matter shows itself in their interior, becoming more divided and broken up as the action of the acetic acid upon the globule is longer continued. (Fig. 61.) This collection of granular matter The Microscope and its Revelations, Philadelphia edition, p. 600. 186 THE BLOOD. often assumes a curved or crescentic form, as at «, and sometimes various other irregular shapes. It does not indicate the existence of a nucleus in the white Fig. 61. Whtte Globhles of the Blood; altered by dilute acetic acid- Ill globule, but is merely an appearance produced by the coagulating and disintegrat- ing action of acetic acid upon the substance of which it is composed. The chemical constitution of the white globules, as distinguished from the red, has never been determined ; owing to the small quantity in which they occur, and the difficulty of separating them from the others for purposes of analysis. The two kinds of blood- globules, white and red, are to be regarded as distinct and inde- pendent anatomical forms. It has been sometimes supposed that the white globules were converted, by a gradual transformation, into the red. There is, however, no direct evidence of this ; as the transformation has never been seen to take place, either in the human subject or in the mammalia, nor even its intermediate stages satisfactorily observed. When, therefore, in default of any such direct evidence, we are reduced to the surmise which has been adopted by some authors, viz., that the change " takes place too rapidly to be detected by our means of observation,"^ it must be acknowledged that the above opinion has no solid founda- tion. It has been stated by some authors (Kcilliker, Gerlach) that in the blood of the batrachian reptiles there are to be seen certain bodies intermediate in appearance between the white and the red globules, and which represent different stages of transi- tion from one form to the other; but this is not a fact which is generally acknowledged. "We have repeatedly examined, with reference to this point, the fresh blood of the frog, as well as that of the menobranchus, in which the large size of the globules would give every opportunity for detecting any such changes, did ' Kolliker, Handbuch der Gewebelelire, Leipzig, 1852, p. 582. BLOOD-GLOBULES. — PLASMA. 187 they really exist; and it is our unavoidable conclusion from these observations, that there is no good evidence, even in the blood of reptiles, of any sucli transformation taking place. There is simply, as in human blood, a certain variation in size and opacit}'- among the red globules; but no such connection with, or resemblance to, the white globules as to indicate a passage from one form to the other. The red and white globules are therefore to be regarded as distinct and independent anatomical elements. They are mingled together in the blood just as capillary bloodvessels and nerves are mino-led in areolar tissue; but there is no other connection between them, so far as their formation is concerned, than that of juxtapo- sition. Neither is it at all probable that the red globules are produced or destroyed in any particular part of the body. One ground for the belief that these bodies were produced by a metamorphosis of the white globules was a supposition that they were continually and rapidly destroyed somewhere in the circulation ; and as this loss must be as rapidly counterbalanced by the formation of new glo- bules, and as no other probable source of their reproduction ap- peared, they were supposed to be produced by transformation of the white globules. But there is no reason for believing that the red globules of the blood are any less permanent, as anatomical forms, than tbe muscular fibres or the nervous filaments. They undergo, it is true, like all the constituent parts of the body, a constant interstitial metamorphosis. They absorb incessantly nu- tritious materials from the blood, and give up to the circulating fluid, at the same time, other substances which result from their internal waste and disintegration. But they do not, so far as we know, perish bodily in any part of the circulation. It is not the anatomical forms^ anywhere, which undergo destruction and reno- vation in the nutritive process; but only the proximate principles of tvhich they are composed. The effect of this interstitial nutrition, therefore, in the blood-globules as in the various solid tissues, is merely to maintain them in a natural and healthy condition of integrity. PLASMA. The plasma of the blood, according to Lehmann, has the follow- ing constitution: — 188 THE BLOOD. Composition of the Plasma in 1000 Parts. Water •. . . . 902.90 Fibrin 4.05 Albumen 78.84 Fatty matters 1.72 Undetermined (extractive) matters . . . . . . 3.94 Chloride of sodium ........ " potassium ....... Phosphates of soda and potass ...... Sulphates " " Phosphate of lime ........ " magnesia ....... 8.55 1000.00 The above ingredients are all intimately mingled in the blood- plasma, in a fluid form, by mutual solution; but they may be separated from each other for examination by appropriate means. The two ingredients belonging to the class of organic substances are the fibrin and the albumen. The fibrin^ though present in small quantity, is evidently an important element in the constitution of the blood. It may be ob- tained in a tolerably pure form by gently stirring freshly drawn blood with a glass rod or a bundle of twigs; upon which the fibrin coagulates, and adheres to the twigs in the form of slender threads and flakes. The fibrin, thus coagulated, is at first colored red by the haematine of the blood globules entangled in it ; but it may be washed colorless by a few hours' soaking in running water. The fibrin then pre- Fig. 62. Coagulated Fibrin, sliowing its fibrillated con- dition. sents itself under the form of nearly white threads and flakes, having a semi-solid consistency, and a consider- able degree of elasticity. The coagulation of fibrin takes place in a peculiar manner. It does not solidify in a perfectly homogeneous mass; but if examined by the microscope in thin layers it is seen to have a fibroid or filamentous texture. In this condition it is said to be " fibrillated." (Fig. 62.) The PLASMA. 189 filaments of which it is composed are colorless and elastic, and when isolated are seen to be exceedingly minute, being not more than 4o^^^Tj or 6ven ^^-J-^xr of ^^ i^ich in diameter. They are in part arranged so as to lie parallel with each other; but are more generally interlaced in a kind of irregular network, crossing each other in every direction. On the addition of dilute acetic acid, they swell up and fuse together into a homogeneous mass, but do not dis- solve. They are often interspersed everywhere with minute granu- lar molecules, which render their outlines more or less obscure. Once coagulated, fibrin is insoluble in water and can only be again liquefied by the action of an alkaline or strongly saline solution, or by prolonged boiling at a very high temperature. These agents, however, produce a complete alteration in the proper- ties of the fibrin, and after being subjected to them it is no longer the same substance as before. The quantity of fibrin in the blood varies in different parts of the body. According to the observations of various writers,' there is more fibrin generally in arterial than in venous blood. The blood of the veins near the heart, again, contains a smaller proportion of fibrin than those at a distance. The blood of the portal vein con- tains less than that of the jugular ; and that of the hepatic vein less than that of the portal. The albumen is undoubtedly the most important ingredient of the plasma, judging both from its nature and the abundance in which it occurs. It coagulates at once on being heated to 160° P., or by contact with alcohol, the mineral acids, the metallic salts, or with ferrocyanide of potassium in an acidulated solution. It exists naturally in the plasma in a fluid form by reason of its union with water. The greater part of the water of the plasma, in fact, is in union with the albumen ; and when the albumen coagulates, the water remains united with it, and assumes at the same time the solid form. If the plasma of the blood, therefore, after the removal of the fibrin, be exposed to the temperature of 160° F., it solidifies almost completely ; so that only a few drops of water remain that can be drained away from the coagulated mass. The phosphates of lime and magnesia are also held in solution principally by the albumen, and are retained by it in coagulation. The fatty matters exist in the blood mostly in a saponified form, excepting soon after the digestion of food rich in fat. At that period, as we have already mentioned, the emulsioned fat finds its way ' Robin and Verdeil, op. cit., vol. ii. p. 202. 190 THE BLOOD. into the blood, and circulates for a time unchanged. Afterward it disappears as free fat, and remains partly in the saponified condition. The saline ingredients of the plasma are of the same nature with those existing in the globules. The chlorides of sodium and potassium, and the phosphates of soda and potass are the most abundant in both, while the sulphates are present only in minute quantity. The proportions in which the various salts are present are very different, according to Lehmann,^ in the blood-globules and in the plasma. Chloride of potassium is most abundant in the globules, chloride of sodium in the plasma. The phos- phates of soda and potass are more abundant in the globules than in the plasma. On the other hand, the phosphates of lime and magnesia are more abundant in the plasma than in the globules. The substances known under the name of extractive matters consist of a mixture of different ingredients, belonging mostly to the class of organic substances, which have not yet been separated in a state of sufficient purity to admit of their being thoroughly examined and distinguished from each other. They do not exist in great abundance, but are -undoubtedly of considerable importance in the constitution of the blood. Beside the substances enumerated in the above list, there are still others which occur in small quantity as ingredients of the blood. Among the most important are the alkaline carbonates, which are held in solution in the serum. It has ■already been mentioned that while the phosphates are most abundant in the blood of the carnivora, the carbonates are most abundant in that of the herbivora. Thus Lehmann^ found car- bonate of soda in the blood of the ox in the proportion of 1.628 per thousand parts. There are also to be found, in solution in the blood, urea, urate of soda, creatine, creatinine, sugar, &c.; all of them ci-ystallizable substances derived from the transformation of other ingredients of the blood, or of the tissues through which it circu- lates. The relative quantity, however, of these substances is very minute, and has not yet been determined with precision. COAGULATION OF THE BLOOD. A few moments after the blood has been withdrawn from the vessels, a remarkable phenomenon presents itself, viz., its coagula- tion or clotting. This process commences at nearly the same time ' Op. cit . voL i. p. 546. ' Op. cit., vol. i. p. 393. COAGULATION OF THE BLOOD. 191 throughout the whole mass of the blood. The blood becomes first somewhat diminished in fluidity, so that it will not run over the edge of the vessel, when slightly inclined ; and its surface may be gently depressed with the end of the finger or a glass rod. It then becomes rapidly thicker, and at last solidifies into a uniformly red, opaque, consistent, gelatinous mass, which takes the form of the vessel in which the blood was received. Its coagulation is then complete. The process usually commences, in the human subject, in about fifteen minutes after the blood has been drawn, and is completed in about twenty minutes. The coagulation of the blood is dependent entirely upon the presence of the fibrin. This fact has been demonstrated in various ways. In the first place, if frog's blood be filtered, so as to separate the globules and leave them upon the filter, while the plasma is allowed to run through, the colorless filtered fluid which contains the fibrin soon coagulates ; while no coagulation takes place in the moist globules remaining on the filter. Again, if the freshly drawn blood be stirred with a bundle of rods, as we have already de- scribed above, the fibrin coagulates upon them by itself, while the rest of the plasma, mixed with the globules, remains perfectly fluid. It is the fibrin, therefore, which, by its own coagulation, induces the solidification of the entire blood. As the fibrin is uniformly distributed throughout the blood, when its coagulation takes place the minute filaments which make their appearance in it entangle in their meshes the globules and the albuminous fluids of the plasma. A very small quantity of fibrin, therefore, is sufficient to entangle by its coagulation all the fluid and semi-fluid parts of the blood, and convert the whole into a volumi- nous, trembling, jelly-like mass, which is Fig. 63. known by the name of the " crassamentum" or " clot." (Fig. 63.) As soon as the clot has fairly formed, it begins to contract and diminish in size. Ex- actly how this contraction of the clot is pro- duced, we are unable to say ; but it is proba- bly a continuation of the same process by which its solidification is at first accomplished, b,^i „f ^^^^^^^^ p^^^^. or at least one very similar to it. As the lated blood, showing the . ,• 1 ,1 11 • n ■ ^ whole mass uniformly solidi- contraction proceeds, the albuminous fluids ged. begin to be pressed out from the meshes in which they were entangled. A few isolated drops first appear on 192 THE BLOOD. Fig. 64. the surface of the clot. These drops soon increase in size and be- come more numerous. They run together and coalesce with each other, as more and more fluid exudes from the coagulated mass, until the whole surface of the clot is covered with a thin layer of fluid. The clot at first adheres pretty strongly to the sides of the vessel into which the blood was drawn ; but as its contraction goes on, its edges are separated, and the fluid continues to exude between it and the sides of the vessel. This exudation continues for ten or twelve hours; the clot growing constantly smaller and firmer, and the expressed fluid more and more abundant. The globules, owing to their greater con- sistency, do not escape with the albuminous fluids, but remain entangled in the fibrinous coagulum. Finally, at the end of ten or twelve hours the whole of the blood has usu- ally separated into two parts, viz., the clot, which is a red, opaque, dense and resisting, semi-solid mass, consisting of the fibrin and the blood-globules; and the serum, which is a transparent, nearly colorless fluid, containing the water, albumen, and saline matters of the plasma. (Fig. 64.) The change of the blood in coagulation may therefore be ex- pressed as follows : — Bowl of Coagulated Blood, after twelve hours ; showing the clot contracted and floating iu the fluid serum. Before coagulation the blood consists of 1st. Globules ; and 2d. Plasma — containing After coagulation it is separated into 1st. Clot, containing i Fibrin and I Globules ; Fibrin, Albumen, Water, Salts. ( Albumen, and 2d. Serum, containing -^ Water, i Salts. The coagulation of the blood is hastened or retarded by various physical conditions, which have been studied with care by various observers, but more particularly by Kobin and Yerdeil. The con- ditions which influence the rapidity of coagulation are as follows : First, the rapidity with which the blood is drawn from the vein, and the size of the orifice from which it flows. If blood be drawn rapidly, in a full stream, from a large orifice, it remains fluid for a comparatively long time ; if it be drawn slowly, from a narrow COAGULATION OF THE BLOOD. 193 orifice, it coagulates quickly. Tlius it usually happens that in the operation of venesection, the blood drawn immediately after the opening of the vein runs freely and coagulates slowly; while that which is drawn toward the end of the operation, when the tension of the veins has been relieved and the blood trickles slowly from the wound, coagulates quickly. Secondly, the shape of the vessel into which the blood is received and the condition of its internal surface. The greater the extent of surface over which the blood comes in contact with the vessel, the more is its coagulation hastened. Thus, if the blood be allowed to flow into a tall, narrow, cylindrical vessel, or into a shallow plate, it coagulates more rapidly than if it be received into a hemispherical bowl, in which the ex- tent of surface is less, in proportion to the entire quantity of blood which it contains. For the same reason, coagulation takes place more rapidly in a vessel with a roughened internal surface, than in one which is smooth and polished. The blood coagulates most rapidly when spread out in thin layers, and entangled among the fibres of cloth or sponges. For the same reason, also, hemorrhage continues longer from an incised wound than from a lacerated one; because the blood, in flowing over the ragged edges of the lacer- ated bloodvessels and tissues, solidifies upon them readily, and thus blocks up the wound. In all these cases, there is an inverse relation between the rapidity of coagulation and the firmness of the clot. When coagulation takes place slowly, the clot afterward becomes small and dense, and the serum is abundant. When coagulation is rapid, there is but little contraction of the coagulum, an imperfect separation of the serum, and the clot remains large, soft and gelatinous. It is well known to practical physicians that a similar relation exists when the coagulation of the blood is hastened or retarded by disease. In cases of -lingering and exhausting illness, or in diseases of a typhoid or exanthematous character, with much depression of the vital powers, the blood coagulates rapidly and the clot remains soft. In cases of active inflammatory disease, as pleurisy or pneumonia, occurring in previously healthy subjects, the blood coagulates slowly, and the clot becomes very firm. In every instance, the blood which has coagulated liquefies again at the commencement of putrefaction. The coagulation of the fibrin is not a commencement of organization. The filaments already described, which show themselves in the clot (Fig. 62), are not, properly speaking, organized fibres, and are en- 13 194 THE BLOOD. tirely different in their character from the fibres of areolar tissue, or any other normal anatomical elements. They are simply the ultimate form which fibrin assumes in coagulating, just as albumen takes the form of granules under the same circumstances. The coagulation of fibrin does not differ in character from that of any other organic substance ; it merely difi'ers in the physical conditions which give rise to it. All the coagulable organic substances are naturally fluid, and coagulate only when they are placed under certain unusual con- ditions. But the particular conditions necessary for coagulation vary with the different organic substances. Thus albumen coagu- lates by the application of heat. Casein, which is not affected by heat, coagulates by contact with an acid body. Pancreatine, again, is coagulated by contact with sulphate of magnesia, which has no efieet on albumen. So fibrin, which is naturally fluid, and which remains fluid so long as it is circulating in the vessels, coagulates when it is withdrawn from them and brought in contact with unnatural surfaces. Its coagulation, therefore, is no more " sponta- neous," properly speaking, than that of any other organic substance. Still less does it indicate anything like organization, or even a commencement of it. On the contrary, in the natural processes of nutrition, fibrin is assimilated by the tissues and takes part in their organization, only when it is absorbed by them from the blood- vessels in a fluid form. As soon as it is once coagulated by any means, it passes into an unnatural condition, and must be again liquefied and absorbed into the blood before it can be assimilated. As the fibrin, therefore, is maintained in its natural condition of fluidity by the movement of the circulating blood in the interior of the vessels, anything which interferes with this circulation will induce its coagulation. If a ligature be placed upon an artery in the living subject, the blood which stagnates above the ligature coagulates just as it would do if entirely removed from the circula- tion. If the vessel be ruptured or lacerated, the blood which escapes from it into the areolar tissue coagulates, because here also it is withdrawn from the circulation. It coagulates also in the interior of the vessels after death owing to the same cause, viz : stoppage of the circulation. During the last moments of life, when the flow of blood through the cavities of the heart is impeded, the fibrin often coagulates, in greater or less abundance, upon the moving chordss tendineas and the edges of the valves, just as it would do if withdrawn from the body and stirred with a bundle of twigs. In every instance, the coagulation of the fibrin is a COAGULATION OF THE BLOOD. 195 Vertical section of a. R e- CENT COAGCLCM, showing the greater accumulation of blood-globules at the bottom. morbid phenomenon, dependent on the cessation or disturbance of the circulation. If the blood be allowed to coagulate in a bowl, and the clot be then divided by a vertical section, it will be seen that its lower portion is softer and of a deeper red than the upper. (Fig. 65.) This is because the globules, which are of greater specific gravity than the plasma, sink toward the bottom of the vessel before coagu- lation takes place, and accumulate in the lower portion of the blood. This deposit of the globules, however, is only partial; be- cause they are soon fixed and entangled by the solid mass of the coagulum, and thus retained in the position in which they hap- pened to be at the moment that coagulation occurred. If the coagulation, however, be delayed longer than usual, or if the globules sink more rapidly than is customary, they will have time to subside entirely from the upper portion of the blood, leaving a layer at the surface which is composed of plasma alone. When coagulation then takes place, this upper portion solidifies at the same time with the rest, and the clot then presents two dif- ferent portions, viz., a lower portion of a dark red color, in which the globules are accumulated, and an upper portion from which the globules have subsided, and which is of a grayish white color and partially transparent. This whitish layer on the surface of the clot is termed the " buffy coat ;" and the blood presenting it is said to be " buffed." It is an appearance which often presents itself in cases of acute inflammatory disease, in which the coagulation of the blood is unusually retarded. When a clot with a buffy coat begins to contract, the contraction takes place perfectly well in its upper portion, but in the lower part it is impeded by the presence of the globules which have accumulated in large quantity at the bottom of the clot. While the lower part of the coagulum, therefore, remains voluminous, and hardly separates from the sides of the vessel, its upper color- less portion diminishes very much in size ; and as its edges separate from the sides of the vessel, they curl over Bowl of CoAOtlLATED Blood, showing the clot buffed and cupped. 196 THE BLOOD. toward each other, so that the upper surface of the clot becomes more or less excavated or cup-shaped. (Fig. 66.) The blood is then said to be " buffed and cupped." These appearances do not present themselves under ordinary conditions, but only when the blood has become altered by disease. The entire quantity of blood existing in the body has never been very accurately ascertained. It is not possible to extract the whole of it by opening the large vessels, since a certain portion will always remain in the cavities of the heart, in the veins, and in the capil- laries of the head and abdominal organs. The other methods which have been practised or proposed from time to time are all liable to some practical objection. We have accordingly only been able thus far to ascertain the minimum quantity of blood existing in the body. Weber and Lehmann^ ascertained as nearly as possible the quantity of blood in two criminals who suflered death by decapitation ; in both of which cases they obtained essen- tially similar results. The body weighed before decapitation 133 pounds avoirdupois. The blood which escaped from the vessels at the time of decapitation amounted to 12.27 pounds. In order to estimate the quantity of blood which remained in the vessels, the experimenters then injected the arteries of the head and trunk with water, collected the watery fluid as it escaped from the veins, and ascertained how much solid matter it held in solution. This amounted to 477.22 grains, which corresponded to 4.38 pounds of blood. The result of the experiment is therefore as follows : — Blood which escaped from the vessels . . . . .12.27 pounds. " remained in the body ..... 4.38 " Whole quantity of blood in the living body, 16.65 The weight of the blood, then, in proportion to the entire weight of the body, was as 1 : 8 ; and the body of a healthy man, weighing 140 pounds, will therefore contain on the average at least 17J pounds of blood. ' Physiological Chemistry, vol. i. p. 638. RESPIRATION". 197 CHAPTER XII. RESPIRATION. The blood as it circulates in the arterial system has a bright scarlet color ; but as it passes through the capillaries it gradually becomes darker, and on its arrival in the veins its color is a deep purple, and in some parts of the body nearly black. There are, therefore, two kinds of blood in the body ; arterial blood, which is of a bright color, and venous blood, which is dark. Now it is found that the dark-colored venous blood, which has been contaminated by passing through the capillaries, is unfit for further circulation. It is incapable, in this state, of supplying the organs with their healthy stimulus and nutrition, and has become, oh the contrary, deleterious and poisonous. It is accordingly carried back to the heart by the veins, and thence sent to the lungs, where it is recon- verted into arterial blood. The process by which the venous blood is thus arterialized and renovated, is known as the process of re- spiration. This process takes place very actively in the higher animals, and probably does so to a greater or less extent in all animals without exception. Its essential conditions are that the circulating fluid should be exposed to the influence of atmospheric air, or of an aerated fluid ; that is, of a fluid holding atmospheric air or oxygen in solution. The respiratory apparatus consists essentially of a moist and permeable animal membrane, the respiratory membrane, with the bloodvessels on one side of it, and the air or aerated fluid on the other. The blood and the air, consequently, do not come in direct contact with each other, but absorption and exhalation take place from one to the other through the thin membrane which lies between. The special anatomical arrangement of the respiratory apparatus differs in different species of animals. In most of those inhabiting the water, the respiratory organs have the form of gills or branchice ; that is, delicate filamentous prolongations of some part of the 198 EESPIEATION. Head and Gills op Menobranchus. integument or mucous membranes, wlaich contain an abundant supply of bloodvessels, and which hang out freely into the sur- rounding water. In many kinds of aquatic lizards, as, for exam- ple, in menohranchus (Fig. 67), '^* there are upon each side of the neck three delicate feathery tufts of thread-like prolonga- tions from the mucous mem- brane of the pharynx, which pass out through fissures in the side of the neck. Each tuft is composed of a prin- cipal stem, upon which the filaments are arranged in a pinnated form, like the plume upon the shaft of a feather. Each filament, when examined by itself, is seen to consist of a thin, rib- bon-shaped, double fold of mucous membrane, in the interior of which there is a plentiful network of minute bloodvessels. The dark blood, as it comes into the filament from the branchial artery, is exposed to the influence of the water in which the filament is bathed, and as it circulates through the capillary network of the gills is reconverted into arterial blood. It is then carried away by the branchial vein, and passes into the general current of the cir- culation. The apparatus is further supplied with a cartilaginous framework, and a set of muscles by which the gills are gently waved about in the surrounding water, and constantly brought into con- tact with fresh portions of the aerated fluid. Most of the aquatic animals breathe by gills similar in all their essential characters to those described above. In terrestrial and air-breathing animals, however, the respiratory apparatus is situated internally. In them, the air is made to penetrate into the interior of the body, into certain cavities or sacs called the lungs, which are lined with a vascular mucous membrane. In the salamanders, for example, which, though aquatic in their habits, are air-breathing animals, the lungs are two long cylindrical sacs, running nearly the entire length of the body, commencing anteriorly by a communi- cation with the pharynx, and terminating by rounded extremities at the posterior part of the abdomen. These lungs, or air-sacs, have a smooth internal surface; and the blood which circulates through their vessels is arterialized by exposure to the air contained in their cavities. The air is forced into the lungs by a kind of EESPIEATION. 199 Fig. 68. swallowing movement, and is after a time regurgitated and dis- charged, in order to make room for a fresh supply. In the frog, tortoise, serpents, &c., the structure of the lung is a little more complicated, since respiration is more active in these animals, and a more perfect organ is requisite to accomplish the arterialization of the blood. In these animals, the cavity of the lung, instead of being simple, is divided by incomplete partitions into a number of smaller cavities or " cells." The cells all communicate with the central pulmonary cavity; and the partitions, which join each other at various angles, are all composed of thin, projecting, double folds of the lining membrane, with bloodvessels ramifying between them. (Fig. 68.) By this arrangement, the extent of surface presented to the air by the pulmonary membrane isjnuch increased, and the arterialization of the blood takes place with a corresponding degree of rapidity. In the human subject, and in all the warm- blooded quadrupeds, the lungs are constructed on a plan which is essentially similar to the above, and which differs from it only in the greater extent to which the pulmonary cavity is subdivided, and the surface of the respiratory membrane increased. The respiratory apparatus (Fig. 69) commences with the larynx, which communicates with the pharynx at the upper part of the neck. Then follows the trachea, a membranous tube with cartilaginous rings; which, upon its entrance into the chest, divides into the right and left bronchus. These again divide successively into second- ary and tertiary bronchi; the subdivision continuing, while the bronchial tubes grow smaller and more numerous, and separate constantly from each other. As they diminish in size, the tubes grow more delicate in structure, and the cartilaginous rings and plates disappear from their walls. They are finally reduced, accord- ing to KoUiker, to the size of ^^ of an inch in diameter ; and are composed only of a thin mucous membrane, lined with pavement epithelium, which rests upon an elastic fibrous layer. They are then known as the "ultimate bronchial tubes." Each ultimate bronchial tube terminates in a division or islet of the pulmonary tissue, about y'^ of an inch in diameter, which is termed a "pulmonary lobule." Each pulmonary lobule resembles in its structure the entire frog's lung in miniature. It consists of a Lung of Froo, showing its internal sur- face 200 EESPIEATIOX. Fig. 69. Fig. 70. Human Larynx, Trachea, Bronchi, and Lungs; showing the ramification of the bronchi, and the division of the lungs into lobules. vascular membrane inclosing a cavity; which cavity is divided into a large number of secondary compartments by thin septa or partitions, which project from its internal surface. (Fig. 70.) These secondary cavities are the " pulmonary cells," or " vesicles." Each vesicle is about ^-g of an inch in diameter; and is covered on its exterior with a close network of ca- pillary bloodvessels, which dip down into the spaces between the adjacent vesicles, and expose in this way a double surface to the air which is contained in their cavities. Between the vesicles, and in the interstices between the lobules, there is a large quan- tity of yellow elastic tissue, which gives firmness and resiliency to the pulmonary structure. The pulmonary vesicles, accord- siNGLE LoBCLE OF Hu- • ^^ ^j^g obscrvatious of Kdlliker, are MAN Lung. — a. Ultimate bron- ° _ ' chiaitube. h. Cavity of lobule, lined cvcrywhcre with a layer of pavement c,j,c. Pulmonary cells, or vesi- epithelium, COUtinUOUS with that in the RESPIKATORY MOVEMENTS OF THE CHEST. 201 ultimate bronchial tubes. The whole extent of respiratory sur- hce in both lungs has been calculated by Lieberkiihn' at 'fourteen hundred square feet. It is plainly impossible to make a precisely accurate calculation of this extent; but there is every reason to believe that the estimate adopted by Lieberkiihn, regarded as approximative, is not by any means an exaggerated one. The great multiplication of the minute pulmonary vesicles, and of the partitions between them, must evidently increase to an extraor- dinary degree the extent of surface over which the blood, spread out in a thin layer, is exposed to the action of the air. These anatomical conditions are, therefore, the most favorable to its rapid and complete arterialization. Respiratory Movements of the Chest. — The air which is con- tained in the pulmonary lobules and vesicles becomes rapidly vitiated in the process of respiration, and requires therefore to be expelled and replaced by a fresh supply. This exchange or renovation of the air is effected by alternate movements of the chest, of expansion and collapse, which are termed the " respiratory movements of the chest." The expansion of the chest is effected by two sets of mus- cles, viz., first, the diaphragm, and, second, the intercostals. While the diaphragm is in a state of relaxation, it has the form of a vaulted partition between the thorax and abdomen, the edges of which are attached to the inferior extremity of the sternum, the inferior costal cartilages, the borders of the lower ribs and the bodies of the lumbar vertebne, while its convexity rises high into the cavity of the chest, as far as the level of the fifth rib. When the fibres of the diaphragm contract, their curvature is necessarily dimi- nished; and they approximate a straight line, exactly in proportion to the extent of their contraction. Consequently, the entire con- vexity of the diaphragm is diminished in the same proportion; and it descends toward the abdomen, enlarging the cavity of the chest from above downward, (i^'ig. 71.) At the same time the inter- costal muscles enlarge it in a lateral direction. For the ribs, arti- culated behind with the bodies of the vertebrge, and joined in front to the sternum by the flexible and elastic costal cartilages, are so arranged that, in a position of rest, their convexities look obliquely outward and downward. When the movement of inspiration is about to commence, the first rib is fixed by the contraction of the ' In Simon's Chemistry of Man, Philada. ed., 1846, p. 109. 202 RESPIRATION. Fig. 71. scaleni muscles, and the intercostal muscles then contracting simul- taneously, the ribs are drawn upward. In this movement, as each rib rotates upon its articulation with the spinal column at one extremity, and with the sternum at the other, its convexity is necessarily carried outward at the same time that it is drawn upward, and the pa- rietes of the chest are, therefore, expanded laterally. The sternum itself rises slightly with the same movement, and enlarges to some extent the antero-posterior diameter of the thorax. By the simultaneous action, therefore, of the diaphragm which descends, and of the intercostal muscles which lift the ribs and the sternum, the cavity of the chest is expanded in every direction, and the air passes inward, through the trachea and bronchial tubes, by the simple force of aspiration. After the movement of inspiration is ac- complished, and the lungs are filled with air, the diaphragm and intercostal muscles relax, and a movement of expiration takes place, by which the chest is partially col- lapsed, and a portion of the air contained in the pulmonary cavity expelled. The movement of expiration is entirely a passive one, and is accomplished by the action of three different forces. First, the abdominal organs, which have been pushed out of their usual position by the descent of the diaphragm, fall backward by their own weight and carry upward the relaxed diaphragm before them. Secondly, the costal cartilages, which are slightly twisted out of shape when the ribs are drawn upward, resume their natural position as soon as the muscles are relaxed, and, drawing the ribs down again, compress the sides of the chest. Thirdly, the pul- monary tissue, as we have already remarked, is abundantly sup- plied with yellow elastic fibres, which retract by virtue of their own elasticity, in every part of the lungs, after they have been forcibly distended, and, compressing the pulmonary vesicles, drive out a portion of the air which they contained. By the constant Diagram illustratingi THE ReSPIRATORT MOVE- MENTS. — a. Cavity of the chest. h. Diaphragm. The dark out- lines show the figure of the chest when collapsed ; the dotted lines show the same when expanded. RESPIEATOEY MOVEMENTS OF THE CHEST. 203 recurrence of these alternating movements of inspiration and expi- ration, fresh portions of air are constantly introduced into and expelled from the chest. The whole of the air, however, is not exchanged at each move- ment of respiration. On the contrary, a very considerable quantity remains in the pulmonary cavity after the most complete expira- tion; and even after the lungs have been removed from the chest, they still contain a large quantity of air which cannot be entirely displaced by any violence short of disintegrating and disorganizing the pulmonary tissue. It is evident, therefore, that only a com- paratively small portion of the air in the lungs passes in and out with each respiratory movement ; and it will require several suc- cessive respirations before all the air in the chest can be entirely changed. It has not been possible to ascertain with certainty the exact proportion in volume which exists between the air which is alternately inspired and expired, or "tidal" air, and that which remains constantly in the chest, or "residual" air, as it is called. It has been estimated, however, by Dr. Carpenter,^ from the reports of various observers, that the volume of inspired and expired air varies from 10 to 13 per cent, of the entire quantity contained in the chest. If this estimate be correct, it will require from eight to ten respirations to change the whole quantity of air in the cavity of the chest. It is evident, however, from the foregoing, that the inspiratory and expiratory movements of the chest cannot be sufficient to change the air at all in the pulmonary l©bules and vesicles. The air which is drawn* in with each inspiration penetrates only into the trachea and bronchial tubes, until it occupies the place of that which was driven out by the last expiration. By the ordinary respiratory movements, therefore, only that small portion of the air lying nearest the exterior, in the trachea and large bronchi would fluctuate backward and forward, without ever penetrating into the deeper parts of the lung, were there no other means pro- vided for its renovation. There are, however, two other forces in play for this purpose. The first of these is the diffusive power of the gases themselves. The air remaining in the deeper parts of the chest is richer in carbonic acid and poorer in oxygen than that which has been recently inspired ; and by the laws of gaseous dif- fusion there must be a constant interchange of these gases between ' Human Physiology, Philada. ed., 1855, p. 300. 204 EESPIEATION. the pulmonary vesicles and the trachea, tending to mix them equally in all parts of the lung. This mutual diffusion and inter- mixture of the gases will therefore tend to renovate, partially at least, the air in the pulmonary lobules and vesicles. Secondly the trachea and bronchial tubes, down to those even of the smallest size, are lined with a mucous membrane which is covered with a ciliated epithelium. The movement of these cilia is found to be directed always from below upward ; and, like ciliary motion wherever it occurs, has the effect of producing a current in the same direction, in the fluids covering the mucous membrane. The air in the tubes must partici- Fig- 72. pate, to a certain extent, in ■|H|BBHB^BH|HBBBB^H this and a double ^^^S^^^^^^^^^H^^^^^^H stream of air therefore is estab- ^KKmljjj^ lished in each bronchial tube; ^/^^^BS^S/K^^^^^^^^^ current passing from with- ^^^^P^^PI^^^^^^^^^^^B in outward along the walls of ^^^^^M^HHMj^^^^^^^HU the tube, and a return ^ . , , passing from without inward, Small Bronchial Tube, showing outward r o i and inward current, produced by ciliary motion. alono" the Central part of itS cavity. (Fig. 72.) By this means a kind of aerial circulation is constantly maintained in the interior of the bronchial tubes ; which, combined with the mutual diffusion of the gases and the alternate expansion and collapse of the chest, effectually accomplish the renovation of the air contained in all parts of the pulmonary cavity. Eespiratory Movements of the Glottis. — Beside the move- ments of expansion and collapse already described, belonging to the chest, there are similar respiratory movements which take place in the larynx. If the respiratory passages be examined after death, in the state of collapse in which they are usually found, it will be noticed that the opening of the glottis is very much smaller than the cavity of the trachea below. The glottis itself presents the appearance of a narrow chink, while the passage for the inspired air widens in the lower part of the larynx, and in the trachea constitutes a spacious tube, nearly cylindrical in shape, and over half an inch in diameter. We have found, for instance, that in the human subject the space included between the vocal chords has an area of only 0.15 to 0.17 square inch ; while the calibre of the trachea in the middle of its length is 0.45 square inch. RESPIRATORY MOVEMENTS OF THE GLOTTIS. 205 This disproportion, however, which is so evident after death, does not exist during life. While respiration is going on, there is a constant and regular movement of the vocal chords, synchronous with the inspiratory and expiratory movements of the chest, by Fig. 73. Fig. 74. Human Larynx, viewed from above in its ordinary post-mortem condition. — a. Vocal chords, b. Thyroid cartilage, cc. Ary- tenoid cartilages, o. Opening of the glottis. The same, with the glottis opened by separation of the vocal chords. — a. Vocal chords. 6. Thyroid cartilage, vc. Aryte- noid cartilages, o. Opening of the glottis. Fig. 75. which the size of the glottis is alternately enlarged and diminished. At every inspiration, the glottis opens and allows the air to pass freely into the trachea ; at every expiration it collapses, and the air is driven out through it from be- low. These movements are called the " respiratory movements of the glottis." They correspond in ev«ry respect with those of the chest, and are excited or retarded by similar causes. When- ever the general movements of respira- tion are hurried and labored, those of the glottis become accelerated and in- creased in intensity at the same time ; and when the movements of the chest are slower or fainter than usual, owing to debility, coma, or the like, those of the glottis are diminished in the same proportion. 1.1 „ „• , ,• ^ii Human Larynx, POSTERIOR n the respiratory motions of the viEw.-a. Thyroid cartilage. 6. Epi- glottis, as in those of the chest, the glottis, cc. Arytenoid cartilages, d. , ry • ... . . Cricoid cartilage, ee. Posterior crico- movement Ot inspiration is an active arytenoid muscles. /. Trachea. 206 EESPIRATION. one, and that of expiration passive. In inspiration, the glottis is opened by contraction of the posterior crico-arytenoid muscles. (Fig. 75.) These muscles originate from the posterior surface of the cricoid cartilage, near the median line; and their fibres, running upward and outward, are inserted into the external angle of the arytenoid cartilages. By the contraction of these muscles, during the movement of inspiration, the arytenoid cartilages are rotated upon their articulations with the cricoid, so that their anterior extremities are carried outward, and the vocal chords stretched and separated from each other. (Fig. 74.) In this way, the size of the glottis may be increased from 0.16 to 0.27 square inch. In expiration, the posterior crico-arytenoid muscles are relaxed, and the elasticity of the vocal chords brings them back to their former position. The motions of respiration consist, therefore, of two sets of move- ments : viz., those of the chest, and those of the glottis. These move- ments, in the natural condition, correspond with each other both in time and intensity. It is at the same time and by the same nervous influence, that the chest expands to inhale the air, while the glottis opens to admit it ; and in expiration, the miiscles of both chest and glottis are relaxed, while the elasticity of the tissues, by a kind of passive contraction, restores the parts to their original condition. CHANGES IN THE AIR DURING RESPIRATION. The atmospheric air, as it is drawn into the cavity of the lungs, is a mixture of oxygen and nitrogen, in the proportion of about 21 per cent., by volume, of oxygen, to 79 per cent, of nitrogen. It also contains about one-twentieth per cent, of carbonic acid, a vary- ing quantity of watery vapor, and some traces of ammonia. If col- lected and examined, after passing through the lungs, it is found to have become altered in the following essential particulars, viz : — 1st. It has lost oxygen. 2d. It has gained carbonic acid. And 3d. It has absorbed the vapor of water. Beside the two latter substances, there are also exhaled with the expired air a very small quantity of nitrogen, over and above what was taken in with inspiration, and a little animal matter in a gaseous form, which communicates a slight but peculiar odor to the breath. The air is also somewhat elevated in temperature, by CHANGES IN THE BLOOD DURING RESPIRATION. 207 contact with the pulmonary mucous membrane. By far the most important part, however, of the above changes suffered by the air, consists in its loss of ox3^gen, and its absorption of carbonic acid. The oxygen which disappears from the inspired air is not entirely replaced in the carbonic acid exhaled ; that is, there is less oxygen in the carbonic acid which is returned to the air by expiration than has been lost during inspiration. There is even more oxygen absorbed than is given off again in both the carbonic acid and aqueous vapor together, which are exhaled from the lungs.^ There is, then, a constant disappearance of oxygen from the air used in respiration, and a constant accumu- lation of carbonic acid. The proportion of oxygen which disappears in the interior of the body, over and above that which is returned in the breath under the form of carbonic acid, varies in different kinds of animals. In the herbivora, it is about 10 per cent, of the whole amount of oxy- gen inspired ; in the carnivora, 20 or 25 per cent,, and even more. It is a very remarkable fact, also, and an important one, as regards the theory of respiration, that, in the same animal, the proportion of oxygen absorbed, to that of carbonic acid exhaled, varies according to the quality of the food. In dogs, for instance, while fed on ani- mal food, according to the experiments of Eegnault and Eeiset, 25 per cent, of the inspired oxygen disappeared in the body of the animal; but when fed on starchy substances, all but 8 per cent, reappeared in the expired carbonic acid. It is already evident, there- fore, from these facts, that the oxygen of the inspired air is not altogether employed in the formation of carbonic acid. CHANGES IN THE BLOOD DURING RESPIRATION. If we pass from the consideration of the changes produced in the air by respiration to those which take place in the blood during the same process, we find, as might have been expected, that the latter correspond inversely with the former. The blood, in passing through the lungs, suffers the following alterations : — 1st. Its color is changed from venous to arterial. 2d. It absorbs oxygen. And 8d. It exhales carbonic acid and the vapor of water. ' Lehmann's Physiological Chemistry, Philada. ed., vol. ii. p. 432. 208 EESPIRATION. The interchange of gases, which takes place during respiration between the air and the blood, is a simple phenomenon of absorp- tion and exhalation. The inspired oxygen does not, as Lavoisier once supposed, immediately combine with carbon in the lungs, and return to the atmosphere under the form of carbonic acid. On the contrary, almost the first fact of importance which has been estab- lished by the examination of the blood in this respect is the fol- lowing, viz : that carbonic acid exists ready formed in the venous blood before its entrance into the lungs ; and, on the other hand, that the oxygen which is absorbed during respiration passes off in a free state with the arterial blood. The real process, as it takes place in the lung, is, therefore, for the most part, as follows : The blood comes to the lungs already charged with carbonic acid. In passing through the pulmonary capillaries, it is exposed to the influence of the air in the cavity of the pulmonary cells, and a transudation of gases takes place through the moist animal membranes of the lung. Since the blood in the capillaries contains a larger proportion of carbonic acid than the air in the air-vesicles, a portion of this gas leaves the blood and passes out through the pulmonary mem- brane ; while the oxygen, being more abundant in the air of the vesicles than in the circulating fluid, passes inward at the same time, and is condensed by the blood. In this double phenomenon of exhalation and absorption, which takes place in the lungs, both parts of the process are equally necessary to life. It is essential for the constant activity and nutri- tion of the tissues that they be steadily supplied with oxygen by the blood ; and if this supply be cut off, their functional activity ceases. On the other hand, the carbonic acid which is produced in. the body by the processes of nutrition becomes a poisonous sub- stance, if it be allowed to collect in large quantity. Under ordinary circumstances, the carbonic acid is removed by exhalation through the lungs as fast as it is produced in the interior of the body ; but if respiration be suspended, or seriously impeded, since the produc- tion of carbonic acid continues, while its elimination is prevented, it accumulates in the blood and in the tissues, and terminates life in a few moments, by a rapid deterioration of the circulating fluid, and more particularly by its poisonous effect on the nervous system. The deleterious effects of breathing in a confined space will therefore very soon become apparent. As respiration goes on, the oxygen of the air constantly diminishes, and the carbonic acid, mingled with it by exhalation, increases in quantity. After a time CHANGES OF THE BLOOD DURING RESPIRATION. 209 the air becomes accordingly so poor in oxygen that, by that fact alone, it is incapable of supporting life. At the same time, the carbonic acid becomes so abundant in the air vesicles that it prevents the escape of that which already exists in the blood ; and the dele- terious effect of its accumulation in the circulating fluid is added to that produced by a diminished supply of oxygen. An increased proportion of carbonic acid in the atmosphere is therefore injurious in a similar manner, although there may be no diminution of oxy- gen; since by its presence it impedes the elimination of the carbonic acid already formed in the blood, and induces the poisonous effects which result from its accumulation. Examination of the blood shows furthermore that the interchange of gases in the lungs is not complete but only partial in its extent. It results from the experiments of Magendie, Magnus, and others, that both oxygen and carbonic acid are contained in both venous and arterial blood. Magnus' found that the proportion of oxygen to carbonic acid, by volume, in arterial blood was as 10 to 25 ; in venous blood as 10 to 40. The venous blood, then, as it arrives at the lungs, still retains a remnant of the oxygen which it had pre- viously absorbed ; and in passing through the pulmonary capillaries it gives off only a part of the carbonic acid with which it has become charged in the general circulation. The. oxygen and carbonic acid of the blood exist in a state of solution in the circulating fluid, and not in a state of intimate chemi- cal combination. This is shown by the fact that both of these substances may be withdrawn from the blood by simple exhaustion with an air pump, or by a stream of any other indifferent gas, such as hydrogen, which possesses sufficient physical displacing power. Magnus found^ that freshly drawn arterial blood yielded by simple agitation with carbonic acid more than 10 per cent, of its volume of oxygen. The carbonic acid may also be expelled from venous blood by a current of pure oxygen, or of hydrogen, or, in great measure, by simple agitation with atmospheric air. There is some difficulty in determining, however, whether the carbonic acid of the blood be altogether in a free state, or whether it be partly in a state of loose chemical combination with a base, under the form of an alkaline bicarbonate. A solution of bicarbonate of soda itself will lose a portion of its carbonic acid, and become reduced to the ' In Lehmann, op. cit., vol. i. p. 570. • ^ In Robin and Verdeil, op. cit., vol. ii. p. 34. 14 210 EESPIRATION". condition of a carbonate, bj simple exhaustion under the air-pump, or by agitation with pure hydrogen at the temperature of the body. Lehmann has found' that after the expulsion of all the carbonic acid removable by the air-pump and a current of hydrogen, there still remains, in ox's blood, 0.1628 per cent, of carbonate of soda; and he estimates that this quantity is sufficient to have retained all tbe carbonic acid, previously given off", in the form of a bicarbonate. It makes little or no difference, however, so far ar regards the pro- cess of respiration, whether the carbonic acid of the blood exist in an entirely free state, or under the form of an alkaline bicarbonate ; since it may be readily removed from this combination, at the tem- perature of the body, by contact with an indifferent gas. The oxygen and carbonic acid of the blood are in solution prin- cipally in the blood- globules, and not in the plasma. The researches of Magnus have shown^ that the blood holds in solution 2| times as much oxygen as pure water could dissolve at the same tempera- ture; and that while the serum of the blood, separated from the globules, exerts no more solvent power on oxygen than pure water, defibrinated blood, that is, the serum and globules mixed, dissolves quite as much oxygen as the fresh blood itself. The same thing is true with regard to the carbonic acid. It is therefore the semi- fluid blood-globules which retain these two gases in solution ; and since the color of the blood depends entirely upon that of the glo- bules, it is easy to understand why the blood should alter its hue from purple to scarlet in passing through the lungs, where the globules give up carbonic acid, and absorb a fresh quantity of oxygen. The above change may readily be produced outside the body. If freshly drawn venous blood be shaken in a bottle with, pure oxygen, its color changes at once from purple to red ; and the same change will take place, though more slowly, if the blood be simply agitated with atmospheric air. It is for this reason that the surface of defibrinated venous blood, and the external parts of a dark-colored clot, exposed to the atmosphere, become rapidly red- dened, while the internal portions retain their original color. The process of respiration, so far as we have considered it, con- sists in an alternate interchange of carbonic acid and oxygen in the blood of the general and pulmonary circulations. In the pulmonary circulation, carbonic acid is given off and oxygen absorbed ; while ' Op. cit., vol. i. p. 393. 2 In Robin and Verdeil, op. cit., vol. ii. pp. 28—32. CHANGES OF THE BLOOD DURING RESPIRATION. 211 in the general circulation tlie oxygen gradually disappears, and is replaced, in the venous blood, by carbonic acid. The oxygen' which thus disappears from the blood in the general circulation does not, for the most part, enter into direct combination in the blood itself. On the contrary, it exists there, as we have already stated, in the form of a simple solution. It is absorbed, however, from the blood of the capillary vessels, and becomes fixed in the substance of the vascular tissues. The blood may be regarded, therefore, in this respect, as a circulating fluid, destined to transport oxygen from the lungs to the tissues ; for it is the tissues themselves which finally appropriate the oxygen, and fix it in their substance. The next important question which presents itself in the study of the respiratory process relates to the origin of the carbonic acid in the venous blood. It was formerly supposed, when Lavoisier first discovered the changes produced in the air by respiration, that the production of the carbonic acid could be accounted for in a very simple manner. It was thought to be produced in the lungs by a direct union of the inspired oxygen with the carbon of the blood in the pulmonary vessels. It was found afterward, however, that this could not be the case; since carbonic acid exists already formed in the blood, previous to its entrance into the lungs. It was then imagined that the oxidation of carbon, and the consequent produc- tion of carbonic acid, took place in the capillaries of the general circulation, since it could not be shown to take place in the lungs, nor between the lungs and the capillaries. The truth is, however, that no direct evidence exists of such a direct oxidation taking place anywhere. The formation of carbonic acid, as it is now understood, takes place in three different modes: 1st, in the lungs; 2d, in the blood; and 3d, in the tissues. First, in the lungs. There exists in the pulmonary tissue a pecu- liar acid substance first described by Yerdeil' under the name of " pneumic" or "pulmonic" acid. It is a crystallizable body, soluble in water, which is produced in the substance of the pulmonary tissue by transformation of some of its other ingredients, in the same manner as sugar is produced in the tissue of the liver. It is on account of the presence of this substance that the fresh tissue of the lung has usually an acid reaction to test-paper, and that it has also the property, which has been noticed by several observers, of ' Robin and Verdeil, op. cit., vol. ii. p. 460. 212 EESPIRATION. decomposing the metallic cyanides, with the production of hydro- cyanic acid ; a property not possessed by sections of areolar tissue, the internal surface of the skin, &c. &c. When the blood, there- fore, comes in contact with the pulmonary tissue, which is permeated everywhere by pneumic acid in a soluble form, its alkaline carbonates and bicarbonates, if any be present, are decom- posed with the production on the one hand of the pneumates of soda and potass, and on the other of free carbonic acid, which is exhaled, M, Bernard has found^ that if a solution of bicarbonate of soda be rapidly injected into the jugular vein of a rabbit, it becomes decomposed in the lungs with so rapid a development of carbonic acid, that the gas accumulates in the pulmonary tissue, and even in the pulmonary vessels and the cavities of the heart, to such an extent as to cause immediate death by stoppage of the circulation. In the normal condition, however, the carbonates and bicarbonates of the blood arrive so slowly at the lungs that as fast as they are decomposed there, the carbonic acid is readily exhaled by expiration, and produces no deleterious efiect on the circulation. Secondly, in the blood. There is little doubt, although the fact has not been directly proved, that some of the oxygen definitely dis- appears, and some of the carbonic acid is also formed, in the sub- stance of the blood-globules during their circulation. Since these globules are anatomical elements, and since they undoubtedly go through with nutritive processes analogous to those which take place in the elements of the solid tissues, there is no reason for dis- believing that they also require oxygen for their support, and that they produce carbonic acid as one of the results of their interstitial decomposition. While the oxygen and carbonic acid, therefore, contained in the globule^, are for the most part transported by these bodies from the lungs to the tissues, and from the tissues back again to the lungs, they probably take part, also, to a certain extent, in the nutrition of the blood-globules themselves. Thirdly, in the tissues. This is by far the most important source of the carbonic acid in the blood. From the experiments of Spal- lanzani, W, Edwards, Marchand and others, the following very important fact has been established, viz., that every organized tissue and even every organic substance, when in a recent condition, has the power of absorbing oxygen and of exhaling carbonic acid. G, Liebig, for example,^ found that frog's muscles, recently prepared and com- ' Archives Gen. de Med,, xvi, 222. ^ In Lehmann, op. cit., vol. ii. p. 474. CHANGES OF THE BLOOD DURING RESPIRATION. 213 pletely freed from blood, continued to absorb oxygen and discharge carbonic acid. Similar experiments with other tissues have led to a similar result. The interchange of gases, therefore, in the process of respiration, takes place mostly in the tissues themselves. It is in their substance that the oxygen becomes fixed and assimi- lated, and that the carbonic acid takes its origin. As the blood in the lungs gives up its carbonic acid to the air, and absorbs oxygen from it, so in the general circulation it gives up its oxygen to the tissues, and absorbs from them carbonic acid. We come lastly to examine the exact mode by which the car- bonic acid originates in the animal tissues. Investigation shows that even here it is not produced hy a process of oxidation, or direct union of oxygen ivith the carbon of the tissues, hut in some other and more indirect mode. This is proved by the fact that animals and fresh animal tissues will continue to exhale carbonic acid in an atmo- sphere of hydrogen or of nitrogen, or even when placed in a vacuum. Marchand found* that frogs would live for from half an hour to an hour in pure hydrogen gas ; and that during this time they exhaled even more carbonic acid than in atmospheric air, owing probably to the superior displacing power of hydrogen for carbonic acid. For while 15,500 grains' weight of frogs exhaled about 1.13 grain of carbonic acid per hour in atmospheric air, they exhaled during the same time in pure hydrogen as much as 4.07 grains. The same observer found that frogs would recover on the admission of air after remaining for nearly half an hour in a nearly complete vacuum ; and that if they were killed by total abstraction of the air, 15,500 grains weight of the animals were found to have eliminated 9.3 grains of carbonic acid. The exhalation of carbonic acid by the tissues does not, therefore, depend directly upon the access of free oxygen. It cannot go on, it is true, for an indefinite time, any more than the other vital processes, without the presence of oxygen. But it may continue long enough to show that the carbonic acid exhaled is not a direct product of oxidation, but that it originates, on the contrary, in all probability, by a decomposi- tion of the organic ingredients of the tissues, resulting in the "pro- duction of carbonic acid on the one hand, and of various other substances on the other, with which we are not yet fully acquainted; in very much the same manner as the decomposition of sugar during fermentation gives rise to alcohol on the one hand and to ' Lehmann, op. cit., vol. ii. p. 442. 214 KESPIRATION. carbonic acid on the other. The fermentation of sugar, when it has once commenced, does not require the continued access of air. It will go on in an atmosphere of hydrogen, or even when confined in a close vessel over mercury; since its carbonic acid is not produced by direct oxidation, but by a decomposition of the sugar already present. For the same reason, carbonic acid will continue to be exhaled by living or recently dead animal tissues, even in an atmo- sphere of hydrogen, or in a vacuum. Carbonic acid makes its appearance, accordingly, in the tissues, as one product of their decomposition in the nutritive process. From them it is taken up by the blood, either in simple solution or in loose combination as a bicarbonate, transported by the circulation to the lungs, and finally exhaled from the pulmonary mucous mem- brane in a gaseous form. • The carbonic acid exhaled from the lungs should accordingly be studied by itself as one of the products of the animal organism, and its quantity ascertained in the different physiological conditions of the body. According to the researches of Vierordt,^ which are regarded as the most accurate on this subject, an adult man gives off 1.62 cubic inch of carbonic acid with each normal expiration. This would give 19.16 cubic inches per minute, 1149.6 cubic inches per hour, and 15.4 cubic feet per day. The amount of carbonic acid exhaled, however, varies from time to time, according to many different circumstances; so that no such estimate can represent correctly its quantity at all times. These variations have been very fully investigated by Andral and Gavarret,^ who found that the principal conditions modifying the amount of this gas produced were age, sex, constitution and development. The variations were very marked in different individuals, notwithstanding that the experiments were made at the same period of the day, and with the subject as nearly as possible in the same condition. Thus they found that the quantity of carbonic acid exhaled per hour in five different individuals was as follows :— Quantity of Carbonic Acid pek houb . In subjec t No. 1 1207 cubic inches (( (I "2 970 11 " 11 li "3 . . . . . 1250 11 11 a u "4 . . ". . . 1250 11 11 (1 (f "5 1591 11 « ' In Lehmann, op. cit., vol. ii. p. 439. ^ Annales de Cliimie et de Pharmacie, 1843, vol. viii. p. 129. CHANGES OF THE BLOOD DUKING RESPIRATION. 215 With regard to the difference produced by age, it was found that from the period of eight years up to puberty the quantity of car- bonic acid increases constantly with the age. Thus a boy of eight years exhales, on the average, 564 cubic inches per hour ; while a boy of fifteen years exhales 981 cubic inches in the same time. Boys exhale during this period more carbonic acid than girls of the same age. In males this augmentation of the quantity of carbonic acid continues till the twenty-fifth or thirtieth year, when it reaches, on the average, 1398 cubic inches per hour. Its quantity then remains stationary for ten or fifteen years ; then diminishes slightly from the fortieth to the sixtieth year; and after sixty years dimi- nishes in a marked degree, so that it may fall so low as 1038 cubic inches. In one superannuated person, 102 years of age, Andral and Gavarret found the hourly quantity of carbonic acid to be only 665 cubic inches. In women, the increase of carbonic acid ceases at the period of puberty; and its production then remains constant until the cessa- tion of menstruation, about the fortieth or forty -fifth year. At that time it increases again until after fifty years, when it subsequently diminishes v^ith the approach of old age, as in men. Pregnancy, occurring at any time in the above period, imm.ediately produces a temporary increase in the quantity of carbonic acid. The strength of the constitution, and more particularly the deve- lopment of the muscular system, was found to have a very great influence in this respect ; increasing the quantity of carbonic acid very much, in proportion to the weight of the individual. The largest production of carbonic acid observed was in a young man, 26 years of age, whose frame presented a remarkably vigorous and athletic development, and who exhaled 1591 cubic inches per hour. This large quantity of carbonic acid, moreover, in well developed persons, is not owing simply to the size of the entire body, but particularly to the development of the muscular system, since an unusually large skeleton, or an abundant deposit of adipose tissue, is not accompanied by any such increase of the carbonic acid. Andral and Gavarret finally sum up the results of their investiga- tions as follows : — 1. The quantity of carbonic acid exhaled from the lungs in a given time varies with the age, the sex, and the constitution of the subject. 2. In the male, as well as in the female, the quantity of carbonic 216 RESPIRATION. acid varies according to the age; and that independently of the weight of the individual subjected to experiment. 3. During all the periods of life, from that of eight years up to the most advanced age, the male and female may be distinguished by the different quantities of carbonic acid which they exhale in a given time. Other things being equal, the male exhales always a larger quantity than the female. This difference is particularly marked between the ages of 16 and 40 years, during which period the male usually exhales twice as much carbonic acid as the female. 4. In the male, the quantity of carbonic acid increases constantly from eight to thirty years ; and the rate of this increase undergoes a rapid augmentation at the period of puberty. Beyond thirty years the exhalation of carbonic acid begins to decrease, and its diminution is more marked as the individual approaches extreme old age ; so that near the termination of life, the quantity of carbonic acid produced may be no greater than at the age of ten years. 5. In the female, the exhalation of carbonic acid increases accord- ing to the same law as in the male, from the age of eight years until puberty. But at the period of puberty, at the same time with the appearance of menstruation, the exhalation of carbonic acid, contrary to what happens in the male, ceases to increase ; and it afterward remains stationary so long as the menstrual periods recur with regularity. At the cessation of the menses, the quantity of carbonic acid exhaled increases in a notable manner ; then it de- creases again, as in the male, as the woman advances toward old age. 6. During the whole period of pregnancy, the exhalation of car- bonic acid rises, for the time, to the same standard as in women whose menses have ceased. 7. In both sexes, and at all ages, the quantity of carbonic acid is greater as the constitution is stronger, and the muscular systeni more fully developed. Prof. Scharling, in a similar series of investigations,' found that the quantity of carbonic acid exhaled was greater during the diges- tion of food than in the fasting condition. It is greater, also, in the waking state than during sleep ; and in a state of activity than in. one of quietude. It is diminished, also, by fatigue, and by most conditions which interfere with perfect health. The process of respiration is not altogether confined to the lungs, ' Annales de Chimie et de Pharmacie, vol. viii. p. 490. CHANGES OF THE BLOOD DURING RESPIRATION. 217 but the interchange of gases takes place, also, to some extent through the skin. It has been found, by inclosing one of the limbs in an air-tight case, that the air in which it is confined loses oxygen and gains in carbonic acid. By an experiment of this sort, performed by Prof Scharling,^ it was ascertained that the carbonic acid given off from the whole cutaneous surface, in the human subject, is from one- sixtieth to one-thirtieth of that discharged during the same period from the lungs. In the true amphibious animals, that is, those which breathe by lungs, and can yet remain under water for an indefinite period without injury (as frogs and salamanders), the respiratory function of the skin is very active. In these animals, the integument is very vascular, moist, and flexible; and is covered, not with dry cuticle, but with a very thin and delicate layer of epithelium. It, therefore, presents all the conditions necessary for the accomplishment of respiration; and while the animal remains beneath the surface, and the lungs are in a state of inactivity, the exhalation and absorption of gases continue to take place through the skin, and the process of respiration goes on in a nearly unin- terrupted manner. ' In Carpenter's Human Physiology, Philada. ed., 1855, p 308. 218 ANIMAL HEAT. CHAPTER XIII. ANIMAL HEAT. One of the most important phenomena presented by animals and vegetables is the property which they possess of maintaining, more or less constantly, a standard temperature, notwithstanding the external vicissitudes of heat and cold to which they may be sub- jected. If a bar of iron, or a jar of water, be heated up to 100° or 200° F., and then exposed to the air at 50° or 60°, it will imme- diately begin to lose heat by radiation and conduction ; and this loss of heat will steadily continue, until, after a certain time, the temperature of the heated body has become reduced to that of the surrounding atmosphere. It then remains stationary at this point, unless the temperature of the atmosphere should happen to rise or fall ; in which case, a similar change takes place in the inorganic body, its temperature remaining constant, or varying with that of the surrounding medium. With living animals, the case is different. If a thermometer be introduced into the stomach of a dog, or placed under the tongue of the human subject, it will indicate a temperature of 100° F., very nearly, whatever may be the condition of the surrounding atmos- phere at the time. This internal temperature is the same in sum- mer and in winter. If the individual upon whom the experiment has been tried be afterward exposed to a cold of zero, or even of 20° or 30° below zero, the thermometer introduced into the interior of the body will still stand at 100° F. As the body, during the whole period of its exposure, must have been losing heat by radiation and conduction, like any inorganic mass, and has, notwithstanding, main- tained a constant temperature, it is plain that a certain amount of heat has been generated in the interior of the body by means of the vital processes, sufficient to compensate for the external loss. The internal heat, so produced, is known by the name of vital or animal heat. There are two classes of animals in which the production of vital ANIMAL HEAT. 219 beat takes place with such activity that their blood and internal organs are nearly always very much above the external temperature; and which are therefore called "warm-blooded animals." Tliese are mammalia and birds. Among the birds, some species, as the gull, have a temperature as low as 100° F.; but in most of them it is higher, sometimes reaching as high as 110° or 111°. In the mam- malians, to which class man belongs, the animal temperature is never far from 100°. In the seal and the Greenland whale, it has been found to be 10-4°; and in the porpoise, which is an air-breathing animal, 99°.5. In the human subject it is 98° to 100.° When the temperature of the air is below this, the external parts of the body, being most exposed to the cooling influences of radiation and con- duction, fall a little below the standard, and may indicate a tempera- ture of 97°, or even several degrees below this point. Thus, on a very cold day, the thinner and more exposed parts, such as the nose, the ears, and the ends of the fingers, may become cooled down con- siderably below the standard temperature, and may even be con- gealed, if the cold be severe; but the temperature of the internal organs and of the blood still remains the same under all ordinary exposures. If the cold be so intense and long continued as to affect the general temperature of the blood, it at once becomes fatal. It has been found that although a warm-blooded animal usually preserves its natural temperature when exposed to external cold, yet if the actual temperature of the blood become reduced by any means more than 5° or 6° below its natural standard, death inevitably results. The animal, under these circumstances, gradually becomes torpid and insensible, and all the vital operations finally cease. Birds, accordingly, whose natural temperature is about 110°, die if the blood be cooled down to 100°, which is the natural temperature of the mammalia ; and the mammalians die if their blood be cooled down below 94° or 95°. Each of these different classes has there- fore a natural temperature, at which the blood must be maintained in order to sustain life; and even the different species of animals, belonging to the same class, have each a specific temperature which is characteristic of them, and which cannot be raised or lowered, to any considerable extent, without producing death. While in the birds and mammalians, however, the internal pro- duction of heat is so active, that their temperature is nearly always considerably above that of the surrounding media, and suffers but little variation ; in reptiles and fish, on the other hand, its produc- 220 ANIMAL HEAT. tion is much less rapid, and the temperature of their bodies differs but little from that of the air or water which they inhabit. Birds and mammalians are therefore called " warm-blooded," and reptiles and fish " cold-blooded" animals. There is, however, no other distinc- tion between them, in this respect, than one of degree. In reptiles and fish there is also an internal source of heat ; only this is not so active as in the other classes. Even in these animals a difierence is usually found to exist between the temperature of their bodies and that of the surrounding media. John Hunter, Sir Humphrey Davy, Czermak, and others,' have found the temperature of Proteus anguinus to be 63°.5, when that of the air was 55° A] that of a frog 48°, in water at 44°.4 ; that of a serpent 88°.46, in air at 81°.5 ; that of a tortoise 84°, in air at 79°.o ; and that of fish to be from 1°.7 to 2°.5 above that of the surrounding water. The following list^ shows the mean temperature belongmg to animals of different classes and species. Birds. Mammalia. ■ Reptile. Fish. { Animal. Swallow Heron Raven Pigeon Fowl ^ Gull Squirrel Goat Cat Hare Ox Dog Man L Ape Toad Carp Tench Mean Temperatdke. 1110.25 1110.2 108O.5 1070.6 106O.7 1000.0 1050 1020.5 1010.3 1000.4 990,5 990.4 980.6 950.9 510.6 510.25 520.10 In the invertebrate animals, as a general rule, the internal heat is produced in too small quantity to be readily estimated. In some of the more active kinds, however, such as insects and arachnida, it is occasionally generated with such activity that it may be appreciated by the thermometer. Thus, the temperature of the butterfly, when in a state of excitement, is from 5° to 9° above ' Simon's Chemistry of Man, Philadelphia edition, p. 124. 2 Ihid., pp. 123—126. ANIMAL HEAT. 221 that of the air ; and that of the humble-bee from 3° to 10° higher than the exterior. According to the experiments of Mr. Newport/ the interior of a hive of bees may have a temperature of 48*^.5, when the external atmosphere is at 34°.5, even while the insects are quiet; but if they be excited, by tapping on the outside of the hive, it may rise to 102°. In all cases, while the insect is at rest, the temperature is very moderate; but if kept in rapid motion in a confined space, it may generate heat enough to affect the thermo- meter sensibly, in the course of a few minutes. Even in vegetables a certain degree of heat-producing power is occasionally manifest. Usually, the exposed surface of a plant is so extensive in proportion to its mass, that whatever caloric may be generated is too rapidly lost by radiation and evaporation, to be appreciated by ordinary means. Under some circumstances, how- ever, it may accumulate to such an extent as to become readily perceptible. In the process of malting, for example, when a large quantity of germinating grain is piled together in a mass, its ele- vated temperature may be readily distinguished, both by the hand and the thermometer. During the flowering process, also, an unu- sual evolution of heat takes place in plants. The flowers of the geranium have been found to have a temperature of 87°, while that of the air was 81°; and the thermometer, placed in the centre of a clump of blossoms of arum cordifolium, has been seen to rise to 111°, and even 121°, while the temperature of the external air was only 66°.^ Dutroctzet has moreover found, by a series of very ingenious and delicate experiments,^ that nearly all parts of a living plant gene- rate a certain amount of heat. The proper beat of the plant is usually so rapidly dissipated by the continuous evaporation of its fluids, that it is mostly imperceptible by ordinary means ; but if this evaporation be prevented, by keeping the air charged with watery vapor, tbe heat becomes sensible and can be appreciated by a delicate thermometer. Dutrochet used for this purpose a thermo- electric apparatus, so constructed that an elevation of temperature of 1° F., in the substance examined, would produce a deviation in the needle of nearly nine degrees. By this means he found that he could appreciate, without difficulty, the proper temperature of the plant. A certain amount of heat was constantly generated, during ' Carpenter's General and Comparative Physiology, Philadelphia, 1851, p, 852, * Carpenter's Gen, and Comp. Physiology, p, 846. ' Annales des Sciences Naturelles, 2d series, xii, p. 277. 222 ANIMAL HEAT. the day, in the green stems, the leaves, the buds, and even the roots and fruit. The maximum temperature of these parts, above that of the surrounding atmosphere, was sometimes a little over one-half a degree, Fahrenheit; though it was often considerably less than this. The different parts of the vegetable fabric, therefore, generate different quantities of caloric. In the same manner, the heat-pro- ducing power is not equally active in different species of animals ; but its existence is nevertheless common to both animals and vege- tables. With regard to the mode of generation of this internal or vital heat, we may start with the assertion that its production depends upon changes of a chemical nature, and is so far to be regarded as a chemical phenomenon. The sources of heat which we meet with in external nature are of various kinds. Sometimes the heat is of a physical origin ; as, for example, that derived from the rays of the sun, the friction of solid substances, or the passage of electric currents. In other instances it is produced by chemical changes ; and the most abundant and useful source of artificial heat is the oxidation, or combustion, of carbon and carbonaceous compounds. Wood and coal, substances rich in carbon, are mostly used for this purpose ; and charcoal, which is nearly pure carbon, is frequently employed by itself. These substances, when burnt, or oxidized, evolve a large amount of heat ; and produce, as the result of their oxidation, carbonic acid. In order that the process may go on, it is of course necessary that oxygen, or atmospheric air, should have free access to the burning body; otherwise the combustion and evolution of heat cease, for want of a necessary agent in the chemi- cal combination. In all these instances, the quantity of heat gene- rated is in direct proportion to the amount of oxidation ; and may be measured, either by the quantity of carbon consumed, or by that of carbonic acid produced. It may be made to go on, also, either rapidly or slowly, according to the abundance and purity in which oxygen is supplied to the carbonaceous substance. Thus, if char- coal be ignited in an atmosphere of pure oxygen, it burns rapidly and violently, raises the temperature to a high point, and is soon entirely consumed. On the other hand, if it be shut up in a close stove, to which the air is admitted but slowly, it produces only a slight elevation of temperature, and may require a much longer time for its complete disappearance. Nevertheless, for the same quantity of carbon consumed, the amount of heat generated, and ANIMAL HEAT. 223 that of carbonic acid produced, will be equal in the two cases. In one instance we have a rapid combustion, in the other a slow com- bustion ; the total effect being, however, the same in both. Such is the mode in which heat is commonly produced by artifi- cial means. Its evolution is here dependent upon two striking conditions, which are essential to it, and by which it is always accompanied, viz., the consumption of oxygen, and the production of carbonic acid. Now, since the two phenomena just mentioned are presented also by the living body, and since they are accompanied here, too, by the production of animal heat, it was very natural to suppose that in the animal organization, as well as elsewhere, the internal heat must be owing to an oxidation or combustion of carbon. According to Lavoisier, the oxygen taken into the lungs was sup- posed to combine immediately with the carbon of the pulmonary tissues and fluids, producing carbonic acid, and to be at once returned under that form to the atmosphere; the same quantity of heat result- ing from the above process as would have been produced by the oxidation of a similar quantity of carbon in wood or coal. Accord- ingly, he regarded the lungs as a sort of stove or furnace, by which the rest of the body was warmed, through the medium of the circu- lating blood. It was soon found, however, that this view was altogether erro- neous ; for the slightest examination shows that the lungs are not perceptibly warmer than the rest of the body ; and that the heat- producing power, whatever it may be, does not reside exclusively in the pulmonary tissue. Furthermore, subsequent investigations showed the following very important facts, which we have already mentioned, viz., that the carbonic acid is not formed in the lungs, but exists in the blood before its arrival in the pulmonary capilla- ries ; and that the oxygen of the inspired air, so far from combining with carbon in the lungs, is taken up in solution by the blood- globules, and carried away by the current of the general circulation. It is evident, therefore, that this oxidation or combustion of the blood must take place, if at all, not in the lungs, but in the capil- laries of the various organs and tissues of the body. Liebig accordingly adopted Lavoisier's theory of the production of animal heat, with the above modification. He believed the heat of the animal body to be produced by the oxidation or combustion of certain elements of the food while still circulating in the blood ; these substances being converted into carbonic acid and water by 224 ANIMAL HEAT. the oxidation of their carbon and hydrogen, and immediately ex- pelled from the body without ever having formed a part of the solid tissues. He therefore divided the food into two different classes of alimentary substances; viz., 1st, the nitrogenous or plastic elements^ which are introduced in comparatively small quantity, and which are to be actually converted into the substance of the tissues, such as albumen, muscular flesh, &c. ; and 2d, the hydro-carhons or respiratory elements, such as sugar, starch, and fat; which, according to his view, are taken into the blood solely to be burned, never being assimilated or converted into the tissues, but only oxidized in the circulation, and immediately expelled, as above, under the form of carbonic acid and water. He therefore regardf^d these elements of the food only as so much fuel ; destined simply to maintain the heat of the body, but taking no part in the proper function of nutrition. The above theory of animal heat has been very generally adopted and acknowledged by the medical profession until within a recent period. A few years ago, however, some of its deficiencies and inconsistencies were pointed out, by Lehmann in Germany, and by Eobin and Verdeil in France ; and since that time it has begun to lose ground and give place to a different mode of explanation, more in accordance with the present state of physiological science. We believe it, in fact, to be altogether erroneous; and incapable of explaining, in a satisfactory manner, the phenomena of animal heat, as exhibited by the living body. We shall now proceed to pass in review the principal objections to the theory of combustion, con- sidered as a physiological doctrine. I. It is not at all necessary to regard the evolution of heat as dependent solely on direct oxidation. This is only one of its sources, as we see constantly in external nature. The sun's rays, mechanical friction, electric currents, and more particularly a great variety of chemical actions, such as various saline combinations and decompositions, are all capable of producing heat ; and even simple solutions, such as the solution of caustic potass in water, the mixture of sulphuric acid and water, or of alcohol and water, will often pro- duce a very sensible elevation of temperature. Now we know that in the interior of the body a thousand different actions of this nature are constantly going on; solutions, combinations and decom- positions in endless variety, all of which, taken together, are amply sufficient to account for the production of animal heat, provided the theory of combustion should be found insufficient or improbable. ANIMAL HEAT. 225 II. In vegetables there is an internal production of heat, as well as in animals; a fact which has been fully demonstrated by the experiments of Dutrochet and. others, already described. In vege- tables, however, the absorption of oxygen and exhalation of car- bonic acid do not take place; excepting, to some extent, during the night. On the contrary, the diurnal process in vegetables, it is well known, is exactly the reverse of this. Under the influence of the solar light they absorb carbonic acid and exhale oxygen. And it is exceedingly remarkable that, in Dutrochet's experiments, he found that the evolution of heat by plants was always accompanied by the disappearance of carbonic acid and the exhalation of oxygen. Plants which, in the daylight, exhale oxygen and evolve heat, if placed in the dark, immediately begin to absorb oxygen and exhale carbonic acid ; and, at the same time, tbe evolution of heat is sus- pended. Dutrochet even found that the evolution of heat by plants presented a regular diurnal variation; and that its maximum of intensity was aboilt the middle of the day, just at the time when the absorption of carbonic acid and the exhalation of oxygen are going on with the greatest activity. The proper heat of plants, therefore, can- not be the result of oxidation or combustion, but must be dependent on an entirely different process. * III. In animals, the quantities of oxygen absorbed and of carbonic acid exhaled do not correspond with each other. Most frequently a certain amount of oxygen disappears in the body, over and above that which is returned in the breath under the form of carbonic acid. This overplus of oxygen has been said to unite with the hydrogen of the food, so as to form water which also passes out by the lungs ; but this is a pure assumption, resting on no direct evidence whatever, for we have no experimental proof that any more watery vapor is exhaled from the lungs than is supplied by the fluids taken into the stomach. It is superfluous, therefore, to assume that any of it is produced by the oxidation of hydrogen. Furthermore, the proportion of overplus oxygen which disap- pears in the body, beside that which is exhaled in the carbonic acid of the breath, varies greatly in the same animal according to the quality of the food. Eegnault and Eeiset' found that in dogs, fed on meat, the oxygen which reappeared under the form of carbonic acid was only 75 per cent, of the whole quantity absorbed ; while ' Aiinales de Chiiiiie et de Physique, 3d series, xxvi. p. 428. 15 226 AXIMAL HEAT. in dogs fed on vegetable substances it amounted to over 90 per cent. In some instances/ where the animals (rabbits and fowls) were fed on bread and grain exclusively, the proportion of expired oxygen amounted to 101 or even 102 per cent.; that is, more oxygen luas actually contaiyied in the carbonic acid exhaled, than had been ab- sorbed in a free state from the atmosphere. A portion, at least, of the carbonic acid must therefore have been produced by other means than direct oxidation. TV. It has already been shown, in a previous chapter, that the carbonic acid which is exhaled from the lungs is not primarily formed in the blood, but makes its appearance in the substance of the tissues themselves ; and furthermore, that even here it does not originate by a direct oxidation, but rather by a process of decom- position, similar to that by which sugar, in fermentation, is resolved into alcohol and carbonic acid. We understand from this how to explain the singular fact alluded to in the last paragraph, viz., the abundant production of carbonic acid, under some circumstances, with a comparatively small supply of free oxygen. The statement made by Liebig, therefore, that starchy and oily matters taken with the food are immediately oxidized in the circulation without ever being assimilated by the tissues, is without foundation. It never, in fact, rested on any other ground than a supposed probability; and as we see that carbonic acid is abundantly produced in the body by other means, we have no longer any reason for assuming, without direct evidence, the existence of a combustive process in the blood. V. The evolution of heat in the animal body is not general, as it would be if it resulted from a combustion of the blood ; but local, since it takes place primarily in the substance of the tissues them- selves. Various causes will therefore produce a local elevation or depression of temperature, by modifying the nutritive changes which take place in the tissues. Thus, in the celebrated experiment of Bernard, which we have often verified, division of the sympa- thetic nerve in the middle of the neck produces very soon a marked elevation of temperature in the corresponding side of the head and face. Local inflammations, also, increase very sensibly the tempera- ture of the part in which they are seated, while that of the general ' Annales de Chimie et de Physique, 3d series, xxvi. pp. 409 — 451. ANIMAL HEAT. 227 mass of the blood is not altered. Finally it has been demonstrated by Bernard that in the natural state of the system there is a marked difference in the temperature of the dififerent organs and of the blood returning from them.* The method adopted by this experimenter was to introduce, in the living animal, the bulb of a fine thermo- meter successively into the bloodvessels entering and those leaving the various internal organs. The difference of temperature in these two situations showed whether the blood had lost or gained in heat Avhile traversing the capillaries of the organ. Bernard found, in the first place, that the blood in passing through the lungs, so far from increasing, was absolutely diminished in temperature; the blood on the left side of the heart being sometimes a little more and sometimes a little less than one-third of a degree Fahr. lower than on the right side. This slight cooling of the blood in the lungs is owing simply to its exposure to the air through the pul- monary membrane, and to the vaporization of water which takes place in these organs. In the abdominal viscera, on the contrary, the blood is increased in temperature. It is sensibly warmer in the portal vein than in the aorta ; and very considerably warmer in the hepatic vein than in either the portal or the vena cava. The blood of the hepatic vein is in fact warmer than that of any other part of the body. The production of heat, therefore, according to Ber- nard's observations, is more active in the liver than in any other portion of the system. As the chemical processes of nutrition are necessarily different in the different tissues and organs, it is easy to understand why a specific amount of heat should be produced in each of them. A similar fact, it will be recollected, was noticed by Dutrochet, in regard to the different parts of the vegetable organ- ization. VI. Animal heat has been supposed to stand in a special relation to the production of carbonic acid, because in warm-blooded animals the respiratory process is more active than in those of a lower temperature; and because, in the same animal, an increase or diminution in the evolution of heat is accompanied by a corre- sponding increase or diminution in the products of respiration. But this is also true of all the other excretory products of the body. An elevation of temperature is accompanied by an increased activity of all the nutritive processes. Not only carbonic acid, but the ' Gazette Hebdomadaiie, Aug. 29 and Sept. 26, 1856. 228 ANIMAL HEAT. ingredients of the urine and the perspiration are discharged in larger quantity than usual. An increased supply of food also is required, as well as a larger quantity of oxygen; and the digestive and secretory processes both go on, at the same time, with unusual activity. Animal heat, then, is a phenomenon which results from the simultaneous activity of many different processes, taking place in many different organs, and dependent, undoubtedly, on different chemical changes in each one. The introduction of oxygen and the exhalation of carbonic acid have no direct connection with each other, but are only the beginning and the end of a long series of continuous changes, in which all the tissues of the body successively take a part. Their relation is precisely that which exists between the food introduced through the stomach, and the urinary ingre- dients eliminated by the kidneys. The tissues require for their nutrition a constant supply of solid and liquid food which is intro- duced through the stomach, and of oxygen which is introduced through the lungs. The disintegration and decomposition of the tissues give rise, on the one hand, to urea, uric acid, &c., which are discharged with the urine, and on the other hand to carbonic acid, which is exhaled from the lungs. But the oxygen is not directly converted into carbonic acid, any more than the food is directly converted into urea and the urates. Animal heat is not to be regarded, therefore, as the result of a combustive process. There is no reason for believing that the greater part of the food is " burned" in the circulation. It is, on the contrary, assimilated by the substance of the tissues; and these, in their subsequent disintegration, give rise to several excretory products, one of which is carbonic acid. The numerous combinations and decompositions which follow each other incessantly during the nutritive process, result in the production of an internal or vital heat, which is present in both animals and vegetables, and which varies in amount in different species, in the same individual at different times, and even in different parts and organs of the same body. THE CIRCULATION". 229 CHAPTER XIV. THE CIRCULATION. The blood may be regarded as a nutritious fluid, holding in solution all the ingredients necessary for the formation of the tissues. In some animals and vegetables, of the lowest organization, such as infusoria, polypes, algte, and the like, neither blood nor circulation is required ; since all parts of the body, having a similar structure, absorb nourishment equally from the surrounding media, and carry on nearly or quite the same chemical processes of growth and assimilation. In the higher animals and vegetables, however, as well as in the human subject, the case is different. In them, the structure of the body is compound. Different organs, with widely different functions, are situated in different parts of the frame ; and each of these functions is more or less essential to the continued existence of the whole. In the intestine, for example, the process of digestion takes place ; and the prepared ingredients of the food are thence absorbed into the bloodvessels, by which they are transported to distant tissues and organs. In the lungs, again, the blood absorbs oxygen which is afterward to be appropriated by the tissues; and carbonic acid, which was produced in the tissues, is exhaled from the lungs. In the liver, the kidneys, and the skin, other substances again are produced or eliminated, and these local processes are all of them necessary to the preservation of the general organization. The circulating fluid is therefore, in the higher animals, a means of transportation, by which the substances pror duced in particular organs are dispersed throughout the body, or by which substances produced generally in the tissues are conveyed to particular organs, in order to be eliminated and expelled. The circulatory apparatus consists of four different parts, viz: 1st. The heart ; a hollow, muscular organ, which receives the blood at one orifice and drives it out, in successive impulses, at another. 2d. The arteries ; a series of branching tubes, which convey the blood from the heart to the different tissues and organs of the body. 230 THE CIRCULATION'. 3d. The capillaries; a network of minute inosculating tubules, whicli are interwoven with the substance of the tissues, and which bring the blood into intimate contact with the cells and fibres of which thej are composed; and, 4th. The veins; a set of converg- ing vessels, destined to collect the blood from the capillaries, and return it to the heart. In each of these four different parts of the circulatory apparatus, the movement of the blood is peculiar and dependent on special conditions. It will therefore require to be studied in each one of them separately. THE HEART. The structure of the heart, and of the large vessels connected with it, varies considerably in different classes of animals, owing to the different arrangement of the respiratory organs. For the respi- ratory apparatus being one of the most important in the body, and the one most closely connected •^'S* '^^' by anatomical relations with the organs of circulation, the latter are necessarily modified in structure to correspond with the former. In fish, for exam- ple (Fig. 76), the heart is an organ consisting of two princi- pal cavities : an auricle (a) into which the blood is received from the central extremity of the vena cava, and a ventricle (b) into which the blood is driven by the contraction of the auricle. The ventricle is considerably larger and more powerful than the auricle, and by its contrac- tion drives the blood into the main artery supplying the gills. In the gills (cc) the blood is arterialized ; after which it is collected by the branchial veins. These veins unite upon the median line to form the aorta {d) by which the blood is finally distributed throughout the frame. In Circulation- of Fish. — a. Auricle, b. Ventricle, cc. Gills, d. Aorta, ee. Vense carse. THE HEART. 231 Fig. 77. these animals the respiratory process is not a very active one; but the gills, which are of small size, being the only respiratory organs, all the blood requires to pass through them for purposes of aeration. The heart here is a single organ, destined only to drive the blood from the termination of the venous system to the capillaries of the gills. In reptiles, the heart is composed of two auricles, placed side by side, and one ventricle. (B'ig. 77.) The venas cavas discharge their blood into the right auricle (a), whence it passes into the ventricle (c). From the ventricle, a part of it is carried into the aorta and distri- buted throughout the body, while a part is sent to the lungs through the pulmonary artery. The arterialized blood, returning from the lungs by the pulmonary vein, is discharged into the left auricle (6), and thence into the ventricle (c), where it mingles with the venous blood which has just arrived by the venae cavae. In the reptile, therefore, the ventricle is a common organ of pro- pulsion, both for the lungs and for the general circulation. In these animals the aeration of the blood in the lungs is only partial ; a certain portion of the blood which leaves the heart being carried to these organs, just as in the human subject, it is only a portion of the blood which is carried to the kidney by the renal artery. This arrangement is sufficient for the reptiles, because in many of them, such as serpents and turtles, the lungs are much more extensive and efficient, as respiratory organs, than the gills of fish ; while in others, such as frogs and water-lizards, the integument itself, which is moist, smooth, and naked, takes an important share in the aeration of the blood. In quadrupeds and the human species, however, 'the respi- ratory process is not only exceedingly active, but the lungs are, at the same time, the only organs in which the aeration of the blood can be fully accomplished. In them, accordingly, we find the two circulations, general and pulmonary, entirely dis- ClRCITLATIOX OF REPTILES. — a. Right auricle. 6. Left auricle, c. Ventricle. d. Lungs, e. Aorta, f. Vena Cava. 232 THE CIRCULATION. tinct from each other. (Fig. 78.) All the blood returning from the body by the veins must pass through the lungs before it is again distributed through the Fig- 78. arterial system. We have therefore a double circula- tion, and also a double heart ; the two sides of which, though united externally, are separate internally. The mammalian heart consists of a right auricle and ventricle ((7, h), receiving the blood from the vena cava (?"), and driving it to the lungs ; and a left auricle and ventricle (/, g) receiving the blood from the lungs and driving it outward through the arte- rial system. In the complete or double mammalian heart, the differ- ent parts of the organ present certain peculiarities and bear certain relations to each other, which it is necessary to understand before we can properly appreciate its action and movements. The entire organ has a more or less conical form, its base being situated on the median line, directed upward and backward ; the whole being suspended in the chest, and loosely fixed to the spinal column, by the great vessels which enter and leave it at this point. The apex, on the contrary, is directed downward, forward, and to the left, sur- rounded by the pericardium and the pericardial fluid, but capable of a very free lateral and rotatory motion. The auricles, which have a smaller capacity and thinner walls^ than the ventricles, are situated at the upper and posterior part of the organ (Figs. 79 and 80); while the ventricles occupy its anterior and lower portions. The two ventricles, moreover, are not situated on the same plane, but the right ventricle occupies a position somewhat in front and above that of the left ; so that in an anterior view of the heart the greater portion of the left ventricle is concealed by the right (Fig. 79), and in a posterior view the greater portion of the right ven- tricle is concealed by the left (Fig. 80); while in both positions the Circulation in Mammalians. — a. Right auricle. 6. Eight ventricle, c. Pulmonary artery. d. Lungs, e. Pulmonary vein. / Left auricle, g. Left ventricle, h. Aorta, i. Vena cava. THE HEART. 233 apex of tlie heart is constituted altogether by the point of the left ventricle. Fig. 80. Human Heart, anterior view. — a. Right ventricle, b. Left ventricle. c. Right auricle, d. Left auricle, e. Pulmonary artery. /. Aorta. Human Heart, posterior view. — a. Right ventricle, b. Left ventricle. c. Right auricle, d. Left auricle. The different cavities of the heart and of the adjacent blood- vessels, though continuous with each other, are partially separated by certain constrictions. These constricted orifices, by which the different cavities communicate, are known by the names of the Fig, 81. RiOHT Auricle and Ventricle; Auriculo-ventricular Valves open. Arterial Valves closed. auricular, auriculo-ventricular, and aortic and pulmonary orifices; the auricular orifices being the passages from the vense cavas and 234 THE CIRCULATION. pulmonary veins into the right and left auricles; the auriculo- ventricular orifices leading from the auricles into the ventricles; and the aortic and pulmonary orifices leading from the ventricles into the aortic and pulmonary arteries respectively. The auriculo-ventricular, aortic, and pulmonary orifices are fur- nished with valves, which allow the blood to pass readily from the auricles to the ventricles, and from the ventricles to the arteries, but shut back with the contractions of the organ, so as to prevent its return in an opposite direction. The course of the blood through the heart is, therefore, as follows. From the vena cava it passes into the right auricle; and from the right auricle into the right ventricle. (Fig. 81.) On the contraction of the right ventricle, the tricuspid valves shut back, preventing its return into the auricle (Fig. 82); and it is thus driven through the pulmonary artery to the Fig. 82. Eight Auricle and Ventricle; Auriculo-ventricular Valves closed, Arterial Valves open. lungs. Returning from the lungs, it enters the left auricle, thence passes into the left ventricle, from which it is finally delivered into the aorta, and distributed throughout the body. (Fig. 83.) This movement of the blood, however, through the cardiac cavities, is not a continuous and steady flow, but is accomplished by alternate contractions and relaxations of the muscular parietes-of the heart; so that with every impulse, successive portions of blood are received by the auricles, delivered into the ventricles, and by them dis- THE HEART. 235 charged into the arteries. Each one of these successive actions is called a beat, or pulsation of the heart. Fig. 83. Course op Blood through the Heart. — a, a. Vena cava, superior and inferior. b. Right ventricle, c. Pulmonary artery, d. Pulmonary vein. e. Left ventricle. /. Aorta. Each pulsation of the heart is accompanied by certain important phenomena, which require to be studied in detail. These are the sounds, the movements, and the impulse. The sounds of the heart are two in number. They can readily be heard by applying the ear ov,er the cardiac region, when they are found to be quite different from each other in position, in tone, and in duration. They are distinguished as the first and second sounds of the heart. The first sound is heard with the greatest intensity over the anterior surface of the heart, and more particularly over the fifth rib and the fifth intercostal space. It is long, dull, and smothered in tone, and occupies one-half the entire duration of a single beat. It corresponds in time with the impulse of the heart in the precordial region, and the stroke of the large arteries in the immediate vicinity of the chest. The second sound follows imme- diately upon the first. It is heard most distinctly at the situation of the aortic and pulmonary valves, viz., over the sternum at the level of the third costal cartilage. It is short, sharp, and distinct in tone, and occupies only about one-quarter of the whole time of 236 THE CIRCULATION. a pulsation. It is followed by an equal interval of silence ; after wliich the first sound again recurs. The whole time of a cardiac pulsation may then be divided into four quarters, of which the first two are occupied by the first sound, the third by the second sound, and the fourth by an interval of silence, as follows : — Time of pulsation. ' 1st quarter \ n , „ V First sound. 3d " Second sound. 4th " Interval of silence. The cause of the second sound is universally acknowledged to be the sudden closure and tension of the aortic and pulmonary valves. This fact is established by the following proofs : 1st, this sound is heard with perfect distinctness, as we have already mentioned, di- rectly over the situation of the above-mentioned valves; 2d, the far- ther we recede in any direction from this point, the fainter becomes the sound ; and 3d, in experiments upon the living animal, often repeated by different observers, it has been found that if a curved needle be introduced into the base of the large vessels, so as to hook back the semilunar valves, the second sound at once disappears, and remains absent until the valve is again liberated. These valves consist of fibrous sheets, covered with a layer of endocardial epithe- lium. They have the form of semilunar festoons, the free edge of which is directed away from the cavity of the ventricle, while the attached edge is fastened to the inner surface of the base of the artery. While the blood is passing from the ventricle to the artery, these valves are thrown forward and relaxed ; but when the artery reacts upon its contents they shut back, and their fibres, becoming suddenly tense, yield a clear, characteristic, snapping sound. The production of the first sound has been attributed by some writers to a combination of various causes ; such as the rush of blood through the cardiac orifices, the muscular contraction of the parietes of the heart, the tension of the auriculo-ventricular valves, the collision of the particles of blood with each other and with the surface of the ventricle, &c. &c. We believe, however, with Andry' and some others, that the first sound of the heart has a similar origin with the second; and that it is dependent altogether on the closure of the auriculo-ventricular valves. The reasons for this con- clusion are the following : — 1st. The second sound is undoubtedly caused by the closure of ' Diseases of the Heart, Kneeland's translation, Boston, 1846. THE HEART. 237 the semilunar valves, and in the action of the heart the shutting back of the two sets of valves alternate with each other precisely as do the first and second sounds; and there is every probability, to say the least, that the sudden tension of the valvular fibres pro- duces a similar effect in each instance. 2d. The first sound is heard most distinctly over the anterior surface of the ventricles, where the tendinous cords supporting the auriculo-ventricular valves are inserted, and where the sound pro- duced by the tension of these valves would be most readily con- ducted to the ear. 8d. There is no reason to believe that the current of blood through the cardiac orifices could give rise to an appreciable sound, so long as these orifices, and the cavities to which they lead, have their normal dimensions. An unnatural souffle may indeed origi- nate from this cause when the orifices of the heart are diminished in size, as by. calcareous or fibrinous deposits; and it may also occur in cases of aneurism. A souffle may even be produced at will in any one of the large arteries by pressing firmly upon it with the end of a stethoscope, so as to diminish its calibre. But in all these instances, the abnormal sound occurs only in consequence of a disturbance in the natural relation existing between the volume of the blood and the size of the orifice through which it passes. In the healthy heart, tbe different orifices of the organ are in exact proportion to the quantity of the circulating blood ; and there is no more reason for believing that its passage should give rise to a sound in the cardiac cavities than in the larger arteries or veins. 4:th. The difference in character between the two sounds of the heart depends, in all probability, on the different arrangement of the two sets of valves. The second sound is short, sharp, and dis- tinct, because the semilunar valves are short and narrow, superficial in their situation, and supported by the highly elastic, dense and fibrous bases of the aortic and pulmonary arteries. The first sound is dull and prolonged, because the auriculo-ventricular valves are broad and deep-seated, and are attached, by their long chordas tendinese to the comparatively soft and yielding fleshy columns of the heart. The difference between the first and second sounds can, in fact, be easily imitated, by simply snapping between the fingers two pieces of tape or ribbon, of the same texture but of different lengths. (Fig. 84.) The short one will give out a distinct and sharp sound ; the long one a comparatively dull and prolonged sound. Together with the first sound of the heart there is also to be 238 THE CIRCULATION. heard a sW^i friction sound, produced by the collision of the point of the heart against the parietes of the chest. This sound, which is heard in the fifth intercostal space, is very faint, and is more or less Fig. 84. masked by the strong valvular sound which occurs at the same time. It is different, however, in character from the latter, and may usually be distinguished from it by careful examination. The movements of the heart during the time of a pulsation are of a peculiar character, and have been very often erroneously described. In fact altogether the best description of the move- ments of the heart which has yet appeared, is that given by Wil- liam Harvey, in his celebrated work on the Motion of the Heart and Blood, published in 1628. He examined the motion of the heart by opening the chest of the living animal; and though the same or similar experiments have been frequently performed since his time, the descriptions given by subsequent observers have been for the most part singularly inferior to his, both in clearness and fidelity. The method which we have adopted for examining the motions of the heart in the dog is as follows: The animal is first rendered insensible by ether, or by the inoculation of woorara. The latter mode is preferable, since a long-continued etherization seems to exert a sensibly depressing effect on the heart's action, which is not the case with woorara. The trachea is then exposed and opened just below the larj'nx, and the nozzle of a bellows inserted and secured by ligature. Finally, the chest is opened on the me- dian line, its two sides widely separated, so as to expose the heart and lungs, the pericardium slit up and, carefull}^ cut away from its attachments, and the lungs inflated by insufflation through the trachea. By keeping up a steady artificial respiration, the move- THE HEART. 239 ments of the heart may be made to continue, in favorable cases, for more than an hour; and its actions may be studied by direct obser- vation, like those of any external organ. The examination, however, requires to be conducted with certain precautions, which are indispensable to success. When the heart is first exposed, its movements are so complicated, and recur with such rapidity, that it is difficult to distinguish them perfectly from each other, and to avoid a certain degree of confusion. Singular as it may seem, it is even difficult at first to determine what period in the heart's pulsation corresponds to contraction, and what to relaxation of the organ. We have even seen several medical men, watching together the pulsations of the same heart, unable to agree upon this point. It is very evident, indeed, that several English and continental observers have mistaken, in their examinations, the contraction for the relaxation, and the relaxation for the contrac- tion. The first point, therefore, which it is necessary to decide, in examining the successive movements of a cardiac pulsation, is the following, viz: Which is the contraction and which the relaxation of the ventricles? The method which we have adopted is to pass a small silver canula directly through the parietes of the left ven- tricle into its cavity. The blood is then driven from the external orifice of the canula in interrupted jets; each jet indicating the time at which the ventricle contracts upon its contents. The canula is then withdrawn, and the difierent muscular layers of the ventricular walls, crossing each other obliquely, close the opening, so that there is little or no subsequent hemorrhage. When the successive actions of contraction and relaxation have by this means been fairly recognized and distinguished from each other, the cardiac pulsations are seen to be characterized by the following phenomena. The changes in form and position of the entire heart are mainly dependent on those of the ventricles, which contract simultaneously with each other, and which constitute much the largest portion of the entire mass of the organ. 1. At the time of its contraction the heart hardens. This pheno- menon is exceedingly well marked, and is easily appreciated by placing the finger upon the ventricles, or by grasping them between the finger and thumb. The muscular fibres become swollen and indurated, and if grasped by the hand communicate the sensation of a somewhat sudden and powerful shock. It is this forcible indu- ration of the heart, at the time of contraction, which has been mis- taken by some writers for an active dilatation, and described as 240 THE CIRCULATION. such. It is, however, a phenomenon precisely similar to that which takes place in the contraction of a voluntary muscle, which be- comes swollen and indurated at the same moment and in the same proportion that it diminishes in length, 2. At the time of contraction, the ventricles elongate and the point of the heart protrudes. This phenomenon was very well described by Dr. Harvey.^ "The heart," he says, "is erected, and rises upward to a point, so that at this time it strikes against the breast and the pulse is felt externally." The elongation of the ventricles during contraction has, however, been frequently denied by subsequent writers. The only modern observers, so far as we are aware, who have recognized its existence, are Drs. C. W. Pen- nock and Edward M. Moore, who performed a series of very careful and interesting experiments on the action of the heart, in Philadel- phia, in the year 1839.^ These experimenters operated upon calves, sheep, and horses, by stunning the animal with a blow upon the head, opening the chest, and keeping up artificial respiration. They observed an elongation of the ventricle at the time of contraction, and were even able to measure its extent by applying a shoemaker's rule to the heart while in active motion. We are able to corroborate entirely the statement of these observers by the result of our own experiments on dogs, rabbits, frogs, &c. The ventricular contrac- tion is an active movement, the relaxation entirely a passive one. When contraction occurs and a stream of blood is thrown out of the ventricle, its sides approximate each other and its point elon- gates ; so that the transverse diameter of the heart is diminished, and its longitudinal diameter increased. This can be readily felt by grasping the base of the heart and the origin of the large vessels gently between the first and middle fingers, and allowing the end of the thumb of the same hand to rest lightly upon its apex. With every contraction the thumb is sensibly lifted and separated from the fingers, by a somewhat forcible elevatiou.of the point of the heart. The same thing can be seen, and even measured by the eye, in the following manner : If the heart of the frog or even of any small warm-blooded animal, as the rabbit, be rapidly removed from the chest, it will continue to beat for some minutes afterward; and when the rhythmical pulsations have finally ceased, contractions ' Works of William Harvey, M. D. Sydenharii ed., Loiulon, 1847, p. 21. ' Philadelphia Medical Kxiuiiiuer, No. 44. THE HEAET. 241 can still be readily excited by toucliing the heart with the point of a steel needle. If the heart be now held by its base between the thumb and finger, with its point directed upward, it will be seen to have a pyramidal or conical form, representing very nearly in its outline an equilateral triangle (Fig. 85) ; its base, while in a condition of rest, bulging out laterally, while the apex is compara- tively obtuse. Fig. 85. Fig. 86. Heart of Froq in a state of relaxa- tion. Heart of Frog iu contiaction. When the heart, held in this position, is touched with the point of a needle (Fig. 86), it starts up, becomes instantly narrower and longer, its sides approximating and its point rising to an acute ancle. This contraction is immediately followed by a relaxation ; the point of the heart sinks down, and its sides again bulge out- ward. Let us now see in what manner this change in the figure of the ventricles during contraction is produced. If the muscular fibres of the heart were arranged in the form of simple loops, running parallel with the axis of the organ, the contraction of these fibres would merely have the effect of di- minishing the size of the heart in every direction. This effect can be seen in the accompanying hypothetical diagram (Fig. 87), where the white outline represents such simple looped fibres in a state of re- laxation, and the dotted internal line indi- cates the form which they would take in contraction. In point of fact, however, none of the muscular fibres of the heart run parallel to its longitudinal axis. They are disposed, on the contrary, in a direction partly spiral and partly circular. The most superficial fibres start from the base of the ventricles, and pass 16 Diagram of Simple Looped Fibres, in relaxation and con- traction. 242 THE CIRCULATION. toward the apex, curling round the heart in such a manner as to pass over its anterior surface in an obliquely spiral direction, from above downward, and from right to left. (Fig. 88.) They converge toward the point of the heart, curl- ing round the centre of its apex, and then, changing their direction, be- come deep-seated, run upward along Fig. 89. Fig. 88. Bullock's Heart, anterior view, showing the superficial muscular litres. Left Ventricle op BdLLOCK's Heart, show- ing the deep fibres. the septum and internal surface of the ventricles, and terminate in the columnae carnese, and in the inner border of the auriculo- ventricular ring. The deeper layers of fibres, on the contrary, are wrapped round the ventricles in a nearly circular direction (Fig. 89); their points of origin and attachment being still the auriculo- ventricular ring, and the points of the fleshy columns. The entire arrangement of the muscular bundles may be readily seen in a heart which has been boiled for six or eight hours, so as to soften the connecting areolar tissue, and enable the fibrous layers to be easily separated from each other. By far the greater part of the mass of the fibres have therefore a circular instead of a longitudinal direction. When they contract, their action tends to draw the lateral walls of the ventricles together, and thus to diminish the transverse diameter of the heart ; but as each muscular fibre becomes thickened in direct proportion to its contraction, their combined lateral swelling necessarily pushes out the apex of the ventricle, and the heart elongates at the same time that its sides are drawn together. This effect is illustrated in the accompanying diagram (Fig. 90), where the white lines show the ficrure of the heart durinor relaxation, with the course of its circular THE HEART. 243 Diagram of Circular Fibres OF THE Heart, and their con- traction. fibres, while the dotted line shows the narrowed and elongated figure necessarily produced by their contraction. This phenomenon, therefore, of the protrusion of the apex of the heart at the time of contraction, is not only fully established by observation, but is readily explained by the anatomical structure of the organ. 3. Simultaneously with the hardening and elongation of the heart, its apex moves slightly from left to right, and rotates also upon its own axis in the same direction. Both these movements result from the peculiar spiral arrangement of the cardiac fibres. If we refer again to the preceding diagrams, we shall see that, provided the fibres were arranged in simple longitudi- nal loops (Fig. 87), their contraction would merely have the efl[ect of drawing the point of the heart directly upward in a straight line toward its base. On the other hand, if they were arranged altogether in a circular direction (Fig. 90), the apex would be simply protruded forward, also in a direct line, without deviating or twisting either to the right or to the left. But iu point of fact, the superficial fibres, as we have already described, run spirally, and curling round the point of the heart, turn inward toward its base ; so that if the apex of the organ be viewed externally, it will be seen that the superficial fibres converge toward its cen- tral point in curved lines, as in Fig. 91. It is well known that every curved muscular fibre, at the time of its shortening, necessa- rily approximates more or less to a straight line. Its curvature is diminished in exact proportion to the extent of its contraction; and if arranged in a spiral form, its contraction tends in the same degree to untwist the spiral. During the con- traction of the heart, therefore, its apex rotates on its own axis in the direction indicated by the arrows in Fig. 91, viz., from left to right anteriorly, and from right to left posteriorly. This produces a twisting movement of the apex in the above direction, which is Fig. 91. Converging Fibres or THE Apex of the Heart. 244 THE CIRCULATION. very perceptible to the eye at every pulsation of the heart, when exposed in the living animal. 4. The protrusion of the point of the heart at the time of con- traction, together with its rotation upon its axis from left to right, brings the apex of the organ in contact with the parietes of the chest, and produces the shock or impulse of the heart, which is readily perceptible externally, both to the eye and to the touch. In the human subject, when in an erect position, the heart strikes the chest in the fifth intercostal space, midway between the edge of the sternum and a line drawn perpendicularly downward from the left nipple. In a supine position of the body, the heart falls away from the anterior parietes of the chest so much that the impulse may disappear for the time altogether. This alternate recession and advance of the point of the heart, in relaxation and contraction, is provided for by the anatomical arrangement of the pericardium, and the existence of the pericardial fluid. As the heart plays back- ward and forward, the pericardial fluid constantly follows its movements, receding as the heart advances, and advancing as the heart recedes. It fulfils, in this respect, the same purpose as the synovial fluid, and the folds of adipose tissue in the cavity of the large articulations ; and allows the cardiac movements to take place to their full extent without disturbing or injuring in any way the adjacent organs. 5. The rhythm of the heart's pulsations is peculiar and somewhat complicated. Each pulsation is made up of a double series of con- tractions and relaxations. The two auricles contract together, and afterward the two ventricles ; and in each case the contraction is immediately followed by a relaxation. The auricular contraction is short and feeble, and occupies the first part of the time of a pulsation. The ventricular contraction is longer and more powerful, and occupies the latter part of the same period. Following the ventricular contraction there comes a short interval of repose, after which the auricular contraction again recurs. The auricular and ventricular contractions, however, do not alternate so distinctly with each other (like the strokes of the two pistons of a fire engine) as we should be led to believe from the accounts which have been given by some observers. On the contrary, they are connected and continuous. The contraction, which commences at the auricle, is immediately propagated to the ventricle, and runs rapidly from the base of the heart to its apex, very much in the manner of a peris- taUic motion, except that it is more sudden and vigorous. THE HEAKT. 245 William Harvey, again, gives a better account of this part of the heart's action than has been published by any subsequent writer. The following exceedingly graphic and appropriate description, taken from his book, shows that he derived his knowledge, not from any secondary or hypothetical sources, but from direct and careful study of the phenomena in the living animal. " First of all," he says,' " the auricle contracts, and in the course of its contraction throws the blood (which it contains in ample quantity as the head of the veins, the storehouse and cistern of the blood) into the ventricle, which being filled, the heart raises itself straightway, makes all its fibres tense, contracts the ventricles, and performs a beat, by which beat it immediately sends the blood supplied to it by the auricle, into the arteries; the right ventricle sending its charge into the lungs by the vessel which is called vena arteriosa, but which, in structure and function, and all things else, is an artery ; the left ventricle sending its charge into the aorta, and through this by the arteries to the body at large. " These two motions, one of the ventricles, another of the auricles, take place consecutively, but in such a manner that there is a kind of harmony or rhythm preserved between them, the two concurring in such wise that but one motion is apparent, especially in the warmer blooded animals, in which the movements in question are rapid. Nor is this for any other reason than it is in a piece of machinery, in which, though one wheel gives motion to another, yet all the wheels seem to move simultaneously ; or in that mechanical contrivance which is adapted to fire-arms, where the trigger being touched, down comes the flint, strikes against the steel, elicits a spark, which falling among the powder, it is ignited, upon which the flame extends, enters the barrel, causes the explo- sion, propels the ball, and the mark is attained ; all of which incidents, by reason of the celerity with which they happen, seem to take place in the twinkling of an eye." The above description indicates precisely the manner in which the contraction of the ventricle follows successively and yet con- tinuously upon that of the auricle. The entire action of the auricles and ventricles during a pulsation is accordingly as follows : The contraction begins, as we have already stated, at the auricle. Thence it runs immediately forward to the apex of the heart. The entire ventricle contracts vigorously, its walls harden, its apex • Op. cit.,p. 31. 246 THE CIRCULATION. protrudes, strikes against the walls of the chest, and twists from left to right, the auriculo- ventricular valves shut back, the first sound is produced, and the blood is driven into the aorta and pulmonary artery. These phenomena occupy about one-half the time of an entire pulsation. Then the ventricle is immediately relaxed, and a short period of repose ensues. During this period the blood flows in a steady stream from the large veins into the auricle, and through the auriculo-ventricular orifice into the ven- tricle ; filling the ventricle, by a kind of passive dilatation, about two-thirds or three-quarters full. Then the auricle contracts with a quick sharp motion, forces the last drop of blood into the ventricle, distending it to its full capacity, and then the ventricular contraction follows, as above described, driving the blood into the large arteries. These movements of contraction and relaxation continue to alternate with each other, and form, by their recurrence, the successive cardiac pulsations. THE ARTERIES AND THE ARTERLAL CIRCULATION. The arteries are a series of branching tubes which commence with the aorta and ramify throughout the body, distributing the blood to all the vascular organs. They are composed of three coats, viz : an internal homogeneous tunic, continuous with the endocardium; a middle coat, composed of elastic and muscular fibres ; and an external or " cellular" coat, composed of condensed layers of areolar tissue. The essential anatomical difference be- tween tbe larger and the smaller arteries consists in the structure of their middle coat. In the smaller arteries this coat is composed exclusively of smooth muscular fibres, arranged in a circular manner around the vessel, like the circular fibres of the muscular coat of the intestine. In arteries of medium size the middle coat contains both muscular and elastic fibres; while in those of the largest calibre it consists of elastic tissue alone. The large arteries, accordingly, possess a remarkable degree of elasticity and little or no contractility ; while the smaller are contractile, and but little or not at all elastic. It is found, by measuring the diameters of the successive arte- rial ramifications, that the combined area of all the branches given off' from a trunk is somewhat greater than that of the original THE ARTERIES AND THE ARTERIAL CIRCULATION. 247 vessel; and therefore that the combined area of all the small arteries must be considerably larger than that of the aorta, from which the arterial system originates. As the blood, consequently, in its passage from the heart outward, flows successively through larger and larger spaces, the rapidity of its circulation must neces- sarily be diminished, in the same proportion as it recedes from the heart. It is driven rapidly through the larger trunks, more slowly through those of medium size, and more slowly still as it approaches the termination of the arterial system and the commencement of the capillaries. The movement of the blood through the arteries is primarily caused by the contractions of the heart ; but is, at the same time, regulated and modified by the elasticity of the vessels. The mode in which the arterial circulation takes place is as follows. At the time of the heart's contraction, the muscular walls of the ventricle act power- fully upon its fluid contents. The auriculo- ventricular valves at the same time shutting back and preventing the blood from regur- gitating into the auricle, it is forced out through the aortic orifice. A charge of blood is therefore driven into that part of the aorta nearest the heart, by which the artery is distended in exact propor- tion to the force of the heart's action and the quantity of blood which it expels. When the ventricle relaxes, the distending force is removed ; and the elastic arterial walls, reacting upon their con- tents, would force the blood back again into the heart, were it not for the semilunar valves which shut together and close the aortic orifice. The column of blood is accordingly forced onward, into the next division of the arterial system, which is distended in its tufn and reacts again upon the blood, driving the blood necessarily farther and farther from the heart, until it arrives at the confines of the capillary system. In this manner a succession of waves or im- pulses is propagated from the heart outward (Fig. 92), consisting of the alternate distension and reaction of different portions of the artery, and which is readily perceptible whenever the vessel occu- pies a superficial position. This phenomenon is known by the name of the arterial 'pulse. When the blood is thus driven by the cardiac pulsations into the artery, the vessel is not only "distended laterally, but is elongated as well as widened, and enlarged in every direction. Particularly when the vessel takes a curved or serpentine course, its elongation and the increase of its curvatures may be observed at every pulsa- 248 THE CIECULATIOISr. tion. This may be seen, for example, in the temporal arteries, or even in the radial arteries, in emaciated persons. It is also very well seen in the mesenteric arteries, when the abdomen is opened Fig. 92. Diagram of Arterial Circulation. in the living animal. At every contraction of the heart the curves of the artery on each side become more strongly pronounced. (Fig. 93.) The vessel even rises up partially out of its bed, particularly where it runs over a bony sur- face, as in the case of the radial artery. In old persons the curves of the vessels become perma- nently enlarged from frequent distension ; and all the arteries tend to assume, with the advance of age, a more serpentine and even spiral course. Owing to the alternating contractions and re- laxations of the heart, the blood passes through the arteries, not in a steady stream, but, as already described, in a series of welling impulses; and the hemorrhage from a wounded artery is readily distinguished from venous or capillary hemor- rhage by the fact that the blood flows in suc- cessive jets, as well as more rapidly and abund- antly. If a puncture be made in the walls of the ventricle, and a slender canula introduced, the flow of the blood through it is seen to be entirely intermittent. A strong jet takes place at each ventricular contraction, and at each relaxation the flow is completely interrupted. If the puncture be made, however, in any of the large arteries near the heart, the flow of blood through the orifice is no longer intermittent, but is continuous ; only it is very much stronger at the time of ventricular contraction, and diminishes, though it does not entirely cease, at the time of relaxa- tion. This is on account of the elasticity of the arterial coats. For if the blood were driven through a series of perfectly rigid and unyielding tubes, its flow would be everywhere intermittent ; and it would be delivered from an orifice situated at any point, in perfectly Elungation and curva- ture of an Artert in PULSATION. THE ARTERIES AND THE ARTERIAL CIRCULATION. 249 interrupted jets. But the arteries are yielding and elastic. When the contraction of the heart drives the blood into the aorta, a part of its force is expended for the time in distending the walls of the vessel; and this force is returned to the blood, when the heart relaxes, by the elastic reaction of the arterial coats. The interrupted or pulsating character of the arterial current, therefore, which is strongly pronounced in the immediate vicinity of the heart, becomes gradually lost and equalized, during its passage through the vessels, until in the smallest arteries it is nearly imperceptible. The same effect of an elastic medium in equalizing the force of an interrupted current may be shown by fitting to the end of a common syringe a long glass or metallic tube. Whatever be the length of the inelastic tubing, the water which is thrown into one extremity of it by the syringe will be delivered from the other end in distinct jets, corresponding with the strokes of the piston ; but if the metallic tube be replaced by one of India rubber, of sufficient length, the elasticity of this substance merges the force of the sepa- rate impulses into each other, and the water is driven out from the farther extremity in a continuous stream. The elasticity of the arteries, however, never entirely equalizes the force of the separate cardiac pulsations, since a pulsating cha- racter can be seen in the flow of the blood through even the smallest arteries, under the microscope; but this pulsating character dimin- ishes very considerably from the heart outward, and the current becomes much more continuous in the smaller vessels than in the larger. The primary cause, therefore, of the motion of the blood in the arteries is the contraction of the ventricles, which, by driving out the blood in interrupted impulses, distends at every stroke the whole arterial system. But the arterial pulse is not exactly syn- chronous everywhere with the beat of the heart; since a certain amount of time is required to propagate the blood-wave from the centre of the circulation outward. The pulse of the radial artery at the wrist is perceptibly later than that of the heart ; and the pulse of the posterior tibial at the ankle, again, perceptibly later than that at the wrist. The arterial circulation, accordingly, is not an entirely simple phenomenon ; but is made up of the combined effects of two different physical forces. In the first place, there is the elasticity of the entire arterial system, by which the blood is subjected to a constant and uniform pressure, quite independent of the action of the heart. Secondly, there is the alternating contrac- 250 THE CIRCULATION". tion and relaxation of the heart, by which the blood is driven in rapid and successive impulses from the centre of the circulation, to be thence distributed throughout the body. The rapidily with which the blood circulates through the arterial system is very great. Its velocity is greatest in the immediate neighborhood of the heart, and diminishes somewhat as the blood recedes farther and farther from the centre of the circulation. This diminution in the rapidity of the arterial current is due to the suc- cessive division of the aorta and its primary branches into smaller and smaller ramifications, by which the total calibre of the arterial system, as we have already mentioned, is somewhat increased. The blood, therefore, flowing through a larger space as it passes outward, necessarily goes more slowly. At the same time the increased extent of the arterial parietes with which the blood comes in con- tact, as well as the mechanical obstacle arising from the division of the vessels and the separation of the streams, undoubtedly contri- bute more or less to retard the currents. The mechanical obstacle, however, arising from the friction of the blood against the walls of the vessels, which would be very serious in the case of water or any similar fluid flowing through glass or metallic tubes, has compara- tively little effect on the rapidity of the arterial circulation. This can readily be seen by microscopic examination of any transparent and vascular tissue. The internal surface of the arteries is so smooth and yielding, and the consistency of the circulating fluid so accu- rately adapted to that of the vessels which contain it, that the retarding effects of friction are reduced to a minimum, and the blood in flowing through the vessels meets with the least possible resistance. It is owing to this fact that the arterial circulation, though some- what slower toward the periphery than near the heart, yet retains a very remarkable velocity throughout ; and even in arteries of the minutest size it is so rapid that the shape of the blood-globules can- not be distinguished in it on microscopic examination, but only a mingled current shooting forward with increased velocity at every cardiac pulsation. Yolkmann, in Germany, has determined, by a very ingenious contrivance, the velocity of the current of blood in some of the large sized arteries in dogs, horses, and calves. The instrument which he employed (Fig. 94) consisted of a metallic cylinder (a), with a perforation running from end to end, and cor- responding in size with the artery to be examined. The artery was then divided transversely, and its cardiac extremity fastened to the THE ARTERIES AND THE ARTERIAL CIRCULATION. 251 upper end (h) of the instrument, while its peripheral extremity was fastened in the same manner to the lower end (c). The blood accordingly still kept on its usual course; only passing for a short distance through the artificial tube (a), between the divided extremi- Fig. 94. Fig- 95. VoLKMANN's APPARATUS for measuring the rapidity of the arterial circulation. ties of the artery. The instrument, however, was provided, as shown in the accompanying figures, with two transverse cylindrical plugs, also perforated f and arranged in such a manner, that when, at u given signal, the two plugs were suddenly turned in opposite directions, the stream of blood would be turned out of its course (Fig. 95), and made to traverse a long bent tube of glass {d, d, d), before again finding its way back to the lower portion of the artery. In this way the distance passed over by the blood in a given tii:ie could be readily measured upon a scale attached to the side of the glass tube. Volkmann found, as the average result of his obser- vations, that the blood moves in the carotid arteries of warm-blooded quadrupeds with a velocity of 12 inches per second. 252 THE CIRCULATION. VENOUS CIRCULATION. The veins, which collect the blood from the tissues and return it to the heart, are composed, like the arteries, of three coats ; an inner, middle, and exterior. In structure, they differ from the arte- ries in containing a much smaller quantity of muscular and elastic fibres, and a larger proportion of simple condensed areolar tissue. They are consequently more flaccid and compressible than the arteries, and less elastic and contractile. They are furthermore distinguished, throughout the limbs, neck, and external portions of the head and trunk, by being provided with valves, consisting of fibrous sheets arranged in the form of festoons, and so placed in the cavity of the vein as to allow the blood to pass readily from the periphery toward the heart, but to prevent altogether its reflex in an opposite direction. The flow of blood through the veins is less powerful and regular than that through the arteries. It depends on the combined action of three different forces. 1. The force of aspiration of the thorax. — When the chest expands, by the lifting of the ribs and the descent of the diaphragm, it has the effect of drawing into the thoracic cavity all the fluids which can gain access to it. The expanded cavity is principally filled by the air, which passes in through the trachea and fills the bronchial tubes and pulmonary vesicles. But the blood in the large veins is also drawn into the chest at the same time and by the same force. It can readily be seen, when the jugular and sub- clavian veins are exposed in the living animal, that these vessels collapse with every inspiration, and fill out again at the moment of expiration. During inspiration, the blood is drawn forward into that part of the vein which occupies the cavity of the chest ; and during expiration, the flow being momentarily checked by the compression of the thorax, the vein fills up from behind, and again becomes distended. This force does not act efficiently at any great distance from the chest, owing to the flaccidity of the venous parietes, which collapse at a short distance from the entrance to the thoracic cavity, as the vein becomes emptied by inspiration. It is active, however, in the neighborhood of the chest, and the respiratory movements exert, therefore, a certain degree of influence on the venous circulation. VENOUS CIECULATION 253 2. Tlie contraction of the voluntary muscles. — The veins which convey the blood through the limbs, and the parietes of the head and trunk, lie among voluntary muscles, which are more or less constantly in a state of alternate contraction and relaxation. At every contraction these muscles become swollen laterally, and, of course, compress the veins which are situated between them. The blood, driven out from the vein by this pressure, cannot regurgitate toward the capillaries, owing to the valves, already described, which shut back and prevent its reflux. It is accordingly forced onward toward the heart; and when the muscle relaxes and the vein is liberated from pressure, it again fills up from behind, and the cir- culation goes on as before. This force is a very efficient one in producing the venous circulation ; since the voluntary muscles are more or less active in every position of the body, and the veins constantly liable to be compressed by them. It is on this account that the veins, in the external parts of the body, communicate so freely with each other by transverse branches ; in order that the current of blood, which is momentarily excluded from one vein by the pressure of the muscles, may readily find a passage through others, which communicate by cross branches with the first. (Figs. 96 and 97.) Fig. 96. Fig. 97. Vein with valves open. Vein with valves closed; stream of blood passing off by a lateral channel. 3. The force of the capillary circulation. — This last cause of the motion of the blood through the veins is the most important of all, as it is the only one which is constantly and universally active. In 254 THE CIRCULATION. fish, for example, respiration is performed altogether by gills; and in reptiles the air is forced down into the lungs by a kind of deglu- tition, instead of being drawn in by the expansion of the chest. In neither of these classes, therefore, can the movements of respiration assist mechanically in the circulation of the blood. In the splanch- nic cavities, again, of all the vertebrate animals, the veins coming from the internal organs, as, for example, the cerebral, pulmonary, portal, hepatic, and renal veins, are unprovided with valves ; and the passage of the blood through them cannot therefore be effected by any lateral pressure. The circulation, however, constantly going on in the capillaries, everywhere tends to crowd the radicles of the veins with blood ; and this vis a tergo, or pressure from behind, fills the whole venous system by a constant and steady accumulation. So long, therefore, as the veins are relieved of blood at their cardiac extremity by the regular pulsations of the heart, there is no back- ward pressure to oppose the impulse derived from the capillary cir- culation ; and the movement of the blood through the veins continues in a steady and uniform course. With regard to the rapidity of the venous circulation, no direct results have been obtained by experiment. Owing to the flaccidity of the venous parietes, and the readiness with which the flow of blood through them is disturbed, it is not possible to determine this point for the veins, in the same manner as it has been determined for the arteries. The only calculation which has been made in this respect is based upon a comparison of the total capacity of the arterial and venous systems. As the same blood which passes out- ward through the arteries, passes inward again through the veins, the rapidity of its flow in each must be in inverse proportion to the capacity of the two sets of vessels. That is to say, a quantity of blood which would pass in a given time, with a velocity of x, through an opening equal to one square inch, would pass during the same time through an opening equal to two square inches, with a velocity of J; and would require, on the other hand, a velocity of 2 «, to pass in the same time through an opening equal to one- half a square inch. Now the capacity of the entire venous system, when distended by injection, is about twice as great as that of the entire arterial system. During life, however, the venous system is at no time so completely filled with blood as is the case with the arteries ; and making allowance for this difference, we find that the entire quantity of venous blood is to the entire quantity of arte- THE CAPILLARY CIKCULATION. 255 rial blood nearly as three to two. The velocity of the venous blood, as compared with the arterial, is therefore as two to three ; or about 8 inches per second. It will be understood, however, that this calculation is altogether approximative, and not exact ; since the venous current varies, according to many different circumstances, in different parts of the body ; being slower near the capillaries, and more rapid near the heart. It expresses, however, with sufl&- cient accuracy, the relative velocity of the arterial and venous cur- rents, at corresponding parts of their course. THE CAPILLARY CIRCULATION. The capillary bloodvessels are minute inosculating tubes, which permeate the vascular organs in every direction, and bring the blood into intimate contact with the substance of the tissues. They are continuous with the terminal ramifications of the arteries on the one hand, and with the com- mencing rootlets of the veins on Fig- 98. the other. They vary somewhat in size in different organs, and in different species of animals ; their average diameter in the human subject being a little over 3 (jVw of an inch. They are composed of a single, transparent, homogene- ous, somewhat elastic, tubular membrane, which is provided at various intervals with flattened, oval nuclei. As the smaller arte- ries approach the capillaries, they diminish constantly in size by successive subdivision, and lose first their external or fibrous tunic. They are then composed only of the internal or homogeneous coat, and the midde or muscu- lar. (Fig. 98, a.) The middle coat then diminishes in thickness, until it is reduced to a single layer of circular, fusiform, unstriped, muscular fibres, which in their turn disappear altogether as the artery merges at last in the capillaries ; leaving only, as we have SiMALL AuTERT, wUh its muscular tiiaic (a), breaking up into capillaries. From the 2Jia ■mater. 256 THE CIECULATIOX. already mentioned, a simple, homogeneous, nucleated, tubular mem- brane, which is continuous with the internal arterial tunic. The capillaries are further distinguished from both arteries and veins by their frequent inosculation. The arteries constantly divide and subdivide, as they pass from within outward; while the veins as constantly unite with each other to form larger and less numerous branches and trunks, as they pass from the circum- ference toward the centre. But the capillaries simply inosculate with each other in every direction, in such a manner as to form an interlacing network or plexus, the capillary plexus (Fig. 99), which is exceedingly rich and abundant in some organs, less so in others. The spaces included between the meshes of the capillary network vary also, in shape as well as in size, in different parts of the body. In the muscular tissue they '^' form long parallelograms ; in the areolar tissue, irregular shapeless figures, correspond- ing with the direction of the fibrous bundles of which the tissue is composed. In the mucous membrane of the large intestine, the capillaries include hexagonal or nearly circular spaces, inclosing the orifices of the follicles. In the papillae of the tongue and of the skin, and in the tufts of the placenta, they are arranged in long spiral loops, and in the adipose tissue in wide meshes, among which the fat vesicles are entangled. The motion of the hlood in the cajnllaries may be studied by examining under the microscope any transparent tissue, of a sufficient degree of vascularity. One of the most convenient parts for this purpose is the web of the frog's foot. "When properly prepared and kept moistened by the occasional addition of water to the integument, the circulation will go on in its vessels for an indefinite length of time. The blood can be seen entering the field by the smaller arteries, shooting along through them with great rapidity and in successive impulses^ and flowing off again by the veins at a somewhat slower rate. In the capillaries themselves Capillary Network from tveb of frog's foot. THE CAPILLARY CIRCULATION. 257 Capillary Circulation in web of frog's foot. the circulation is considerably less rapid than in either the arteries or the veins. It is also perfectly steady and uninterrupted in its flow. The blood passes along in a uniform and continuous current, without any apparent contraction or dilatation of the vessels, very much as if it were flowing through glass tubes. An- ^^s- ^^^• other very remarkable pe- culiarity of the capillary circulation is that it has no definite direction. The nu- merous streams of which it is composed (Fig. 100) do not tend to the right or to the left, nor toward any one particular point. On the contrary, they pass above and below each other, at right angles to each other's course, or even in opposite directions; so that the blood, while in the capillaries, merely circulates promiscuously among the tissues, in such a manner as to come intimately in contact with every part of their substance. The motion of the white and red globules in the circulating blood is also peculiar, and shows very distinctly the difference in their consistency and other physical properties. In the larger vessels the red globules are carried along in a dense column, in the central part of the stream ; while near the edges of the vessel there is a transparent space occupied only by the clear plasma of the blood, in which no red globules are to be seen. In the smaller vessels, the globules pass along in a narrower column, two by two, or following each other in single file. The flexibility and semi-fluid consistency of these globules are here very apparent, from the readiness with which they become folded up, bent or twisted in turning corners, and the ease with which they glide through minute branches of communication, smaller in diameter than themselves. The white globules, on the other hand, flow more slowly and with greater difficulty through the vessels. They drag along the exter- nal portions of the current, and are sometimes momentarily arrested; apparently adhering for a few seconds to the internal surface of the vessel. Whenever the current is obstructed or retarded in any 17 258 THE CIRCULATION. manner, the white globules accumulate in the affected portion, and become more numerous there in proportion to the red. It is during the capillary circulation that the blood serves for the nutrition of the vascular organs. Its fluid portions slowly transude through the walls of the vessels, and are absorbed by the tissues in such proportion as is requisite for their nourishment. The saline substances enter at once into the composition of the surrounding parts, generally without undergoing any change. The phosphate of lime, for example, is taken up in large quantity by the bones and cartilages, and in smaller quantity by the softer parts; while the chlorides of sodium and potassium, the carbonates, sul- phates, &c., are appropriated in special proportions by the different tissues, according to the quantity necessary for their organization. The albuminous ingredients of the blood, on the other hand, are not only absorbed in a similar manner by the animal tissues, but at the same time are transformed by catalysis, and converted into new materials, characteristic of tbe different tissues. In this way are produced the musculine of the muscles, the osteine of the bones, the cartilagine of the cartilages, &c. &c. It is probable that this trans- formation does not take place in the interior of the vessels them- selves ; but that the organic ingredients of the blood are absorbed by the tissues, and at the same moment converted into new mate- rials, by contact with their substance. The blood in this way fur- nishes, directly or indirectly, all the materials necessary for the nutrition of the body. The physical forces which produce the movement of the blood through the capillary vessels are different from those which operate in producing the venous and arterial circulations. The force of the heart's action, whicb drives the blood througb the arteries, merely secures a constant supply of blood to the commencement of the capillaries, but is not of itself the cause of the continued current through the latter vessels. This is shown by the fact that a capil- lary circulation exists in vegetables, where there is no central con- tractile organ, and where the circulation of the sap must necessarily depend on other forces. In fish, again, the heart is separated from the general circulation by the capillary system of the gills, which intervenes between them, and by which the impulsive force of the cardiac contractions must be, to say the least, very much diminished. In the quadrupeds, also, and in the human subject, the bepatic capillary circulation goes on with the same regularity as elsewhere, notwithstanding that it is supplied by the portal vein with blood THE CAPILLARY CIRCULATION. 259 which has already passed through the capillaries of the digestive apparatus before reaching those of the liver. Furthermore, it is evi- dent, from the commonest observation, that the capillary circulation, in various parts of the body, is liable to be increased or diminished in activity from causes of a purely local character, and entirely inde- pendent of the action of the heart. The countenance may become flushed or pale, and temporary congestions may take place in any of the internal organs, as, for example, in the stomach and intestine during the digestive process ; so that a much larger quantity of blood may circulate through them in a given time, though the force of the heart's action remains constant. The study of the microscopic appearances of inflammation in transparent tissues, so well observed by Lebert^ and others, leads to a similar result. At the time when the inflammatory process begins to be established, the circulation is seen to be retarded. The blood moves more slowly through the capillaries, and its current becomes more and more sluggish as the morbid action proceeds. The blood- globules become impacted in the vessels, and finally, at those points where the inflammation is most active, the circulation stops alto- gether, and a local stasis of the blood is produced. This, however, is not owing to any simple mechanical obstacle to its passage, since the vessels, during the entire period of the inflammatory congestion, are actually dilated by the blood which accumulates in them ; nor, on the other hand, is it owing to a diminution in the force of the heart's action, since the cardiac pulsations are still as powerful as ever, and in the healthy tissues, in the immediate neighborhood of the inflamed parts, the circulation may be seen going on at the same time in its ordinary manner. All these facts show beyond a doubt that the force which regu- lates and controls the capillary circulation is a local one, and resident evidently in the capillaries and tissues of the part itself. This force not only carries the blood through the capillaries, but delivers it also into the veins ; and by supplying the requisite vis a iergo^ con- stitutes, as we have already mentioned, one of the most eflBeient causes of the venous circulation. It is not only independent to a certain extent, during life, of the heart's action, but will even con- tinue for a time after the heart has stopped; so that at the moment of death, when the cardiac pulsations cease, the capillaries empty themselves of blood, which accumulates in the large trunks and ' Pliysiologie Pathol ogique, Paris, 1845, voL i. p. 2. 260 THE CIRCULATION. branches of the veins. Precisely what is the nature of this force, thus active in producing the capillary circulation, it is not easy to determine. In all probability it results, in great measure, from the actions of endosmosis and exosmosis, which are constantly going on between the blood contained in the capillaries, and the tissues situated outside of them ; and which, varying in intensity in different organs, and even in the same organ at different times, produce those local variations in the activity of the capillary circulation which con- stantly present themselves to the observation of the physiologist. The rapidity of the circulation in the capillary vessels is much less than in the arteries or the veins. It may be measured, with a tolerable approach to accuracy, during the microscopic examina- tion of transparent and vascular tissues, as, for example, the web of the frog's foot, or the mesentery of the rat. The results obtained in this way by different observers (Valentine, Weber, Volkmann, &c.) show that the rate of movement of the blood through the capillaries is rather less than one-thirtieth of an inch per second ; or not quite two inches per minute. Since the rapidity of the current, as we have mentioned above, must be in inverse ratio to the entire calibre of the vessels through which it moves, it follows that the united calibre of all the capillaries of the body must be from 350 to 400 times greater than that of the arteries. It must not be supposed from this, however, that the whole quantity of blood contained in the capillaries at any one time is so much greater than that in the arteries ; since, although the united calibre of the capil- laries is very large, their length is very small. The effect of the anatomical structure of the capillary system is, therefore, merely to disseminate a comparatively small quantity of blood over a very large space, so that the chemico-physiological reactions, necessary to nutrition, may take place with promptitude and energy. For the same reason, although the rate of movement of the blood in these vessels is very slow, yet as the distance to be passed over between the arteries and veins is very small, the blood really re- quires but a short time to traverse the capillary system, and to commence its returning passage by the veins. GENERAL CONSIDERATIONS. The rapidity with which the blood passes through the entire round of the circulation is a point of great interest, and one which has KAPIDITY OF THE CIRCULATION. 261 received a considerable share of attention. The results of such experiments, as have been tried, show that this rapidity is much greater than would have been anticipated. Hering, Poisseuille, and Matteucci,* have all experimented on this subject in the following manner. A solution of ferrocyanide of potassium was injected into the right jugular vein of a horse, at the same time that a liga- ture was placed upon the corresponding vein on the left side, and an opening made in it above the ligature. The blood flowing from the left jugular vein was then received in separate vessels, which were changed every five seconds, and the contents afterward exa- mined. It was thus found that the blood drawn from the first to the twentieth second contained no traces of the ferrocyanide ; but that which escaped from the vein at the end of from twenty to twenty-five seconds, showed unmistakable evidence of the presence of the foreign salt. The ferrocyanide of potassium must therefore, during this time, have passed from the point of injection to the right side of the heart, thence to the lungs and through the pulmo- nary circulation, returned to the heart, passed out again through the arteries to the capillary system of the head and neck, and thence have commenced its returning passage to the right side of the heart, through the jugular vein. If this experiment were altogether decisive, it would demonstrate that the blood performs the entire round of the circulation in from 20 to 25 seconds. But it is not so conclusive in this respect as might at first be supposed. In reality, it only shows that the solution of the ferrocyanide passes round to the opposite vein during this period, but it does not necessarily follow that the entire blood moves with the same rapidity ; since the injected saline solution may diffuse itself through the blood, so as to travel faster than the blood itself. Sub- sequent experiments of Poisseuille showed, in fact, that other sub- stances injected at the same time may either accelerate or retard the movement of the ferrocyanide. If a little nitrate of potass, for example, were injected together with the ferrocyanide, the latter salt appeared in the blood flowing from the opposite jugular at the end of twenty seconds. A solution of acetate of ammonia, again, shortened the period to eighteen seconds. On the other hand, a little alcohol, injected at the same time, retarded its motion to such a degree, that the ferrocyanide could not be detected till the end of ' Physical Phenomena of Living Beings, Pereira's translation, Philada. ed , 1848, p. 317. 262 THE CIECULATION. forty to forty-five seconds. These facts show conclusively that the time required for a solution of ferrocyanide of potassium to appear in the opposite jugular vein does not depend altogether on the rate of movement of the blood itself, but is influenced very considerably by the chemical constitution and physical properties of the injected fluid, and its physical relations to the blood and to the walls of the bloodvessels. Furthermore, we have already seen that the different ingredients of the blood do not all circulate with the same rapidity. In a microscopic examination, for example, it is evident that the white globules of the blood move much more slowly than the red; and it is very possible that the red globules themselves pass less rapidly from one point to another than those portions of the blood which are entirely fluid. The truth is, however, that we cannot fix upon any uniform rate which shall express exactly the time required by the entire blood to pass the round of the whole vascular system, and return to a given point. The circulation of the blood, far from being a simple phenomenon, like a current of water through a circular tube, is, on the contrary, extremely complicated in all its anatomical and physiological conditions; and it differs in rapidity, as well as in its physical and chemical phenomena, in different parts of the circu- latory apparatus. We have already seen how much the form of the capillary plexus varies in different organs. In some the vascu- lar network is close, in others comparatively open. In some its meshes are circular in shape, in others polygonal, in others rectan- gular. In some the vessels are arranged in twisted loops, in others they communicate by irregular but abundant inosculations. The mere distance at which an organ is situated from the heart must modify to some extent the time required for its blood to return again to the centre of the circulation. The blood which passes through the coronary arteries, for example, and the capillaries of the heart itself, must be returned to the right auricle in a compara- tively short time ; while that which is carried by the carotids into the capillary system of the head and neck, to return by the jugulars, will require a longer interval. That, again, which descends by the abdominal aorta and its divisions, to the lower extremities, and which, after circulating through the tissues of the leg and foot, mounts upward through the whole course of the saphena, femoral, iliac and abdominal veins, must be still, longer on its way ; while that which circulates through the abdominal digestive organs and is then collected by the portal system, to be again dispersed through LOCAL VARIATIONS. 263 the glandular tissue of the liver, requires undoubtedly a longer period still to perform its double capillary circulation. The blood, therefore, arrives at the right side of the heart, from different parts of the body, at successive intervals; and may pass several times through one organ while performing a single circulation through another. Furthermore, the chemical phenomena taking place in the blood and the tissues vary to a similar extent in different organs. The actions of transformation and decomposition, of nutrition and secre- tion, of endosmosis and exosmosis, which go on simultaneously throughout the body, are yet extremely varied in their character, and produced a similar variation in the phenomena of the circula- tion. In one organ the blood loses more fluid than it absorbs ; in another it absorbs more than it loses. The venous blood, conse- quently, has a different composition as it returns from different organs. In the brain and spinal cord it gives up those ingredients necessary for the nutrition of the nervous matter, and absorbs cho- lesterine and other materials resulting from its waste; in the muscles it loses those substances necessary for the supply of the muscular tissue, and in the bones those which are requisite for the osseous system. In the parotid gland it yields the ingredients of the saliva; in the kidneys, those of the urine. In the intestine it absorbs in large quantity the nutritious elements of the digested food ; and in the liver, gives up substances destined finally to produce the bile, at the same time that it absorbs sugar, which has been produced in the hepatic tissue. In the lungs, again, it is the elimination of carbonic acid and the absorption of oxygen that constitute its prin- cipal changes. It has been already remarked that the temperature of the blood varies in different veins, according to the peculiar chemical and nutritive changes going on in the organs from which they originate. Its color, even, which is also dependent on the chemical and nutritive actions taking place in the capillaries, varies in a similar manner. In the lungs, it changes from blue to red ; in the capillaries of the general system, from red to blue. But its tinge also varies very considerably in different parts of the general circulation. The blood of the hepatic veins is darker than that of the femoral or brachial vein. In the renal veins it is very much brighter than in the vena cava; and when the circulation through the kidneys is free, the blood returning from them is nearly as red as arterial blood. We must regard the circulation of the blood, therefore, not as a 264 THE CIRCULATION. simple process, but as made up of many different circulations, going on simultaneously in different organs. It has been customary to illustrate it, in diagram, by a double circle, or figure of 8, of which the upper arc is used to represent the pulmonary, the lower the general cir- culation. This, however, gives but a very imperfect idea of the entire cir- culation, as it really takes place. It would be much more accurately re- presented by such a diagram as that given in Fig. 101, in which its varia- tions in different parts of the body are indicated in such a manner as to show, in some degree, the complicated cha- racter of its phenomena. The circula- tion is modified in these different parts, not only in its mechanism, but also in its rapidity and quantity, and in the nutritive functions performed by the blood. In one part, it stimulates the nervous centres and the organs of special sense; in others it supplies the fluid secretions, or the ingredients of the solid tissues. One portion, in passing through the digestive appara- tus, absorbs the materials requisite for the nourishment of the body ; an- other, in circulating through the lungs, exhales the carbonic acid which it has accumulated elsewhere, and absorbs the oxygen which is afterward trans- ported to distant tissues by the current of arterial blood. The phenomena of the circulation are even liable, as we have already seen, to periodical va- riations in the same organ; increas- ing or diminishing in intensity with the condition of rest or activity of the whole body, or of the particular organ which is the subject of observation. Diagram of the Circulation. — 1. Heart. 2. Lungs. 3. Head and upper extremities. 4. Spleen. 5. Intestine. 6. Kidney. 7. Lower extremities. 8 Liver. SECRETION. 265 CHAPTER XV. SECRETION. We have already seen, in the last chapter, how the elements of the blood are absorbed by the tissues during the capillary circula- tion, and assimilated by them or converted into their own substance. In this process, the inorganic or saline matters are mostly taken up unchanged, and are merely appropriated by the surrounding parts in particular quantities ; while the organic substances are transformed into new compounds, characteristic of the dififerent tissues by which they are assimilated. In this way the various tissues of the body, though they have a diiferent chemical composition from the blood, are nevertheless supplied by it with appropriate ingredients, and their nutrition constantly maintained. Beside this process, which is known by the name of " assimila- tion," there is another somewhat similar to it, which takes place in the different glandular organs, known as the process of secretion. It is the object of this function to supply certain fluids, differing in chemical constitution from the blood, which are required to assist in various physical and chemical actions going on in the body. These secreted fluids, or "secretions," as they are called, vary in consistency, density, color, quantity, and reaction. Some of them are thin and watery, like the tears and the perspiration ; others are viscid and glutinous, like mucus and the pancreatic fluid. They are alkaline like the saliva, acid like the gastric juice, or neutral like the bile. Each secretion contains water and the inorganic salts of the blood, in varying proportions; and is furthermore distin- guished by the presence of some peculiar animal substance which does not exist in the blood, but which is produced by the secreting action of the glandular organ. As the blood circulates through the capillaries of the gland, its watery and saline constituents transude in certain quantities, and are discharged into the excretory duct. At the same time, the glandular cells, which have themselves been nourished by the blood, produce a new substance by the catalytic 266 SECRETION. transformation of their organic constituents; and this new substance is discharged also into the excretory duct and mingled with the other ingredients of the secreted fluid. A true secretion, therefore, is produced only in its own particular gland, and cannot be formed elsewhere, since the glandular cells of that organ are the only ones capable of producing its most characteristic ingredient. Thus pepsine is formed only in the tubules of the gastric mucous mem- brane, pancreatine only in the pancreas, tauro-cholate of soda only in the liver. One secreting gland, consequently, can never perform vicariously the office of another. Those instances which have been from time to time reported of such an unnatural action are not, properly speaking, instances of "vicarious secretion;" but only cases in which certain substances, already existing in the blood, have made their appearance in secretions to which they do not naturally belong. Thus cholesterine, which is produced in the brain and is taken up from it by the blood, usually passes out with the bile ; but it may also appear in the fluid of hydrocele, or in inflammatory exuda- tions. The sugar, again, which is produced in the liver and taken up by the blood, when it accumulates in large quantity in the cir- culating fluid, may pass out with the urine. The coloring matter of the bile, in cases of biliary obstruction, may be reabsorbed, and so make its appearance in the serous fluids, or even in the perspira- tion. In these instances, however, the unnatural ingredient is not actually produced by the kidneys, or the perspiratory glands, but is merely supplied to them, already formed, by the blood. Cases of *' vicarious menstruation" are simply capillary hemorrhages which take place from various mucous membranes, owing to the general disturbance of the circulation in amenorrhcea. A true secretion, however, is always confined to the gland in which it naturally originates. The force by which the different secreted fluids are. prepared in the glandular organs, and discharged into their ducts, is a peculiar one, and resident only in the glands themselves. It is not simply a process of filtration, in which the ingredients of the secretion exude from the bloodvessels by exosmosis under the influence of pressure ; since the most characteristic of these ingredients, as we have already mentioned, do not pre-exist in the blood, but are formed in the substance of the gland itself. Substances, even, which already exist in the blood in a soluble form, may not have the power of passing out through the glandular tissue. Bernard SECRETION. 267 has found' that ferrocjanide of potassium, when injected into the jugular vein, though it appears with great facility in the urine, does not pass out with the saliva; and even that a solution of the same salt, injected into the duct of the parotid gland, is ab- sorbed, taken up by the blood, and discharged with the urine ; but does not appear in the saliva, even of the gland into which it has been injected. The force with which the secreted fluids accumulate in the salivary ducts has also been shown by Ludwig's experi- ments'* to be sometimes greater than the pressure in the bloodves- sels. This author found, by applying mercurial gauges at the same time to the duct of Steno and to the artery of the parotid gland, that the pressure in the duct from the secreted saliva was considerably greater than that in the artery from the circulating blood ; so that the passage of the secreted fluids had really taken place in a direc- tion contrary to that which would have been caused by the simple influence of pressure. The process of secretion, therefore, is one which depends upon the peculiar anatomical and chemical constitution of the glandular tissue and its secreting cells. These cells have the property of absorbing and transmitting from the blood certain inorganic and saline substances, and of producing, by chemical metamorphosis, certain peculiar animal matters from their own tissue. These sub- stances are then mingled together, dissolved in the watery fluids of the secretion, arud discharged simultaneously by the excretory duct. All the secreting organs vary in activity at different periods. Sometimes they are nearly at rest ; while at certain periods they become excited, under the influence of an occasional or periodical stimulus, and then pour out their secretion with great rapidity and in large quantity. The perspiration, for example, is usually so slowly secreted that it evaporates as rapidly as it is poured out, and the surface of the, skin remains dry; but under the influence of unusual bodily exercise or mental excitement it is secreted much faster than it can evaporate, and the whole integument becomes covered with moisture. The gastric juice, again, in the intervals of digestion is either not secreted at all, or is produced in a nearly inappreciable quantity ; but on the introduction of food into the stomach, it is immediately poured out in such abundance, that between two and three ounces may be collected in a quarter of an hour. ' Lecjons de Physiologie ExiJerimentale. Paris, 1856, tome ii. p. 96, et seq. 2 Ibid , p. 106. 268 SECRETION. « The principal secretions met with in the animal body are as follows : — 1. Mucus. 6. Saliva. 2. Sebaceous matter. 7. Gastric juice. 3. Perspiration. 8. Pancreatic juice. 4. The tears. 9. Intestinal juice. 5. The milk. 10. Bile. The last five of these fluids have already been described in the preceding chapters. We shall therefore only require to examine at present the five following, viz., mucus, sebaceous matter, per- spiration, the tears, and the milk, together with some peculiarities in the secretion of the bile. 1. Mucus. — Nearly all the mucous membranes are provided with follicles or glandulas, in which the mucus is prepared. These folli- cles are most abundant in the lining membrane of the mouth, nares, pharynx, oesophagus, trachea and bronchi, vagina, and male urethra. They are generally of a compound form, consisting of a number of secreting sacs or cavities, terminating at one end in a blind ex- tremity, and opening by the other into a common diict by which the secreted fluid is discharged. Each ultimate secreting sac or follicle is lined with glandular epithelium (Fig. 102), and surround- ed on its external surface by a network of capillary bloodvessels. These vessels, penetrating deeply into the interstices between the follicles, bring the blood nearly into contact with the epithelial cells lining its cavity. It is these cells which prepare the secretion, and discharge it afterward into the commencement of the „ excretory duct. Follicles of a Com- J POUND Mucous Glandule. The mucus, produccd in the manner From the human subject. (After ., -i ., ■, . , ,, n • ^ K;3iiiker.)_a. Membrane of the . ^bovc dcscribcd, IS a clcar, colorlcss fluid, follicle, h, c. Epithelium of the which is pourcd out in larger or smaller quantity on the surface of the mucous membranes. It is distinguished from other secretions by its vis- cidity, which is its most marked physical property, and which depends on the presence of a peculiar animal matter, known under the name of mucosine. When unmixed with other animal fluids, this viscidity is so great that the mucus has nearly a semi-solid or gelatinous consistency. Thus, the mucus of the mouth, when ob- tained unmixed with the secretions of the salivary glands, is so SEBACEOUS MATTER. 269 tougli and adhesive that the vessel containing it may be turned upside down without its running out. The mucus of the cervix uteri has a similar firm consistency, so as to' block up the cavity of this part of the organ with a semi-solid gelatinous mass. Mucus is at the same time exceedingly smooth and slippery to the touch, so that it lubricates readily the surfaces upon which it is exuded, and facilitates the passage of foreign substances, while it defends the mucous membrane itself from injury. The composition of mucus, according to the analyses of Nasse,^ is as follows : — Composition of Pulmonary Mucus. Water 955.52 Animal matter .......... 33.57 Fat 2.89 Chloride of sodium ......... 5.83 Phosphates of soda and potass ...... 1.05 Sulphates " " 0.65 Carbonates " " 0.49 1000.00 The animal matter of mucus is insoluble in water ; and conse- quently mucus, when dropped into water, does not mix with it, but is merely broken up by agitation into gelatinous threads and flakes, which subside after a time to the bottom. It is miscible, however, to some extent, with other animal fluids, and may be incorporated with them, so as to become thinner and more dilute. It readily takes on putrefactive changes, and communicates them to other organic substances with which it may be in contact. The varieties of mucus found in different parts of the body are probably not identical, but differ a little in the character of their principal organic ingredient, as well as in the proportions of their saline constituents. The function of mucus is for the most part a physical one, viz., to lubricate the mucous surfaces, to defend them from injury, and to facilitate the passage of foreign substances through their cavities. 2. Sebaceous Matter, — The sebaceous matter is distinguished by containing a very large proportion of fatty or oily ingredients. There are three varieties of this secretion met with in the body, viz., one produced by the sebaceous glands of the skin, another by the ceruminous glands of the external auditory meatus, and a third by the Meibomian glands of the eyelid. The sebaceous ' Simon's Chemistry of Man, Philada., 1846, p. 352. 270 SECRETION. glands of the skin are found most abundantly in those parts which are thickly covered with hairs, as well as on the face, the labia minora of the female generative organs, the glans penis, and the prepuce. They consist sometimes of a simple follicle, or flask- shaped cavity, opening by a single orifice ; but more frequently of a number of such follicles grouped round a common excretory duct. The duct nearly always opens just at the root of one of the hairs, which is smeared more or less abundantly Fig- 103. with its secretion. Each follicle, as in the case of the mucous glandules, is lined with epithelium, and its cavity is filled yi^^ .i'"'^^*f^^^B ^^^'^ ^^ secreted sebaceous matter. "?^'^'»"^V- .^^H In the Meibomian glands of the eye- pf '-'>- '^^*"'^^B lid (Fig. 103), the follicles are ranged ''.. ,- :' " ^^H along the sides of an excretory duet, y, ,. '-'"i ' . ^^^H situated just beneath the conjunctiva, on |'£? ;' .1 '^'^'''^^B th® posterior surface of the tarsus, and >' "^ '"^ "* i^^l opening upon its free edge, a little be- l^iiRJI^ffli^^H ^i^d *^® roots of the eyelashes. The ceruminous glands of the external audi- tory meatus, again, have .the form of long tubes, which terminate, at the lower part Meibomian Glands, after o ^ • t • i Liidovic. 01 the integument Iming the meatus, in a globular coil, or convolution, covered externally by a network of capillary bloodvessels. The sebaceous matter of the skin has the following composition, according to Esenbeck.' Composition of Sebaceous Mattek. Animal substances ....... 358 Fatty matters ........ 868 Phosphate of lime ........ 200 Carbonate of lime ........ 21 Carbonate of magnesia ....... 16 Chloride of sodium ^ I . ■ 37 Acetate of soda, &c. j • • • • • • 1000 Owing to the large proportion of stearine in the fatty ingredients of the sebaceous matters, they have a considerable degree of con- sistency. Their office is to lubricate the integument and the hairs, to keep them soft and pliable, and to prevent their drying up by ' Simon's Chemistry of Man, p. 379. PEESPIRATION. 271 too rapid evaporation. Wlien the sebaceous glands of the scalp are inactive or atrophied, the hairs become dry and brittle, are easily split or broken off, and finally cease growing altogether. The ceruminous matter of the ear is intended without doubt partly to obstruct the cavity of the meatus, and by its glutinous consist- ency and strong odor to prevent small insects from accidentally introducing themselves into the meatus. The secretion of the Meibomian glands, by being smeared on the edges of the eyelids, prevents the tears from running over upon the cheeks, and confines them within the cavity of the lachrymal canals. Fiff. 104. 3. Perspiration. — The perspiratory glands of the skin are scattered everywhere throughout the integument, being most abundant on the anterior portions of the body. They consist each of a slender tube, about -^l^ of an inch in diameter, lined with glandular epithelium, which penetrates nearly through the entire thickness of the skin, and terminates below in a globular coil, very similar in appearance to that of the ceruminous glands of the ear. (Fig. 104.) A network of capil- lary vessels envelops the tubular coil and supplies the gland with the materials necessary to its se- cretion. These glands are very abundant in some parts. On the posterior portion of the trunk, the cheeks, and the skin of the thigh and leg there are, according to Krause,* about 500 to the square inch ; on the anterior part of the trunk, the forehead, the neck, the forearm, and the back of the hand and foot 1000 to the square inch ; and on the sole of the foot and the palm of the hand about 2700 in the same space. According to the same observer, the whole number of per- spiratory glands is not less than 2,300,000, and the length of each tubular coil, when unravelled, about y'g of an inch. The entire length of the glandular tubing must therefore be not less than 153,000 inches, or about two miles and a half. A Perspiratory Gland, -with its ves- sels ; magnified 35 times. (After Todd and Bow- man.) — a. Glandular coil. 6. Plexus of vessels, c. Excretory duct. Kcilliker, Handbucli der Gewebelelire, Leipzig, 1852, p. 147. 272 SECEETioisr. It is easy to understand, therefore, that a very large quantity of fluid may be supplied from so extensive a glandular apparatus. It results from the researches of Lavoisier and Seguin^ that the ave- rage quantity of fluid lost by cutaneous perspiration during 24 hours is 18,770 grains, or nearly two pounds avoirdupois. A still larger quantity than this may be discharged during a shorter time, when the external temperature is high and the circulation active. Dr. Southwood Smith^ found that the laborers employed in gas works lost sometimes as much as 3 J pounds' weight, by both cata- neous and pulmonary exhalation, in less than an hour. In these cases, as Seguin has shown, the amount of cutaneous transpiration is about twice as great as that which takes place through the lungs. The perspiration is a colorless watery fluid, generally with a distinctly acid reaction, and > having a peculiar odor, which varies somewhat according to the part of the body from which the speci- men is obtained. Its chemical constitution, according to Anselmino.^ is as follows : — C'JIPOSITION OF THE PeKSPIBATION. Water 995.00 Animal matters, with lime ........ .10 Sulphates, and substances soluble in water .... 1.05 Chlorides of sodium and potassium, and spirit-extract . . 2.40 Acetic acid, acetates, lactates, and alcohol-extract . . . 1.45 1000.00 The office of the cutaneous perspiration is principally to regulate the temperature of the body. We have already seen, in a preced- ing chapter, that the living body will maintain the temperature of 100° F., though subjected to a much lower temperature by the surrounding atmosphere, in consequence of the continued genera- tion of heat which takes place in its interior ; and that if, by long continued or severe exposure, the blood become cooled down much below its natural standard, death inevitably results. But the body has also the power of resisting an unnaturally high temperature, as well as an unnaturally low one. If exposed to the influence of an atmosphere warmer than 100° F., the body does not become heated up to the temperature of the air, but remains at its natural standard. This is provided for by the action of the cutaneous glands, which are excited to unusual activity, and pour out a large quantity of watery fluid upon the skin. This fluid immediately ' Robin and Verdeil. Op. cit., vol. ii. p. 145. ^ Philosophy of Health, London, 1838, chap. xiii. ^ Simon. Op. cit., p. 374. THE TEARS. 273 evaporates, and in assuming the gaseous form causes so much heat to become latent that the cutaneous surfaces are cooled down to their natural temperature. . So long as the air is dry, so that evaporation from the surface can go on rapidly, a very elevated temperature can be borne with impunity. The workmen of the sculptor Chantrey were in the habit, according to Dr. Carpenter, of entering a furnace in which the air was heated up to 350°; and other instances have been known in which a temperature of 400° to 600° has been borne for a time without much inconvenience. But if the air be saturated with moisture, and evaporation from the skin in this way retarded, the body soon becomes unnaturally warm ; and if the exposure be long continued, death is the result. It is easily seen that horses, when fast driven, suffer much more from a warm and moist atmosphere than from a warm and dry one. The experiments of Magendie and others have shown' that quadrupeds confined in a dry atmosphere suffer at first but little inQonvenience, even when the temperature is much above that of their own bodies ; but as soon as the atmo- sphere is loaded with moisture, or the supply of perspiration is ex- hausted, the blood becomes heated, and the animal dies. Death follows in these cases as soon as the blood has become heated up to 8° or 9° F., above its natural standard. The temperature of 110°, therefore, which is the natural temperature of birds, is fatal to quad- rupeds ; and we have found that frogs, whose natural temperature is 50° or 60°, die very soon if they are kept in water at 100° F. The amount of perspiration is liable to variation, as we have already intimated, from the variations in temperature of the sur- rounding atmosphere. It is excited also by unusual muscular exertion, and increased or diminished by various nervous condi- tions, such as anxiety, irritation, lassitude, or excitement. 4. The Tears. — The tears are produced by lobulated glands situated at the upper and outer part of the orbit of the eye, and opening, by from six to twelve ducts, upon the surface of the con- junctiva, in the fold between the eyeball and the outer portion of the upper lid. The secretion is extremely watery in its composition, and contains only about one part per thousand of solid matters, consisting mostly of chloride of sodium and animal extractive matter. The office of the lachrymal secretion is simply to keep the ' Bernard, Lectures on the Blood. Atlee's translation, p. 25. lb 274 SECRETION. surfaces of the cornea and conjunctiva moist and polished, and to preserve in this way the transparency of the parts. The tears, which are constantly secreted, are spread out uniformly over the anterior part of the eyeball by the movement of the lids in wink- ing, and are gradually conducted to the inner angle of the eye. Here they are taken up by the puncta lachrymalia, pass through the lachrymal canals, and are finally discharged into the nasal pas- sages beneath the inferior turbinated bones. A constant supply of fresh fluid is thus kept passing over the transparent parts of the eyeball, and the bad results avoided Avhich would follow from its accumulation and putrefactive alteration. 5. The Milk. — The mammary glands are conglomerate glands, resembling closely in their structure the pancreas, the salivary, and the lachrymal glands. They consist of numerous secreting sacs or follicles, grouped together in lobules, each lobule being supplied with a common excretory duct, which joins those coming from adjacent parts of the gland. (Fig. 105.) In this way, by their successive union, they form larger branches and trunks, until they are reduced in numbers to some 15 or 20 cylindrical ducts, the lactife- rous ducts, which open finally by as many minute orifices upon the extremity of the nipple. The secretion of the milk becomes fairly established at the end of ^two or three days after delivery, though the breasts often contain a milky fluid during the latter part of pregnancy. At first, the fluid dis- charged from the nipple is a yellowish turbid mixture, which is called the colostrum. It has the appearance of being thinner than the milk, but chemical examinations have shown' that it really con- tains a larger amount of solid ingredients than the perfect secre- tion. When examined under the microscope it is seen to contain, beside the milk-globules proper, a large amount of irregularly glo- Gi.ANDULAR Structure of Mamma. ' Lehmann, op. cit., vol. ii. p. 63. THE MILK. 275 bular or oval bodies, from y^^-go to ^^^ of an inch in diameter, which are termed the *' colostrum corpuscles." (B'ig. 106.) These bodies are more yellow and opaque than the true milk- globules, as well as being very much larger. They have a well defined outline, and con- sist apparently of a group of minute oily granules or glo- bules, imbedded in a mass of organic substance. The milk-globules at this time are less abundant than after- ward, and of larger size, measuring mostly from ^^'^^ to Ys'oo o^ ^^ ^^^^ ^^ ^'^' meter. At the end of a day or two from its first appearance the colostrum ceases to be discharged, and is replaced by the true milky secretion. The milk, as it is discharged from the nipple, is a white, opaque fluid, with a slightly alkaline reaction, and a specific gravity of about 1030. Its proximate chemical constitution, according to Pereira and Lehmann, is as follows : — Composition of Cow's Milk. . . . 9,1^.2 44.8 31.3 47.7 1 Chlorides of sodium and potassium ..... Phosphates of soda and potass ...... i Pliosphate of lime " " magnesia " " iron Alkaline carbonates . Colostrum Corpuscles, with milk-globules ; from a woman, one day after delivery. Water Casein Butter Sugar Soda 6.0 1000.0 Human milk is distinguished from the above by containing less casein, and a larger proportion of oily and saccharine ingredients. The entire amount of solid ingredients is also somewhat less than in cow's milk. 276 SECRETION. The casein is one of the most important ingredients of the milk. It is an extremely nutritious, organic substance, which is held in a fluid form by union with the water of the secretion. Casein is not coagulable by heat, and, consequently, milk may be boiled without changing its consistency to any considerable extent. It becomes a little thinner and more fluid during ebullition, owing to the melt- ing of its oleaginous ingredients ; and a thin, membranous film forms upon its surface, consisting probably of a very little albumen which the milk contains, mingled with the casein. The addition of any of the acids, however, mineral, animal, or vegetable, at once coagulates the casein, and the milk becomes curdled. Milk is coagulated, furthermore, by the gastric juice in the natural process of digestion, immediately after being taken into the stomach ; and if vomiting occur soon after a meal containing milk, it is thrown off in the form of semi-solid, curd-like flakes. The mucous membrane of the calves' stomach, or rennet, also has the power of coagulating casein ; and when milk has been curdled in this way, and its watery, saccharine, and inorganic in- gredients separated by mechanical pressure, it is converted into cheese. The peculiar flavor of the different varieties of cheese depends on the quantity and quality of the oleaginous ingredients which have been entangled with the coagulated casein, and on the alterations which these sub- Fig. 107. stances have undergone by the lapse of time and ex- posure to the atmosphere. The sugar and saline sub- stances of the milk are in solution, together with the casein and water, forming a clear, colorless, homogene- ous, serous fluid. The but- ter, or oleaginous ingredient, however, is suspended in this serous fluid in the form of minute granules and globules, the true "milk globules." (Fig. 107.) These globules are nearly fluid at th3 temperature of the body, and have a perfectly circular out- line. In the perfect milk, they are very much more abundant and ?o°o o°o? O.oo, o o°» @:J&ro^L a o o°o ^ol^.'o " Oo 'O O ^ O O"^ oooo O O o^ o'oo °oO bPoOO ZoOo - - o Milk-Globules; from same woman as above, four days after delivery. Secretion fully established. THE MILK. 277 smaller in size than in the colostrum; as the largest of them are not over v^^V^ of an inch in diameter, and the greater number about YxrrriyTT of an inch. The following is the composition of the butter of cow's milk, according to Eobin and Verdeil: — Margarine . . . . . . . . . .68 Oleine 30 Butyrine .......... 2 100 It is the last of these ingredients, the butyrine, which gives the peculiar flavor to the butter of milk. The niilk-globules have sometimes been described as if each one were separately covered with a thin layer of coagulated casein or albumen. No such investing membrane, however, is to be seen. The milk-globules are simply small masses of semi-fluid fat, sus- pended by admixture in the watery and serous portions of the secretion, so as to make an opaque, whitish emulsion. They do not fuse together when they come in contact under the microscope, simply because they are not quite fluid, but contain a large pro- portion of margarine, which is solid at ordinary temperatures of the body, and is only retained in a partially fluid form by the oleine with which it is associated. The globules may be made to fuse with each other, however, by simply heating the milk and subjecting it to gentle pressure between two slips of glass. "When fresh milk is allowed to remain at rest for twelve to twenty- four hours, a large portion of its fatty matters rise to the surface, and form there a dense and rich looking yellowish-white layer, the cream, which may be removed, leaving the remainder still opaline, but less opaque than before. At the end of thirty-six to forty-eight hours, if the weather be warm, the casein begins to take on a putrefactive change. In this condition it exerts a catalytic action upon the other ingredients of the milk, and particularly upon the sugar. A pure watery solution of milk-sugar {C^^S.^/)^) may be kept for an indefinite length of time, at ordinary temperatures, without undergoing any change. But if kept in contact with the partially altered casein, it suffers a catalytic transformation, and is converted into lactic acid (CgHgOg). This unites with the free soda, and decomposes the alkaline carbonates, forming lactates of soda and potass. After the neutralization of these substances has been accomplished, the milk loses its alkaline reaction and begins to turn sour. The free lactic acid then coagulates the casein, and the milk 278 SECRETION. is curdled. The altered organic matter also acts upon the oleagi- nous ingredients, which are partly decomposed; and the milk begins to give off a rancid odor, owing to the development of various volatile fatty acids, among which are butyric acid, and the like. These changes are very much hastened by a moderately elevated temperature, and also by a highly electric state of the atmosphere. The production of the milk, like that of other secretions, is liable to be much influenced by nervous impressions. It may be increased or diminished in quantity, or vitiated in quality by sudden emo- tions ; and it is even said to have been sometimes so much altered in this way as to produce indigestion, diarrhoea, and convulsions in the infant. Simon found' that the constitution of the milk varies from day to day, owing to temporary causes ; and that it undergoes also more permanent modifications, corresponding with the age of the infant. He analyzed the milk of a nursing woman during a period of nearly six months, commencing with the second day after delivery, and repeating his examinations at intervals of eight or ten days. It appears, from these observations, that the casein is at first in small quantity ; but that it increases during the first two months, and then attains a nearly uniform standard. The saline matters also increase in a nearly similar manner. The sugar, on the contrary, diminishes during the same period ; so that it is less abundant in the third, fourth, fifth and sixth months, than it is in the first and second. These changes are undoubtedly connected with the in- creasing development of the infant, which requires a corresponding alteration in the character of the food supplied to it. Finally, the quantity of butter in the milk varies so much from day to day, owing to incidental causes, that it cannot be said to follow any regular course of increase or diminution. 6. Secretion of the Bile. — The anatomical peculiarities in the structure of the liver are such as to distinguish it in a marked degree from the other glandular organs. Its first peculiarity is that it is furnished principally with venous blood. For, although it receives its blood from the hepatic artery as well as from the portal vein, the quantity of arterial blood with which it is supplied is extremely small in comparison with that which it receives from ' Op. cit., p. 337. SECRETION OF THE BILE. 279 the portal system. The blood which has circulated through the capillaries of the stomach, spleen, pancreas, and intestine is collected by the roots of the corresponding veins, and is discharged into the great portal vein, which enters the liver at the great transverse fissure of the organ. Immediately upon its entrance, the portal vein divides into two branches, right and left, which supply the corresponding portions of the liver ; and these branches success- ively subdivide into smaller twigs and ramifications, until they are reduced to the size, according to Kolliker, of ysVo of ^^ i^ch in diameter. These veins, with their terminal branches, are now arranged in such a manner as to include between them pentagonal or hexagonal spaces, or portions of the hepatic substance g'g to Jg of an inch in diameter in the human subject, which can readily be distinguished by the naked eye, both on the exterior of the organ and by the inspection of cut surfaces. The portions of hepatic substance included in this way between the terminal branches of the portal vein (Fig. 108) are termed the " acini" or the Fig- 108. "lobules" of the liver; and the terminal venous branches, occupying the spaces between the adjacent lobules, are the " interlobular" veins. In the spaces between the lobules we also find the minute branches of the hepatic ar- tery, and ,th e commencing rootlets of the hepatic^ducts. As the portal vein, the he- patic artery, and the hepatic duct enter the liver at the n -I Ramification of Portal Ye IN in Livkk. — a. transverse riSSUre, they are Twig of poital vein. &, 6. interlobular veins, c. Acini. closely invested by a fibrous sheath, Glisson's capsule, which accompanies them in their divisions and ramifications. In some of the lower animals, as in the pig, this sheath extends even to the interlobular spaces, inclosing each lobule in a -thin fibrous investment, by which it is distinctly separated from the neighboring parts. In the human subject, how- ever, Glisson's capsule becomes gradually thinner as it penetrates the liver, and disappears altogether before reaching the interlobular spaces; so that here the lobules are nearly in contact with each 280 SECEETION. other by their adjacent surfaces, being separated only by the inter- lobular veins and the minute branches «[' the hepatic artery and duct previously mentioned. From the sides of the interlobular veins, and also from their terminal extremities, there are given oft' capillary vessels, which penetrate the substance of each lobule and converge from its cir- cumference toward its centre, inosculating at the same time freely with each other, so as to form a minute vascular plexus, the " lobu- lar" capillary plexus. (Fig. 109.) At the centre of each lobule, the Fig. 109. Lobule of Liver, showing distribution of bloodvessels; magnified 22 diameters — a, a. In- terlobular veins. 6. Intralobular vein, c, c, c. Lobular capillary plexus, d, d. Twigs of inter- lobular vein passing to adjacent lobules. converging capillaries unite into a small vein (b), the " intralobu- lar" vein, which is one of the commencing rootlets of the hepatic vein. These rootlets, uniting successivelj'' with each other, so as to form larger and larger branches, finally leave the liver at its posterior edge, to empty into the ascending vena cava. Beside the capillary bloodvessels of the lobular plexus, each acinus is made up of an abundance of minute cellular bodies, about T2V0 of an inch in diameter, the "hepatic cells." (Fig. 110.) These cells have an irregularly pentagonal figure, and a soft consistency. They are composed of a homogeneous organic substance, in the midst of which are imbedded a large number of minute granules, and generally several well defined oil-globules. There is also a round or oval nucleus, with a nucleolus, imbedded in the substance SECRETION OF THE BILE. 281 of the cell, sometimes more or less obscured bj the granules and oil drops with which it is surrounded. The exact mode in which these cells are connected with the hepatic duct was for a long time the most obscure point in the minute anatomy of the liver. It has now been ascertained, Fig. 110. however, by the researches of Dr. Leidy, of Philadelphia,' and Dr. Beale, of London,^ that they are really contained in the interior of secreting tubules, which pass off from the smaller hepatic ducts, and penetrate everywhere the substance of the lobules. The cells fill nearly or com- pletely the whole cavity of the tubules, and the tubules themselves lie in close proxi- mity with each other, so as to leave no space between them except that which is occupied by the capillary bloodvessels of the lobular plexus. These cells are the active agents in accomplishing the function of the liver. It is by their influence that the blood which is brought in contact with them suffers certain changes which give rise to the secreted products of the organ. The ingredients of the bile first make their appearance in the substance of the cells. They are then transuded from one to the other, until they are at last dis- charged into the small biliary ducts seated in the interlobular spaces. Each lobule of the liver must accordingly be regarded as a mass of secreting tubules, lined with glandular cells, and invested with a close network of capillary bloodvessels. It follows, there- fore, from the abundant inosculation of the lobular capillaries, and the manner in which they are entangled with the hepatic tissue, that the blood, in passing through the circulation of the liver, comes into the most intimate relation with the glandular cells of the organ, and gives up to them the nutritious materials which are afterward converted into the iuOTedients of the bile. Hepatic Cells. From the human subject. American Journal Med. Sci., January, 1848. On Some Points in the Minute Anatomy of tlie Liver. London, 1856. 282 EXCKETION CHAPTER XVI. EXCRETION. "We have now come to the last division of the great nutritive function, viz., the process of excretion. In order to understand fairly the nature of this process we must remember that all the component parts of a living organism are necessarily in a state of constant change. It is one of the essential conditions of their existence and activity that they should go through with this incessant transforma- tion and renovation of their component substances. Every living animal and vegetable, therefore, constantly absorbs certain materials from the exterior, which are modified and assimilated by the pro- cess of nutrition, and converted into the natural ingredients of the organized tissues. But at the same time with this incessant growth and supply, there goes on in the same tissues an equally incessant process of waste and decomposition. For though the elements of the food are absorbed by the tissues, and converted into musculine, osteine, haematine and the Hire, they do not remain permanently in this condition, but almost immediately begin to pass over, by a con- tinuance of the alterative process, into new forms and combinations, which are destined to be expelled from the body, as others continue to be absorbed. Thus Spallanzani and Edwards found that every organized tissue not only absorbs oxygen from the atmosphere and fixes it in its own substance; but at the same time exhales carbonic acid, which has been produced by internal metamorphosis. This process, by which the ingredients of the organic tissues, al- ready formed, are decomposed and converted into new substances, is called the process of Destructive Assimilation. Accordingly we find that certain substances are constantly mak- ing their appearance in the tissues and fluids of the body, which did not exist there originally, and which have not been introduced with the food, but which have been produced by the process of in- ternal metamorphosis. These substances represent the waste, or physiological detritus of the animal organism. They are the forms EXCRETION, 283 under wbich those materials present themselves, which have once formed a part of the living tissues, but which have become altered by the incessant changes characteristic of organized bodies, and which are consequently no longer capable of exhibiting vital pro- perties, or of performing the vital functions. They are, therefore, destined to be removed and discharged from the animal frame, and are known accordingly by the name oi Excrementitious Substances. These excrementitious substances have peculiar characters by which they may be distinguished from the other ingredients of the living body; and they might, therefore, be made to constitute a fourth class of proximate principles, in addition to the three which we have enumerated in the preceding chapters. They are all sub- stances of definite chemical composition, and all susceptible of crystallization. Some of the most important of them contain nitro- gen, while a few are non-nitrogenous in their composition. They originate in the interior of living bodies, and are not found else- where, except occasionally as the result of decomposition. They are nearly all soluble in water, and are soluble without exception in the animal fluids. They are formed in the substance of the tissues, from which they are absorbed by the blood, to be afterward conveyed by the circulating fluid to certain excretory organs, particularly the kidneys, from which they are finally discharged and expelled from the body. This entire process, made np of the production of the excrementitious substances, their absorption by the blood, and their final elimination, is known as the process of excretion. The importance of this process to the maintenance of life is readily shown by the injurious effects which follow upon its disturbance. If the discharge of the excrementitious substances be in any way impeded or suspended, these substances accumulate, either in the blood or in the tissues or in both. In consequence of this retention and accumulation, they become poisonous, and rapidly produce a derangement of the vital functions. Their influence is principally exerted upon the nervous system, through which they produce most frequently irritability, disturbance of the special senses, deli- rium, insensibility, coma, and finally death. The readiness with which these effects are produced depends on the character of the excrementitious substance, and the rapidity with whicli it is pro- duced in the body. Thus, if the elimination of carbonic acid be stopped, by overloading the atmosphere with an abundance of the same gas, death takes place at the end of a few minutes ; but if the elimination of urea by the kidneys be checked, it requires three or 284 EXCRETION. four days to produce a fatal result. A fatal result, however, is cer- tain to follow, at the end of a longer or shorter time, if any one of these substances be compelled to remain in the body, and accumu- late in the animal tissues and fluids. The principal excrementitious substances known to exist in the human body are as follows ; 1. Carbonic acid 2. Cholesterine 3. Urea . 4. Creatine 5. Creatinine . 6. Urate of soda 7. Urate of potass 8. Urate of ammon CO, C2 H2,0 C^H.N^O^ C.HgNaO, CgH.NjO.^ NaO.C.HN^Oj+HO KO.CjHN.O^ NH,0,2C5HN202-fHO Of these substances the first two have already been studied at sufficient length in the preceding chapters. We will merely repeat here that carbonic acid is produced in large quantity in nearly all the tissues of the body, from which it is absorbed by the blood, conveyed to the lungs, and there exhaled at the sarrie time that oxygen is absorbed. Cholesterine is a non-saponifiable fatty sub- stance, originating in the brain and spinal cord, in the tissue of which organs it exists in the proportion of 58 parts per thousand. It is thence taken up by the blood, conveyed to the liver and dis- charged with the bile. Cholesterine is extremely insoluble in water, but is held in solution in the blood and the bile, by some of the other ingredients present in these animal fluids. The remaining excrementitious substances ma}^ be examined together, with the more propriety, since they are all ingredients of a single excretory fluid, viz., the urine. UREA. This is a neutral, crystallizable, nitrogenous substance, very readily soluble in water, and easily decomposed by various external influences. It occurs in the urine in the proportion of 30 parts per thousand; and in the blood in much smaller quantity. The blood, however, is the source from which this substance is sup- plied to the urine ; and it exists, accordingly, in but small quantity in the circulating fluid, for the reason that it is constantly drained off by the kidneys. But if the kidneys be extirpated, or the renal arteries tied, or the excretion of urine suspended by inflammation UEEA. 285 or otherwise, the urea then accumulates in the blood, and presents itself there in considerable quantity. It has been found in the blood, under these circum- stances, in the proportion of Fig- m. 1.4 per thousand.' It is not yet known from what source the urea is originally derived ; whether it be produced in the blood itself, or whether it be formed in some of the solid tissues, and thence taken up by the blood. It has not yet been found, however, in any of the solid tissues, in a state of health. Urea is obtained most readily from the urine. For this pur- pose the fresh urine is evapo- rated, in the water bath until it has a syrupy consistency. It is then mixed with an equal volume of nitric acid, which forms nitrate of urea. This salt, being less soluble than pure urea, rapidly crystallizes, after which it is separated by filtration from the other ingredients. It is then dissolved in water and decomposed by carbonate of lead, forming nitrate of lead which remains in solution, and carbonic acid which escapes. The solution is then evaporated, the urea dissolved out by alcohol, and finally crystallized in a pure state. Urea has no tendency to spontaneous decomposition, and may be kept, when perfectly pure, in a dry state or dissolved in water, for an indefinite length of time. If the watery solution be boiled, however, the urea is converted, during the process of ebullition, into carbonate of ammonia. One equivalent of urea unites with two equivalents of water, and becomes transformed into two equiva- lents of carbonate of ammonia, as follows : — Urea, prepared from urine, and crystallized by slow evaporation. (After Lehmann.) C,H^N202=Urea. H, 0,=Water. NH3,C02=Carbonate of ammonia. 2 Various impurities, also, by acting as catalytic bodies, will in- duce the same change, if water be present. Animal substances in a state of commencing decomposition are particularly liable to act Robin and Verdeil, vol. ii. p. 502. 286 EXCRETION. in this way. In order that the conversion of the urea be thus produced, it is necessary that the temperature of the mixture be not far from 70° to 100° F. The quantity of urea produced and discharged daily by a healthy adult is, according to the experiments of Lehmann, about 500 grains. It varies to some extent, like all the other secreted and excreted products, with the size and development of the body. Lehmann, in experiments on his own person, found the average daily quantity to be 487 grains. Dr. William A. Hammond,' of the U. S. Array, who is a very large man, by similar experiments found it to be 670 grains. Dr. John C. Draper'^ found it 408 grains. No urea is to be detected in the urine of very young children ;^ but it soon makes its appearance, and afterward increases in quantity with the development of body. The daily quantity of urea varies also with the degree of mental and bodily activity. Lehmann and Hammond both found it very sensibly increased by muscular exertion and diminished by repose. It has been thought, from these facts, that this substance must be directly produced from disintegration of the muscular tissue. This, however, is by no means certain ; since in a state of general bodily activity it is not only the urea, but the excretions generally, carbonic acid, perspiration, &c., which are increased in quantity simultane- ously with each other. Hammond has also shown that continued mental application will raise the quantity of urea above its normal standard, though the muscular system remain comparatively in- active. The quantity of urea varies also with the nature of the food. Lehmann, by experiments on his own person, found that the quan- tity was larger while living exclusively on animal food than with a mixed, or vegetable diet ; and that its quantity was smallest when confined to a diet of purely non-nitrogenous substances, as starch, sugar, and oil. The following table* gives the result of these ex- periments. Kind of Food. Daily Quantity of Urea. Animal 798 grains. Mixed 487 " Vegetable 337 " Non-nitrogenous ...... 231 " ' American Journal Med. Sci., Jan., 1855, and April, 1856. ^ New York Journal of Medicine, March, 1856. ^ Robin and Veideil, vol. ii. p. 500. '' Lehmann, op. cit., vol. ii. p. 163. CREATINE, 287 Finally, it has been shown by Dr. John C. Draper' that there is also a diurnal variation in the normal quantity of urea. A smaller quantity is produced during the night than during the day ; and this difference exists even in patients who are confined to the bed during the whole twenty-four hours, as in the case of a man under treatment for fracture of the leg. This is probably owing to the greater activity, during the waking hours, of both the mental and digestive functions. More urea is produced in the latter half than in the earlier half of the day; and the greatest quantity is dis- charged during the four hours from 6| to lOJ P. M. Urea exists in the urine of the carnivorous and many of the herbivorous quadrupeds ; but there is little or none to be found in that of birds and reptiles. CREATINE. This is a neutral crystallizable substance, found in the muscles, the blood, and the urine. It is soluble in water, very slightly solu- ble in alcohol, and not at all so in ether. By boiling with ^'S* ^^?: an alkali, it is either converted into carbonic acid and ammonia, or is decomposed with the pro- duction of urea and an artificial nitrogenous crystallizable sub- stance, termed sarcosine. By being heated with strong acids, it loses two equivalents of water, and is converted into the sub- stance next to be described, viz., creatinine. Creatine exists in the urine, in the human subject, in the proportion of about 1.25 parts, and in the muscles in the proportion of 0.67 parts per thousand. Its quantity in the blood has not been determined. In the muscu- lar tissue it is simply in solution in the interstitial fluid of the parts, so that it may be extracted by simply cutting the muscle into Creatine, crystallized from hot water. (After Lehmauu ) ' Loc. cit. 288 EXCRETION. small pieces, treating it with distilled water, and subjecting it to pressure. Creatine evidently originates in the muscular tissue, is absorbed thence bj the blood, and is finally discharged with the urine. CREATININE. This is also a crystallizable substance. It differs in composition from creatine by containing two equivalents less of the elements of water. It is more soluble in water and in spirit than creatine, and dissolves slightly also in ether. It has a distinctly alkaline reaction. It occurs, like crea- tine, in the muscles, the blood, §' and the urine ; and is undoubt- edly first produced in the muscular tissue, to be dis- charged finally by the kidneys. It is very possible that it ori- ginates, not directly from the muscles, but indirectly, by trans- formation of a part of the crea- tine; since it may be artificially produced, as we have already mentioned, by transformation of the latter substance under the influence of strong acids, and since, furthermore, while creatine is more abundant in the muscles than creatinine, in the urine, on the contrary, there is a larger quantity of creatinine than of creatine. Both these substances have been found in the muscles and in the urine of the lower animals. C RE AT IX IN E, crystallized from hot water. (Afier Lehmann.) URATE OF SODA. As its name implies, this substance is a neutral salt, formed by the union of soda, as a base, with a nitrogenous animal acid, viz., uric acid {G^^Jd^^O). Uric acid is sometimes spoken of as though it were itself a proximate principle, and a constituent of the urine; but it cannot properly be regarded as such, since it never occurs in URATES OF POTASS AND AMMONIA. 289 a free state, in a natural condition of the fluids. When present, it has always been produced by decomposition of the urate of soda. Urate of soda is readily soluble in hot water, from which a large portion again deposits on cooling. It is slightly soluble in alcohol, and insoluble in ether. It crystallizes in small globular '^" masses, with projecting, curv- ed, conical, wart-like excres- cences. (Fig. 114.) It dis- solves readily in the alkalies; and by most acid solutions it is decomposed, with the pro- duction of free nric acid. Urate of soda exists in the urine and in the blood. It is either produced originally in the blood, or is formed in some of the solid tissues, and absorbed from them by the circulating fluid. It is con- stantly eliminated by the kidneys, in company with the other ingre- dients of the urine. The average daily quantity of urate of soda discharged by the healthy human subject is, according to Lehmann, about 25 grains. This substance exists in the urine of the carnivo- rous and omnivorous animals, but not in that of the herbivora. In the latter, it is replaced by another substance, differing some- what from it in composition and properties, viz., hippurate of soda. The urine of herbivora, however, while still very young, and living upon the milk of the, mother, has been found to contain urates. But when the young animal is weaned, and becomes herbivorous, the urate of soda disappears, and is replaced by the hippurate. TJeate of Soda: from a urinary deposit. URATES OF POTASS AND AMMONIA. The urates of potass and ammonia resemble the preceding salt very closely in their physiological relations. They are formed in very much smaller quantity than the urate of soda, and appear like it as ingredients of the urine. The substances above enumerated resemble each other closely in 19 290 EXCRETION. their most striking and important characters. They all contain nitrogen, are all crystallizable, and all readily soluble in water. They all originate in the interior of the body by the decomposition or catalytic transformation of its organic ingredients, and are all conveyed by the blood to the kidneys, to be finally expelled with the urine. These are the substances which represent, to a great extent, the final transformation of the organic or albuminoid in- gredients of the tissues. It has already been mentioned, in a pre- vious chapter, that these organic or albuminoid substances are not discharged from the body, under their own form, in quantity at all proportionate to the abundance with which they are introduced. By far the greater part of the mass of the frame is made up of organic substances: albumen, musculine, osteine, &c. Similar materials are taken daily in large quantity with the food, in order to supply the nutrition and waste of those already composing the tissues; and yet only a very insignificant quantity of similar material is expelled with the excretions. A minute proportion of volatile animal matter is exhaled with the breath, and a minute proportion also with the perspiration. A very small quantity is discharged under the form of mucus and coloring matter, with the urine and feces ; but all these taken together are entirely insuffi- cient to account for the constant and rapid disappearance of organic matters in the interior of the body. These matters, in fact, before being discharged, are converted by catalysis and decomposition into new substances. Carbonic acid, under which form 3500 grains of carbon are daily expelled from the body, is one of these substances ; the others are urea, creatine, creatinine, and the urates. We see, then, in what way the organic matters, in ceasing to form a part of the living body, lose their characteristic properties, and are converted into crystallizable substances, of definite chemical composition. It is a kind of retrograde metamorphosis, by which they return more or less to the condition of ordinary inorganic materials. These excrementitious matters are themselves decom- posed, after being expelled from the body, under the influence of the atmospheric air and moisture ; so that the decomposition and destruction of the organic substances is at last complete. It will be seen, consequently, that the urine has a character altogether peculiar, and one which distinguishes it completely from every other animal fluid. All the others are either nutritive fluids, like the blood and milk, or are destined, like the secretions GENERAL CHARACTERS OF THE URINE. 291 generally, to take some direct and essential part in the vital opera- tions. Many of them, like the gastric and pancreatic juices, are reabsorbed after they have done their work, and again enter the current of the circulation. But the urine is merely a solution of excrementitious substances. Its materials exist beforehand in the circulation, and are simply drained away by the kidneys from the blood. There is a wide difference, accordingly, between the action of the kidneys and that of the true glandular organs, in which certain new and peculiar substances are produced by the action of the glandular tissue. The kidneys, on the contrary, do not secrete anything, properly speaking, and are not, therefore, glands. In their mode of action, so far as regards the excretory function, they have more resemblance to the lungs than to any other of the internal organs. But this resemblance is not complete ; since the lungs perform a double function, absorbing oxygen at the same time that they exhale carbonic acid. The kidneys alone are purely excretory in their office. The urine is not intended to fulfil any function, mechanical, chemical, or otherwise ; but is des- tined only to be eliminated and expelled. Since it possesses so peculiar and important a character, it will require to be carefully studied in detail. The urine is a clear, watery, amber-colored fluid, with a distinct acid reaction. It has, while still warm, a peculiar odor, which dis- appears more or less completely on cooling, and returns when the urine is gently heated. The ordinary quantity of urine discharged daily by a healthy adult is about ^xxxv, and its mean specific gravity, 1024. Both its total quantity, however, and its mean specific gravity are liable to vary somewhat from day to day, owing to the different proportion of water and solid ingredients entering into its constitution. Ordinarily the water of the urine is more than sufficient to hold all its solid matters in solution ; and its pro- portion may therefore be diminished by accidental causes without the urine becoming turbid by the formation of a deposit. Under such circumstances, it merely becomes deeper in color, and of a higher specific gravity. Thus, if a smaller quantity of water than usual be taken into the system with the drink, or if the fluid ex- halations from the lungs and skin, or the intestinal discharges, be increased, a smaller quantity of water will necessarily pass off by the kidneys; and the urine will be diminished in quantity, while its specific gravity is increased. We have observed the urine to be 292 EXCEETION. reduced in this way to eighteen or twenty ounces per day, its specific gravity rising at the same time to 1030. On the other hand, if the fluid ingesta be unusually abundant, or if the perspiration be dimi- nished, the surplus quantity of water will pass oif by the kidneys; so that the amount of urine in twenty-four hours may be increased to forty-five or forty-six ounces, and its specific gravity reduced at the same time to 1020 or even 1017. Under these conditions the total amount of solid matter discharged daily remains about the same. The changes above mentioned depend simply upon the fluctuating quantity of water, which may pass off by the kidneys in larger or smaller quantity, according to accidental circumstances. In these purely normal or physiological variations, therefore, the entire quantity of the urine and its mean specific gravity vary always in an inverse direction with each other ; the former increas- ing while the latter diminishes, and vice versa. If, however, it should be found that both the quantity and specific gravity of the urine were increased or diminished at the same time, or if either one were increased or diminished while the other remained station- ary, such an alteration would show an actual change in the total amount of solid ingredients, and would indicate an unnatural and pathological condition. This actually takes place in many forms of disease. The amount of variation in the quantity of water, even, may be so great as to constitute by itself a pathological condition. Thus, in hysterical attacks there is sometimes a very abundant flow of limpid, nearly colorless urine, with a specific gravity not over 1005 or 1006. On the other hand, in the onset of febrile attacks, the quantity of water is often so much diminished that it is no longer sufficient to retain in solution all the solid ingredients of the urine, and the urate of soda is thrown down, after cooling, as a fine red or yellowish sediment. So long, however, as the variation is con- fined within strictly physiological limits, all the solid ingredients are held in solution, and the urine remains clear. There is also, in a state of health, a diurnal variation of the urine, both in regard to its specific gravity and its degree of acidity. The urine is generally discharged from the bladder five or six times during the twenty-four hours, and at each of these periods shows more or less variation in its physical characters. We have found that the urine which collects in the bladder during the night, and is discharged the first thing in the morning, is usually dense, DIURNAL YARIATI0X3 OF THE URINE. 293 highly colored, of a strongly acid reaction, and a high specific gravity. That passed during the forenoon is pale, and of a low specific gravit}'', sometimes not more than 1018 or even 1015. It is at the same time neutral or slightly alkaline in reaction. Toward the middle of the day, its density and depth of color increase, and its acidity returns. All these properties become more strongl}- marked during the afternoon and evening, and toward night the urine is again deeply colored and strongly acid, and has a specific gravity of 1028 or 1030. The following instances will serve to show the general characters of this variation : — Observation First. March 20 fh. Urine of 1st discharge, acid, sp. gr. 1025. " 2d " alkaline, " 1015. •' 3d " neutral, " 1018. " 4th " acid, " 1018. " 5th " acid, " 1027. Observation Second. March 21st. Urine of 1st discharge, acid, sp. gr. 1029. " 2d " neutral, " 1022. " 3d " neutral, " 1025. " 4th " acid, " 1027. « 5th " acid, " 1030. These variations do not always follow the perfectly regular course manifested in the above instances, since they are somewhat liable, as we have already mentioned, to temporary modification from accidental causes during the day ; but their general tendency nearly always corresponds with that given above. It is evident, therefore, that whenever we wish to test the specific gravity and acidity of the urine in cases of disease, it will not be sufficient to examine any single specimen taken at random ; but all the different portions discharged during the day should be collected and examined together. Otherwise, we should incur the risk of regarding as a permanently morbid symptom what might be nothing more than a purely accidental and temporary variation. The chemical constitution of the urine as it is discharged from the bladder, according to the analyses of Berzelius, Lehmann, Becquerel, and others, is as follows : — 294 EXCKETION. Composition of the Urine. Water 938.00 Urea 30.00 Creatine ........... 1.25 Creatinine .......... 1.50 Urate of soda -^ " potass I 1.80 " ammonia J Coloring matter and •> o^ Mucus J Biphosphate of soda -\ Phosphate of soda j " potass I- 12.45 " magnesia I " lime J Chlorides of sodium and potassium 7.80 Sulphates of soda and potass ....... 6.90 1000.00 We need not repeat that the proportionate quantity of these different ingredients, as given above, is not absolute, but only approximative ; and that they vary, from time to time, within cer- tain physiological limits, like the ingredients of all other animal fluids. The urea, creatine, creatinine, and urates have all been suffi- ciently described above. The mucus and coloring matter, unlike the other ingredients of the urine, belong to the class of organic substances proper. They are both present, as may be seen by the analysis quoted above, in very small quantity. The coloring matter, or urosacine, is in solution in a natural condition of the urine, but is apt to be entangled by any accidental deposits which may be thrown down, and more particularly by those consisting of the urates. These deposits, from being often strongly colored red or pink by the urosacine thus thrown down with them, are known under the name of " brick-dust" sediments. The mvcus of the urine comes from the lining membrane of the urinary bladder. When first discharged it is not visible, owing to its being uniformly disseminated through the urine by mechanical agitation ; but if the fluid be allowed to remain at rest for some hours in a cylindrical glass vessel, the mucus collects at the bottom, and may then be seen as a light cottony cloud, interspersed often with minute semi-opaque points. It plays, as we shall hereafter see a very important part in the subsequent fermentation and decomposition of the urine. EEACTIONS OF THE URINE. 295 Biphosphate of soda exists in the urine by direct solution, since it is readily soluble in water. It is this salt which gives to the urine its acid reaction, as there is no free acid present in the recent condition. It is probably derived from the neutral phosphate of soda in the blood, which is decomposed by the uric acid at the time of its for- mation ; producing, on the one hand, a urate of soda, and converting a part of the neutral phosphate of soda into the acid biphosphate. The phosphates of lime and magnesia, or the " earthy phosphates," as they are called, exist in the urine by indirect solution. Though insoluble, or very nearly so, in pure water, they are held in solu- tion in the urine by the acid phosphate of soda, above described. They are derived from the blood, in which they exist in considera- ble quantity. When the urine is alkaline, these phosphates are deposited as a light colored precipitate, and thus communicate a turbid appearance to the fluid. When the urine is neutral, they may still be held in solution, to some extent, by the chloride of sodium, which has the property of dissolving a small quantity of phosphate of lime. The remaining ingredients, phosphates of soda and potass, sul- phates and chlorides, are all derived from the blood, and are held directly in solution by the water of the urine. The urine, constituted by the above ingredients, forms, as we have already described, a clear amber colored fluid, with a reaction for the most part distinctly acid, sometimes neutral, and occasion- ally slightly alkaline. In its healthy condition it is affected by chemical and physical reagents in the following manner. Boiling the urine does not produce any visible change, provided its reaction be acid. If it be neutral or alkaline, and if, at the same time, it contain a larger quantity than usual of the earthy phos- phates, it will become turbid on boiling ; since these salts are less soluble at a high than at a low temperature. The addition of nitric, or other mineral acid, produces at first only a slight darkening of the color, owing to the action of the acid upon the organic coloring matter of the urine. If the mixture, however, be allowed to stand for some time, the urates of soda, potass, &c., will be decomposed, and pure uric acid, which is very insoluble, will be deposited in a crystalline form upon the sides and bottom of the glass vessel. The crystals of uric acid have most frequently the form of transparent rhomboidal plates, or oval laminae with pointed extremities. They are usually tinged of a yellowish hue by the coloring matter of the urine which is entangled with them 296 EXCEETION. Uric Acid; deposited from urine. at the time of their deposit. They are frequently arranged in radiated clusters, or small spheroidal masses, so as to present the appearance of minute calcu- Fig- 115- lous concretions. (Fig. 115.) The crystals vary very much in size and regularity, ac- cording to the time occupied in their formation. If a free alkali, such as potass or soda, be added to the urine, so as to neutralize its acid reaction, it becomes immediately turbid from a deposit of the earthy phos- phates, which are insoluble in alkaline fluids. The addition of nitrate of baryta, chloride of barium, or subacetate of lead to healthy urine, produces a dense precipi- tate, owing to the presence of the alkaline sulphates. Nitrate of silver produces a precipitate with the chlorides of sodium and potassium. Subacetate of lead and nitrate of silver precipitate also the or- ganic substances, mucus and coloring matter, present in the urine. All the above reactions, it will be seen, are owing to the presence of the natural ingredients of the urine, and do not, therefore, indi- cate any abnormal condition of the excretion. Besides the properties mentioned above, the urine has several others which are of some importance, and which have not been usually noticed in previous descriptions. It contains, among other ingredients, certain organic substances which have the power of in- terfering with the mutual reaction of starch and iodine, and even of decomposing the iodide of starch, after it has once been formed. This peculiar action of the urine was first noticed and described by us in 1856.^ If 5j of iodine water be mixed with a solution of starch, it strikes an opaque blue color ; but if 5j of fresh urine be afterward added to the mixture, the color is entirely destroyed at the end of four or five seconds. If fresh urine again be mixed with four or five times its volume of iodine water, and starch be ' American Journal Med. Sci., April, 1856. ACCIDENTAL INGEEDIENTS OF THE UKINE. 297 subsequently added, no union takes place between the starch and iodine, and no blue color is produced. In these instances, the iodine unites with the animal matters of the urine in preference to com- bining with the starch, and is consequently prevented from striking its ordinary blue color with the latter. This interference occurs whether the urine be acid or alkaline in reaction. In all cases in which iodine exists in the urine, as for example where it has been administered as a medicine, it is under the form of an organic com- bination ; and in order to detect its presence by means of starch, a few drops of nitric acid must be added at the same time, so as to destroy the organic matters, after which the blue color immediately appears, if iodine be present. This reaction with starch and iodine belongs also, to some extent, to most of the other animal fluids, as the saliva, gastric and pancreatic juices, serum of the blood, &c. ; but it is most strongly marked in the urine. Another remarkable property of the urine, also dependent on its organic ingredients, is that of interfering with Trommer's test for grape sugar. If clarified honey be mixed with fresh urine, and sul- phate of copper with an excess of potass be afterward added, the mixture takes a dingy, grayish-blue color. On boiling, the color turns yellowisb or yellowish-brown, but the suboxide of copper is not deposited. In order to remove the organic matter and detect the sugar, the urine must be first treated with an excess of animal charcoal and filtered. By this means the organic substances are retained upon the filter, while the sugar passes through in solution, and may then be detected as usual by Trommer's test. Accidental Ingeedients of the Ueine. — Since the urine, in its natural state, consists of materials which are already prepared in the blood, and which merely pass out through the kidneys by a kind of filtration, it is not surprising that most medicinal and poisonous substances, introduced into the circulation, should be expelled from the body by the same channel. Those substances which tend to unite strongly with the animal matters, and to form with them insoluble compounds, such as the preparations of iron, lead, silver, arsenic, mercury, &c., are least liable to appear in the urine. They may occasionally be detected in this fluid when they have been given in large doses, but when administered in moderate quantity are not usually to be found there. Most other substances, however, accidentally present in the circulation, pass off readily by 298 EXCRETION. the kidneys, either in their original form, or after undergoing cer- tain chemical modifications. The salts of the organic acids, such as lactates^ acetates, malates, &G., of soda and potass, when introduced into the circulation, are replaced by the carbonates of the same bases, and appear under that form in the urine. The urine accordingly becomes alkaline from the presence of the carbonates, whenever the above salts have been taken in large quantity, or after the ingestion of fruits and vegetables which contain them. We have already spoken (Chap. II.) of the experiments of Lehmann, in which he found the urine exhi- biting an alkaline reaction, a very few minutes after the adminis- tration of lactates and acetates. In one instance, by experimenting upon a person with congenital extroversion of the bladder, in whom the orifices of the ureters were exposed,^ he found that the urine became alkaline in the course of seven minutes after the ingestion of half an ounce of acetate of potass. The 2^ure alkalies and their carbonates, according to the same ob- server, produce a similar effect. Bicarbonate of potass, for example, administered in doses of two or three drachms, causes the urine to become neutral in from thirty to forty-five minutes, and alkaline in the course of an hour. It is in this way that certain "anti-cal- culous" or " anti-lithic" nostrums operate, when given with a view of dissolving concretions in the bladder. These remedies, which are usually strongly alkaline, pass into the urine, and by giving it an alkaline reaction, produce a precipitation of the earthy phos- phates. Such a precipitate, however, so far from indicating the successful disintegration and discharge of the calculus, can only tend to increase its size by additional deposits. Ferrocyanide of potassium, when introduced into the circulation, appears readily in the urine. Bernard^ observed that a solution of this salt, after being injected into the duct of the submaxillary gland, could be detected in the urine at the end of twenty minutes. Iodine^ in all its combinations, passes out by the same channel. We have found that after the administration of half a drachm of the syrup of iodide of iron, iodine appears in the urine at the end of thirty minutes, and continues to be present for nearly twenty- four hours. In the case of two patients who had been taking iodide of potassium freel}--, one of them for two months, the other for six ' Physiological Chemistry, vol. ii. p. 133. 2 Leqons de Physiologie Experimentale, 1855, vol. ii. p. 111. ACCIDENTAL INGREDIENTS OF THE URINE. 299 weeks, the urine still contained iodine at the end of three days after the suspension of the medicine. In three days and a half, however, it was no longer to be detected. Iodine appears also, after being introduced into the circulation, both in the saliva and the perspiration. Quinine^ when taken as a remedy, has also been detected in the urine. Ether passes out of the circulation in the same way. We have observed the odor of this substance very perceptible in the urine, after it had been inhaled for the purpose of producing anees- thesia. The bile-pigment passes into the urine in great abundance in some cases of jaundice, so that the urine may have a deep yellow or yellowish brown tinge, and may even stain linen clothes, with which it comes in contact, of a similar color. The saline biliary substances, viz., glyko-cholate and tauro-cholate of soda, have occa- sionally, according to Lehmann, been also found in the urine. In these instances the biliary matters are reabsorbed from the hepatic ducts, and afterward conveyed by the blood to the kidneys. Sugar. — When sugar exists in unnatural quantity in the blood, it passes out with the urine. We have repeatedly found that if sugar be artificially introduced into the circulation in rabbits, or injected into the subcutaneous areolar tissue so as to be absorbed by the blood, it is soon discharged by the kidneys. It has been shown by Bernard* that the rapidity with which this substance appears in the urine under these circumstances varies with the quantity injected and the kind of sugar used for the experiment. If a solution of 15 grains of glucose be injected into the areolar tissue of a rabbit weighing a little over two pounds, it is entirely destroyed in the cir- culation, and does not pass out with the urine. A dose of 23 grains, however, injected in the same way, appears in the urine at the end of two hours, 30 grains in an hour and a half, 38 grains in an hour, and 188 grains in fifteen minutes. Again, the kind of sugar used causes a difference in this respect. For while 15 grains of glucose may be injected without passing out by the kidneys, 7J grains of cane sugar, introduced in the same way, fail to be completely de- stroyed in the circulation, and may be detected in the urine. In certain forms of disease (diabetes), where sugar accumulates in the blood, it is eliminated by the same channel ; and a saccharine condition of the urine, accompanied by an increase in its quantity ' Legons de Phys. Exp., 1855, vol. i. p. 214 et seq. 300 EXCEETION. and specific gravity, constitutes the most cbaracteristic feature of the disease. Finally, albumen sometimes shows itself in the urine in conse- quence of various morbid conditions. Most acute inflammations of the internal organs, as pneumonia, pleurisy, &c., are liable to be accompanied, at their outset, by a congestion of the kidneys, which produces a temporary exudation of the albuminous elements of the blood. Albumen has been found in the urine, according to Simon, Becquerel, and others, in pericarditis, pneumonia, pleurisy, bron- chitis, hepatitis, inflammation of the brain, peritonitis, metritis, &c. "We have observed it, as a temporary condition, in pneumonia and after amputation of the thigh. Albuminous urine also occurs fre- quently in pregnant women, and in those affected with abdominal tumors, where the pressure upon the renal veins is sufficient to produce passive congestion of the kidneys. When the renal con- gestion is spontaneous in its origin, and goes on to produce actual degeneration of the tissue of the kidneys, as in Bright's disease, the same symptom occurs, and remains as a permanent condition. In all such instances, however, as the above, where foreign ingre- dients exist in the urine, these substances do not originate in the kidneys themselves, but are derived from the blood, in the same manner as the natural ingredients of the excretion. Changes in the Urine during Decomposition, — When the urine is allowed to remain exposed, after its discharge, at ordinary temperatures, it becomes decomposed, after a time, like any other animal fluid ; and this decomposition is characterized by certain changes which take place in a regular order of succession, as fol- lows: — After a few hours of repose, the mucus of the urine, as we have mentioned above, collects near the bottom of the vessel as a light, nearly transparent, cloudy layer. This mucus, being an organic sub- stance, is liable to putrefaction ; and if the temperature to which it is exposed be between 60° and 100° F., it soon becomes altered and communicates these alterations, more or less rapidly, to the superna- tant fluid. The first of these changes is called the acid fermentation of the urine. It consists in the production of a free acid, usually lactic acid, from some of the undetermined animal matters con- tained in the excretion. This fermentation takes place very early ; within the first twelve, twenty-four, or forty-eight hours, according to the elevation of the surrounding temperature. Perfectly fresh ACID FERMENTATION OF THE URINE. 301 urine, as we have already stated, contains no free acid, its acid reaction to test paper being dependent entirely on the presence of biphosphate of soda. Lactic acid nevertheless has been so fre- quently found in nearly fresh urine as to lead some eminent chemists (Berzelius, Lehmann) to regard it as a natural constituent of the excretion. It has been subsequently found, however, that urine, though entirely free from lactic when first passed, may fre- quently present traces of this substance after some hours' exposure to the air. The lactic acid is undoubtedly formed, in these cases, by the decomposition of some animal substance contained in the urine. Its production in this way, though not constant, seems to be sufficiently frequent to be regarded as a normal process. In consequence of the presence of this acid, the urates are par- tially decomposed ; and a crystalline deposit of free uric acid slowly takes place, in the same manner as if a little nitric or muriatic acid had been artificially mixed with the urine. It is for this reason that urine which is abundant in the urates frequently shows a de- posit of crystallized uric acid some hours after it has been passed, though it may have been perfectly free from deposit at the time of its emission. During the period of the "acid fermentation," there is reason to believe that oxalic acid is also sometimes produced in a similar manner with the lactic. It is very certain that the deposit of oxa- late of lime, far from being a dangerous or even morbid symptom, as it was at one time regarded, is frequently present in perfectly normal urine after a day or two of exposure to the atmosphere. We have often observed it, under these circumstances, when no morbid symptom could be detected in connection with either the kidneys or any other bodily organ. Now, whenever oxalic acid is formed in the urine, it must necessarily be deposited under the form of oxalate of lime ; since this salt is entirely insoluble both in water and in the urine, even when heated to the boiling point. It is difficult to understand, therefore, when oxalate of lime is^ found as a deposit in the urine, how it can previously have been held in solution. Its oxalic acid is in all probability gradually formed, as we have said, in the urine itself; uniting, as fast as it is produced, with the lime previously in solution, and thus appearing as a crystalline deposit of oxalate of lime. It is much more probable that this is the true explanation, since, in the cases to which we allude, the crystals of oxalate of lime grow, as it were, in the cloud of mucus which collects at the bottom of the vessel, while the 302 EXCRETION. Oxalate op Lime; deposited from healthy urine, during the acid fevmentatiou. supernatant fluid remains clear. These crystals are of minute size, transparent, and colorless, ^'S- ^^^- and have tlie form of regular octohedra, or double quad- rangular pyramids, united base to base. (Fig. 116.) They make their appearance usu- ally about the commence- ment of the second day, the urine at the same time con- tinuing clear and retaining its acid reaction. This depo- sit is of frequent occurrence when no substance contain- ing oxalic acid or oxalates has been taken with the food. At the end of some days the changes above described come to an end, and are succeeded by a different process known as the alkaline fermentation. This consists essentially in the decompo- sition or metamorphosis of the urea into carbonate of ammonia. As the alteration of the mucus advances, it loses the power of pro- ducing lactic and oxalic acids, and becomes a ferment capable of acting by catalysis upon the urea, and of exciting its decomposition as above. We have already mentioned that urea may be converted into carbonate of ammonia by prolonged boiling or by contact with decomposing animal substances. In this conversion, the urea unites with the elements of two equivalents of water ; and conse- quently it is not susceptible of the transformation when in a dry state, but only when in solution or supplied with a sufficient quan- tity of moisture. The presence of mucus, in a state of incipient decomposition, is also necessary, to act the part of a catalytic body. Consequently if the urine when first discharged be passed through a succession of close filters, so as to separate and retain its mucus, it may be afterward kept, for an almost indefinite time, without altera- tion. But under ordinary circumstances, the mucus, as soon as its putrefaction has commenced, excites the decomposition of the urea, and carbonate of ammonia begins to be developed. The first portions of the ammoniacal salt thus produced begin to neutralize the biphosphate of soda, so that the acid reaction of the urine diminishes in intensity. This reaction gradually becomes ALKALINE FERMENTATION OF THE URINE. 303 weaker and weaker, as the fermentation proceeds, until it at last disappears altogether, and the urine becomes neutral. The produc- tion of carbonate of ammonia still continuing, the reaction of the fluid then becomes alkaline, and its alkalescence grows more strongly pronounced with the constant accumulation of the ammoniacal salt. The rapidity with which this alteration proceeds depends on the character of the urine, the quantity and quality of the mucus which it contains, and the elevation of the surrounding temperature. The urine passed early in the forenoon, which is often neutral at the time of its discharge, will of course become alkaline more readily than that which has at first a strongly acid reaction. In the summer, urine will become alkaline, if freely exposed, on the third, fourth, or fifth day ; while in the winter, a specimen kept in a cool place may still be neutral at the end of fifteen days. In cases of paralysis of the bladder, on the other hand, accompanied with cystitis, where the mucus is increased in quantity and altered in quality, and the urine retained in the bladder for ten or twelve hours at the tem- perature of the body, the change may go on much more rapidly, so that the urine may be distinctly alkaline and ammoniacal at the time of its discharge. In these cases, however, it is really acid when first secreted by the kidneys, and becomes alkaline while retained in the interior of the bladder. The first effect of the alkaline condition of the urine, thus pro- duced, is the precipitation of the earthy phosphates. These salts, being insoluble in neutral and alkaline fluids, begin to precipitate as soon as the natural acid reaction of the urine has fairly disappeared, and thus produce in the fluid a whitish turbidity. This precipitate slowly settles upon the sides and bottom of the vessel, or is partly entangled with certain animal matters which rise to the surface and form a thin, opaline scum upon the urine. There are no crystals to be seen at this time, but the deposit is entirely amorphous and granular in character. The next change consists in the production of two new double salts by the action of carbonate of ammonia on the phosphates of soda and magnesia. One of these is the " triple phosphate," phos- phate of magnesia and ammonia (2MgO,NHp,P05-f 2H0). The other is the phosphate of soda and ammonia (NaO,NIip,IIO,POj-f 8H0). The phosphate of magnesia and ammonia is formed from the phosphate of magnesia in the urine (SMgOjPO^+THO) by the replacement of one equivalent of magnesia by one of ammonia. The crystals of this salt are very elegant and characteristic. They 304 EXCRETION. Fig. 117. Phosphate of Magnesia and Ammonia; deposited from healthy urine, during alkaline fermen- tation. show themselves throughout all parts of the mixture ; growing gra- dually in the mucus at the bottom, adhering to the sides of the glass, and scattered abund- antly over the film which col- lects, as we have mentioned, upon the surface. By their refractive power, they give to this film a peculiar glisten- ing or iridescent appearance, which is nearly always visi- ble at the end of six or seven days. The crystals are per- fectly colorless and transpa- rent, and have the form of triangular prisms, generally with bevelled extremities. (Fig. 117.) Frequentlj'-, also, their edges and angles are replaced by secondary facets. They are insoluble in alkalies, but are easily dissolved by acids, even in a very dilute form. At first they are of minute size, but gradually increase, so that after seven or eight days they may become visible to the naked eye. The phosphate of soda and ammonia is formed, in a similar man- ner to the above, by the union of ammonia with the phosphate of soda previously existing in the urine. Its crystals resemble very much those just described, except that their prisms are of a quad- rangular form, or some figure derived from it. They are inter- mingled with the preceding in the putrefying urine, and are affected in the same way by chemical reagents. As the putrefaction of the urine continues, the carbonate of am- monia which is produced, after saturating all the other ingredients with which it is capable of entering into combination, begins to be given off in a free form. The urine then acquires a strong ammoniacal odor; and a piece of moistened test paper, held a little above its surface, will have its color immediately turned by the alkaline gas escaping from the fluid. This is the source of the ammoniacal vapor which is so freely given off from stables and from dung heaps, or wherever urine is allowed to remain and decompose. This process continues until all the urea has been destroyed, and until the products of its decomposition have either united with other substances, or have finally escaped in a gaseous form. SECTION 11. NERVOUS SYSTEM. CHAPTER I. GENERAL STRUCTURE AND FUNCTIONS OF THE NERVOUS SYSTEM Iisr entering upon the study of the nervous system, we commence, the examination of an entirely different order of phenomena from those which have thus far engaged our attention. Hitherto we have studied the physical and chemical actions taking place in the body and constituting together the process of nutrition. We have seen how the lungs absorb and exhale different gases; how the stomach dissolves the food introduced into it, and how the tissues produce and destroy different substances by virtue of the varied transformations which take place in their interior. In all these instances, we have found each organ and each tissue possessing certain properties and performing certain functions, of a physical or chemical nature, which belong exclusively to it, and are charac- teristic of its action. The functions of the nervous system, however, are neither phy- sical nor chemical in their nature. They do not correspond, in their mode of operation, with any known phenomena belonging to these two orders. The nervous system, on the contrary, acts only upon other organs, in some unexplained manner, so as to excite or modify the functions peculiar to them. It is not therefore an appa- ratus which acts for itself, but is intended entirely for the purpose of influencing, in an indirect manner, the action of other organs. Its object is to connect and associate the functions of different parts of the body, and to cause them to act in harmony with each other. 20 306 GENERAL STRUCTURE AND FUNCTIONS This object may be more fully exemplified as follows: — Each organ and tissue in the body has certain properties peculiar to it, which may be called into activity by the operation of a stimu- lus or exciting cause. This capacity, which all the organs possess, of reacting under the influence of a stimulus, is called their excita- bility, or irntability. "We have often had occasion to notice this pro- perty of irritability, in experiments related in the foregoing pages. We have seen, for example, that if the heart of a frog, after being removed from the body, be touched with the point of a needle, it immediately contracts, and repeats the movement of an ordinary pulsation. If the leg of a frog be separated from the thigh, its integument removed, and the poles of a galvanic battery brought in contact with the exposed surface of the muscles, a violent con- traction takes place every time the electric circuit is completed. In this instance, the stimulus to the muscles is supplied by the electric discharge, as, in the case of the heart above mentioned, it is supplied by the contact of the steel needle ; and in both, a muscu- lar contraction is the immediate consequence. If we introduce a metallic catheter into the empty stomach of a dog through a gastric fistula, and gently irritate with it the mucous membrane, a secretion of gastric juice at once begins to take place ; and if food be intro- duced the fluid is poured out in still greater abundance. We know also that if the integument be exposed to contact with a heated body, or to friction with an irritating liquid, an excitement of the circulation is at once produced, which again passes away after the removal of the irritating cause. In all these instances we find that the organ which is called into activity is excited by the direct application of some stimulus to its own tissues. But this is not usually the manner in which the dif- ferent functions are excited during life. The stimulus which calls into action the organs of the living body is usually not direct, but indirect in its operation. Generally speaking, the organs which are situated in distant parts are connected with each other by such a sympathy, that the activity of one is influenced by the condition of the others. The muscles, for example, are almost never called into action by an external stimulus operating directly upon their own fibres, but by one which is applied to some other organ, either adja- cent or remote. Thus the peristaltic action of the muscular coat of the intestine commences when the food is brought in contact with its mucous membrane. The lachrymal gland is excited to an in- creased activity by anything which causes irritation of the conjunc- OF THE NERVOUS SYSTEM. 307 tiva. In all such instances, the physiological connection between two different organs is established through the medium of the nervous system. The function of the nervous system may therefore be defined, in the simplest terms, as follows : It is intended to associate the different parts of the body in such a manner^ that an action may be excited in one organ by means of a stimulus applied to another. The instances of this mode of action are exceedingly numerous. Thus, the light which falls upon the retina produces a contraction of the pupil. The presence of food in the stomach causes the gall- bladder to discharge its contents into the duodenum. The expul- sive efforts of coughing are excited by a foreign body entangled in the glottis. It is easy to understand the great importance of this function, particularly in the higher animals and in man, whose organization is an exceedingly complicated one. For the different organs of the body, in order to preserve the integrity of the whole frame, must not only act and perform their functions, but they must act in har- mony with each other, and at the right time, and in the right direc- tion. The functions of circulation, of respiration, and of digestion, are so mutually dependent, that if their actions do not take place harmoniously, and in proper order, a serious disturbance must inevitably follow. When the muscular system is excited by unu- sual exertion, the circulation is also quickened. The blood arrives more rapidly at the heart, and is sent in greater quantity to the lungs. If the movements of respiration were not accelerated, at the same time, through the connections of the nervous system, there would immediately follow deficiency of aeration, pulmonary conges- tion, and accumulation of blood on the right side of the heart. If the iris were not stimulated to contract by the influence of the light falling on the retina, the delicate expansion of the optic nerve would be dazzled by any unusual brilliancy, and vision would be obscured or confused. In all the higher animals, therefore, where the different functions of the body are performed by distinct organs, situated in different parts of the frame, it is necessary that their action should be thus regulated and harmonized by the operation of the nervous system. The manner in which this is accomplished is as follows: — The nervous system, however simple or however complicated it may be, consists always of two different kinds of tissue, which are 308 GENERAL STRUCTURE AND FUNCTIONS distinguished from each other by their color, their structure, and their mode of action. One of these is known as the white substance, or i\iQ fibrous tissue. It constitutes the whole of the substance of the nervous trunks and branches, and is found in large quantity on the exterior ot the spinal cord, and in the central parts of the brain and cerebellum. In the latter situations, it is of a soft consistency, like curdled cream, and of a uniform, opaque white color. In the trunks and branches of the nerves it has the same opaque white color, but is at the same time of a firmer consistency, owing to its being mingled with condensed areolar tissue. Examined by the microscope, the white substance is seen to be composed every- where of minute fibres or filaments, the "ultimate nervous fila- ments," running in a direction very nearly parallel with each other. These filaments are cylindrical in shape, and vary considerably in size. Those which are met with in the spinal cord and the brain are the smallest, and have an average diameter of yo^oo of an inch. In the trunks and branches of the nerves they average ^^^-^ of an inch. The structure of the ultimate nervous filament is as follows: The exterior of each filament consists of a colorless, transparent tubular membrane, which is seen with some difficulty in the natural condition of the fibre, owing to the extreme delicacy of its texture, and to its cavity being completely filled with a substance yerj similar to it in refractive power. In the interior of this tubular membrane there is contained a thick, softish, semi-fluid nervous matter, which is white and glistening by reflected light, and is called the "white substance of Schwann." Finally, running longi- tudinally through the central part of each filament, is a narrow ribbon-shaped cord, of rather firm consistency, and of a semi- transparent grayish color. This central portion is called the "axis cylinder," or the " flattened band." It is enveloped everywhere by the semi-fluid white substance, and the whole invested by the ex- ternal tubular membrane. When nervous matter is prepared for the microscope and exa- mined by transmitted light, two remarkable appearances are observed in its filaments, produced by the contact of foreign sub- stances. In the first place the unequal pressure, to which the fila- ments are accidentally subjected in the process of dissection and preparation, produces an irregularly bulging or varicose appearance in them at various points, owing to the readiness with which the OF THE NERVOUS SYSTEM. 309 Nervous Filaments from white substance of brain. — a, a, a. Soft substance of the filaments pressed out, and floating in irregularly rounded drops. semi-fluid white substance in their interior is displaced in different directions. (Fig. 118.) Sometimes spots may be seen here and there, where the nervous matter has been entirely pressed apart in the centre of a filament, so that there appears to be an entire break in its continuity, while the investing membrane may be still seen, passing across from one portion to the other. When a nervous filament is torn across under the micro- scope and subjected to pres sure, a certain quantity of the semi-fluid white substance is pressed out from its torn extremity, and may be en- tirely separated from it, so as to present itself under the form of irregularly rounded drops of various sizes (a, a, a), scat- tered over the field of the microscope. The varicose appearance above alluded to is more frequently seen in the smaller nervous filaments from the brain and spinal cord, owing to their soft con- sistency and the readiness with which they yield to pressure. The second effect produced by the artificial preparation of the nervous matter is a partial coagulation of the white substance of Schv/ann. In its natural condition this substance has the same consistency throughout, and appears perfectly transparent and homogeneous by transmitted light. As soon, however, as the nerv- ous filament is removed from its natural situation, and brought in contact with air, water, or other unnatural fluids, the soft substance immediately under the investing membrane begins to coagulate. It increases in consistency, and at the same time becomes more highly refractive ; so that it presents on each side, immediately underneath the investing membrane, a thin layer of a peculiar glistening aspect. (Fig. 119.) At first, this change takes place only in the outer portions of the white substance of Schwann. The coagulating process, however, subsequently goes on, and gradually advances from the edges of the filament toward its 310 GENERAL STRUCTURE AND FUNCTIONS centre, until its entire thickness after a time presents the same ap- pearance. The effect of the same process can also be seen in those portions of the white sub- stance which have been pressed out from the interior of the filaments, and which float about in the form of drops. (Fig. 118, a.) These drops are always covered with a layer of coagulated material which is thicker and more opaque in propor- tion to the length of time which has elapsed since the commencement of the alter- ation. The nervous filaments have essentially the same structure in the brain and spinal cord as in the nervous trunks and branches; only they are of much smaller size in the former than in the latter situation. In the nervous trunks and branches, however, outside the cranial and spinal cavities, there exists, superadded to the nervous filaments and interwoven with them, a large amount of ordinary areolar or fibrous tissue, which protects them from injury, and gives to this portion of the nervous system a peculiar density and resistance. This difference in consistency between the white sub- stance of the nerves and that of the brain and spinal cord is owing, therefore, exclusively to the presence of ordinary fibrous tissue in the nerves, while it is wanting in the brain and spinal cord. The consistency of the nervous filaments themselves is the same in each situation. The nervous filaments are arranged, in the nervous trunks and branches, in a direction nearly parallel with each other. A certain number of them are collected in the form of a bundle, which is invested with a layer of fibrous tissue, in which run the small bloodvessels, destined for the nutrition of the nerve. These pri- mary bundles are united again into secondary, the secondary into tertiary, &;c. A nerve, therefore, consists of a large bundle of ulti- Nhrvohs Filaments from sciatic nerve, showing their coagulation. — At a, the torn extremity of a nervous filament with the axis cylindA' (6) protruding from it. At c, the white substance of Schwann is nearly separated by accidental compression, but the axis- cyliuder passes across the ruptured portion. The out- line of the tubular membrane is also seen at c on the outside of the nervous filament. OF THE NERVOUS SYSTEM. 311 mate filaments, associated with each other in larger or smaller packets, and bound together by the investing fibrous layers. When a nerve is said to become branch- ed or " divided" in any part of its course, this division merely implies that some of its filaments leave the bundles with which they were previously associated, and pursue a different direction. (Fig. 120.) A nerve which ori- ginates, for example, from the spinal cord in the region- of the neck, and runs down the upper extremity, dividing and subdivid- ing, to be finally distributed to the integument and muscles of the hand, contains at its point of (jrigin all the filaments into which it is afterward divided, and which are merely separated at succes- sive points from the main bundle. The ultimate filaments, accord- ingly, are continuous throughout, and do not themselves divide at any point between their origin and their final distribution. When a nerve, furthermore, is said to "inosculate" with another nerve, as when the infra-orbital inosculates with the facial, or the cervical nerves inosculate with each other, this means simply that some of the filaments composing the first nervous bundle separate from it, and cross over to form a part of the second, while some of those belonging to the second cross over and join the first (Fig. 121); but the individual filaments in each instance remain continuous and preserve their identity throughout. This fact is of great physiological importance ; since the white or fibrous nerve-substance is everywhere simply an organ of transmission. It serves to convey the nervous impulse in various directions, from without inward, or from within outward ; and as each nervous filament acts independently of the others, it will convey an impression or a stimulus continuously from its origin to its termination, and will always have the same character and function in every part of its course. Division of a Nerve, showiug portioa of nervous trunk {a), and the separation of its filaments (6, c, d, e). 312 GENERAL STRUCTURE AND FUNCTIONS The other variety of nervous matter is known as the gray sub- stance. It is sometimes called " cineritious matter," and sometimes Fig. 121. Inosculatiou of Nerves. " vesicular neurine," It is found in the central parts of the spinal cord, at the base of the brain in isolated masses, and is also spread out as a continuous layer on the external portions of the cerebrum and cerebellum. It also constitutes the substance of all the gan- glia of the great syrapathe- Fig. 122. Nerve Cells, intermingled witli fibres; from teiiiilunar ganglion of cat. tic. Examined by the micro- scope, it consists of vesicles or cells, of various forms and sizes, imbedded in a grayish, granular, intercellular sub- stance, and containing also, very frequently, granules of grayish pigmentary matter. It is to the presence of this granular pigment that this kind of nervous matter owes the ashy or "cineritious" color from which it derives its name. The cells composing it vary in size, according to OF THE NERVOUS SYSTEM. 313 Kolliker, from -^-q^^ to -^^jj of an inch. The largest of them have a very distinct nucleus and nucleolus. (Fig. 122.) Many of them are provided with long processes or projections, which are some- times divided into two or three smaller branches. These cells are intermingled, in all the collections of gray matter, with nervous filaments, and are entangled with their extremities in such a man- ner that it is exceedingly difficult to ascertain the exact nature of the anatomical relations existing between them. It is certain that in some instances the slender processes running out from the nerv- ous vesicles become at last continuous with the filaments ; but it is not known whether this be the case in all or even in a majority of instances. The extremities of the filaments, however, are at all events brought into very close relation with the vesicles or cells of the gray matter. Every collection of gray matter, whatever be its situation or relative size in the nervous system, is called a ganglion or nervous centre. Its function is to receive impressions conveyed to it by the nervous filaments, and to send out by them impulses which are to be transmitted to distant organs. The ganglia, therefore, originate nervous power, so to speak ; while the filaments and the nerves only transmit it. Now we shall find that, in the structure of every nervous system, the ganglia are connected, first with the different organs, by bundles of filaments which are called nerves; and secondly with each other, by other bundles which are termed commissures. The entire system is accordingly made up of ganglia^ nerves^ and commissures. The simplest form of nervous system is probably that found in the five-rayed starfish. This animal belongs to the type known as radiata; that is, animals whose organs radiate from a central point, Pi?- 123. so as to form a circular series of similar parts, each organ being re- peated at different points of the circumference. The starfish (Fig. 123) consists of a central mass, with five arms or limbs radiating from it. In the centre is the mouth, and immediately beneath it the sto- mach or digestive cavity, which sends prolongations into every one of the projecting limbs. There is also contained in each limb a portion n^hvocs system of starp,. 314 GENERAL STRUCTURE AND FUNCTIONS of the glandular and muscular systems, and the whole is covered by a sensitive integument. The nervous system consists of five similar ganglia, situated in the central portion, at the base of the arms. These ganglia are connected with each other by commis- sures, so as to form a nervous collar or chain, surrounding the orifice of the digestive cavity. Each ganglion also sends off nerves, the filaments of which are distributed to the organs contained in the corresponding limb. We have already stated that the proper function of the nervous system is to enable a stimulus, acting upon one organ, to produce motion or excitement in another. This is accomplished, in the starfish, in the following manner: — When any stimulus or irritation is applied to the integument of one of the arms, it is transmitted by the nerves of the integument to the ganglion situated near the mouth. Arrived here, it is received by the gray matter of the ganglion, and immediately con- verted into an impulse which is sent out by other filaments to the muscles of the corresponding limb; and a muscular contraction and movement consequently take place. The muscles therefore contract in consequence of an irritation which has been applied to the skin. This is called the "reflex action" of the nervous system; because the stimulus is first sent inward by the nerves of the integument, and then returned or reflected back from the ganglion upon the muscles. It must be recollected that this action does not necessarily indicate any sensation or volition, nor even any consciousness on the part of the animal. The function of the gray matter is simply to receive the impulse conveyed to it, and to reflect or send back another ; and this may be accomplished altogether involuntarily, and without the existence of any conscious perception. Where the irritation applied to the integument is of an ordinary character and not very intense, it is simply reflected, as above described, from the corresponding ganglion back to the same limb. But if it be of a peculiar character, or of greater intensity than usual, it may be also transmitted by the commissures to the neighboring ganglia; and so two, three, four, or even all five of the limbs may be set in motion by a stimulus applied to the integument of one of them. Now, as all the limbs of the animal have the same structure and contain the same organs, their action will also be the same ; and the effect of this communication of the stimulus from one to the other by means of commissures will be a repetition, or rather a simultaneous production of similar movements in different parts OF THE NERVOUS SYSTEM. 315 of the body. According to the character and intensity, therefore, of the original stimulus, it will be followed by a response from one, several, or all of the different parts of the animal frame. It will be seen also that there are two kinds of nervous filaments, differing essentially in their functions. One set of these fibres run from the sensitive surfaces to the ganglion, and convey the nervous impression inward. These are called sensitive fibres. The other set run from the ganglion to the muscles, and carry the nervous im- pression outward. These are called motor fibres. In the starfish, wbere the body is composed of a repetition of simi- lar parts arranged round a common centre, and where all the limbs are precisely alike in structure, the several ganglia composing the nervous system are also similar to each other, and act in the same way. But in animals which are constructed upon a different plan, and whose bodies are composed of distinct organs, situated in dif- ferent regions, we find that the nervous ganglia, presiding over the function of these organs, present a corresponding degree of dissimilarity. In Ajylysia, for example, which belongs to the type of mollusca, or soft-bodied animals, the digestive apparatus consists of a mouth, an oesophagus, a triple stomach, and a somewhat convoluted intes- tine. The liver is large, and placed on one side of the body, while the gills, in the form of vascular laminae, occupy the opposite side. There are both testicles and ovaries in the same animal, the male and female functions co-existing, as in many other invertebrate species. All the organs, furthermore, are here arranged without any reference to a regular or symmetrical plan. The body is covered with a muscular man- tle, which expands at the ventral surface into a tolerably well developed " foot," or organ of locomotion, by whicb the animal is enabled to change its position and move from one locality to another. The nervous system of this animal is con- structed upon a plan corresponding with that of the entire body. (Fig. 124.) There is a small ganglion (i) situated anteriorly, which sends nerves to the commencement nervol-s svstem of of the digestive apparatus, and is regarded i^pL^itua^. "rrL;. as the oesophageal or digestive ganglion, i^r^^i gangUon. 3.3. Pedai or T J'il l_l-T-^- 1 /x locomotory ganglia -t. Respi- Immeaiately behmd it is a larger one (i) latory ganguou. Fig. 124. 316 GENERAL STRUCTURE AND FUNCTIONS Fig. 125. called the cephalic or cerebral ganglion, which sends nerves to the organs of special sense, and which is regarded as the seat of volition and general sensation for the entire body. Following this is a pair of ganglia (3, s), the pedal or locomotory ganglia, which supply the muscular mantle and its foot-like expansion, and which regulate the movement of these organs. Finally, another ganglion (4), situated at the posterior part of the body, sends nerves to the branchiae or gills, and is termed the branchial or respiratory ganglion. All these nervous centres are connected by commissures with the central or cerebral ganglion, and may therefore act either independently or in association with each other, by means of these connecting fibres. In the third type of animals, again, viz., the articulata, the gene- ral plan of structure of the body is different from the foregoing, and the nervous system is accordingly modified to correspond with it. In these animals, the body is composed of a number of rings or sections, which are articulated with each other in linear series. A very good example of this type may be found in the common centipede, or scolopendra. Here the body is com- posed of twenty-two successive and nearly simi- lar articulations, each of which has a pair of legs attached, and contains a portion of the glandular, respiratory, digestive and reproductive appara- tuses. The animal, therefore, consists of a repe- tition of similar compound parts, arranged in a longitudinal chain or series. The only exceptions to this similarity are in the first and last articula- tions. The first is large, and contains the mouth; the last is small, and contains the anus. The first articulation, which is called the "head," is also furnished with eyes, with antennae, and with a pair of jaws, or mandibles. The nervous system of the centipede (Fig. 125), corresponding in structure with the above plan, consists of a linear series of nearly equal and similar ganglia arranged in pairs, situated upon the median line, along the ventral surface of the alimentary canal. Each pair of ganglia is con- nected with the integument and muscles of its own articulation by sensitive and motor filaments ; and with those which precede and follow by a double cord of lonojitudinal commissural fibres. In the first articu- NERVons System OF Centipede. OF THE NERVOUS SYSTEM. 317 lation, moreover, or the head, the ganglia are larger than elsewhere, and send nerves to the antennae and to the organs of special sense. This pair is termed the cerebral ganglion, or the " brain." A reflex action may take place, in these animals, through either one or all of the ganglia composing the nervous chain. An impression received bj the integument of any part of the body may be transmitted inward to its own ganglion and thence reflected immediately outward, so as to produce a movement of the limbs belonging to that articulation alone; or it may be propagated, through the longitudinal commissures, forward or back, and pro- duce simultaneous movements in several neighboring articulations; or, finally, it may be propagated quite up to the anterior pair of ganglia, or "brain," where its reception will be accompanied with consciousness, and a voluntary movement reflected back upon any or all of the limbs at once. The organs of special sense, also, com- municate directly with the cerebral ganglia; and impressions con- veyed through them may accordingly give rise to movements in any distant part of the body. In these animals the ventral ganglia, or those which simply stand as a medium of communication be- tween the integument and the muscles, are nearly similar through- out; while the first pair, or those which receive the nerves of special sense, and which exercise a general controlling power over the rest of the nervous system, are distinguished from the remainder by a well-marked preponderance in size. In the centipede it will be noticed that nearly all the organs and functions are distributed in an equal degree throughout the whole length of the body. The organs of special sense alone, with those of mastication and the functions of perception and volition, are confined to the head. The ganglia occupying this part are there- fore the only ones which are distinguished by any external pecu- liarities; the remainder being nearly uniform both in size and activity. In some kinds of articulated animals, however, particular functions are concentrated, to a greater or less extent, in particular parts of the body ; and the nervous ganglia which preside over them are modified in a corresponding manner. In the insects, for example, the body is divided into three distinct sections, viz : the head, containing the organs of prehension, mastication, tact and special sense ; the chest, upon which are concentrated the or- gans of locomotion, the legs and wings; and the abdomen, contain- ing the greater part of the alimentary canal, together with the glandular and generative organs. As the insects have a greater 318 GENERAL STRUCTURE AND FUNCTIONS amount of intelligence and activity, than the centipedes and other worm-like articulata, and as the organs of special sense are more perfect in them, the cerebral ganglia are also unusually developed, and are evidently composed of several pairs, connected by commissures so as to form a compound mass. As the organs of locomotion, fur- thermore, instead of being distributed, as in the centipede, through- out the entire length of the animal, are concentrated upon the chest, the locomotory ganglia also preponderate in size in this region of the body ; while the ganglia which preside over the secretory and generative functions are situated together, in the cavity of the ab- domen. All the above parts, however, are connected, in the same manner as previously described, with the anterior or cerebral pair of gan- glia. In all articulate animals, moreover, the general arrangement of the body is symmetrical. The right side is, for the most part, precisely like the left, as well in the internal organs as in the ex- ternal covering and the locomotory appendages. The only marked variation between different parts of the body is in an antero-pos- terior direction ; owing to difierent organs being concentrated, in some cases, in the head, chest, and abdomen. Finally, in the vertebrate type of animals, comprising man, the quadrupeds, birds, reptiles and fish, the external parts of the body, togetlier with the locomotory apparatus and the organs of special sense, are symmetrical, as in the articulata; but the internal organs, especially those concerned in the digestive and secretory functions, are unsymraetrical and irregular, as in the molluscs. The organs of respiration, however, are nearly symmetrical in the vertebrata, for the reason that the respiratory movements, upon which the function of these organs is immediately dependent, are performed by muscles belonging to the general locomotory apparatus. The nervous system of the vertebrata partakes, accordingly, of the structural arrangement of the organs under its control. That por- tion which presides over the locomotory, respiratory, sensitive, and intellectual functions forms a system by itself, called the cerebro- spinal, system. This system is arranged in a manner very similar to that of the articulata. It is composed of two equal and symmetri- cal halves, running along the median line of the body, the different parts of which are connected by transverse and longitudinal com- missures. Its ganglia occupy the cavities of the cranium and the spinal canal, and send out their nerves through openings in the bony walls of these cavities. OF THE NERVOUS SYSTEM. 319 Fig. 126. The other portion of the nervous system of vertebrata is that which presides over the functions of vegetative life. It is called the ganglionic^ or great sympathetic system. Its ganglia are situated anteriorly to the spinal column, in the visceral cavities of the body, and are connected, like the others, by transverse and longitudinal commissures. This part of the nervous system is symmetrical in the neck and thorax, but is unsymmetrical in the abdomen, where it attains its largest size and its most complete development. The vertebrate animals, as a general rule, are very much superior to the other classes, in intelligence and activity, as well as in the variety and complicated character of their motions; while their nutritive or vegetative functions, on the other hand, are not particu- larly well developed. Accordingly we find that in these animals the cerebro-spinal sj'^stem of nerves preponderates very much, in im- portance and extent, over that of the great sympathetic. The quan- tity of nervous matter contained in the brain and spinal cord is, even in the lowest vertebrate animal, very much greater than that con- tained in the system of the great sympathetic; and this preponderance increases, in the higher classes, just in proportion to their superiority in intelligence, sensation, power of mo- tion, and other functions of a purely animal character. The spinal cord is very nearly alike in the different classes of ver- tebrate animals. It is a nearly cylindrical cord, running from one end of the spinal canal to the other, and connected at its anterior ex- tremity with the ganglia of the brain. (Fig. 126.) It is divided, by an anterior and posterior median fissure, into two lateral halves, which still remain connected with each other by a central mass or commis- sure. Its inner portions are occupied by gray matter, which forms a con- tinuous ganglionic chain, running cerebro-spi^al sy.tkm op max. from one extremity of the cord to — l- cerebrum. 2. Cerebellum 3,3, .3. Spinal .i ,■! -r . cord aad nerves. 4, 4. Brachial nerves. the other. Its outer portions are 5, 5. sacrai nerves. 320 GENERAL STRUCTURE AND FUNCTIONS composed of white substance, the filaments of which run for the most part in a longitudinal direction, connecting the different parts of the cord with each other, and the cord itself with the ganglia of the brain. The spinal nerves are given off from the spinal cord at regular intervals, and in symmetrical pairs; one pair to each successive portion of the body. Their filaments are distributed to the integu- ment and muscles of the corresponding regions. In serpents, where locomotion is performed by simple, alternate, lateral movements of the spinal column, the spinal cord and its nerves are of the same size throughout. But in the other vertebrate classes, where there exist special organs of locomotion, such as fore and hind legs, wings, and the like, the spinal cord is increased in size at the points where the nerves of these organs are given off; and the nerves themselves, which supply the limbs, are larger than those originating from other parts of the spinal cord. Thus, in the hu- man subject (Fig. 126), the cervical nerves, which go to the arms, and the sacral nerves, which are distributed to the legs, are larger than the dorsal and lumbar nerves. They form also, by frequent inosculation, two remarkable plexuses, before entering their corre- sponding limbs, viz., the brachial plexus above, and the sacral plexus below. The cord itself, moreover, presents two enlargements at the point of origin of these nerves, viz., the cervical enlargement from which the brachial nerves (4, 4) are given off, and the lum- bar enlargement from which the sacral nerves (5, 5) originate. If the spinal cord be examined in transverse section (Fig. 127), it will be seen that the gray Fig. 127. matter in its central portion forms a double crescentic- shaped mass, with the con- cavity of the crescents turn- ed outward. These crescentic masses of gray matter, occu- pying the two lateral halves of the cord, are united with each other by a transverse band of the same substance, which is called the gray commissure of the cord. Di- rectly in front of this is a transverse band of white substance, connecting in a similar manner Transverse Section of Spinal Cokd. — a,h. Spinal nerves of right and left side, showing their two roots, d. Origin of anterior root. e. Origin of posterior root. c. Ganglion of posterior root. OF THE NERVOUS SYSTEM. 321 the white portions of the two lateral halves. It is called the white commissure of the cord. The spinal nerves originate from the cord on each side by two distinct roots ; one anterior, and one posterior. The anterior root (Fig. 127, d) arises from the surface of the cord near the extremity of the anterior peak of gray matter. The posterior root (e) origi- nates at the point corresponding with the posterior peak of gray matter. Both roots are composed of a considerable number of ultimate nervous filaments, united with each other in parallel bundles. The posterior root is distinguished by the presence of a small ganglion (c) which appears to be incorporated with it, and through which its fibres pass. There is no such ganglion on the anterior root. The two roots unite with each other shortly after leaving the cavity of the spinal canal, and mingle their filaments in a single trunk. It will be seen, on referring to the diagram (Fig. 127), that each lateral half of the spinal cord is divided into two portions, an anterior and a posterior portion. The posterior peak of gray mat- ter comes quite up to the surface of the cord, and it is just at this point (e) that the posterior roots of the nerves have their origin. The whole of the white substance included between this point and the posterior median fissure is called the posterior column of the cord. That which is included between the same point and the anterior median fissure is the arderior column of the cord. The white substance of the cord may then be regarded as consisting for the most part of four longitudinal bundles of nervous filaments, viz., the right and left anterior, and the right and left posterior columns. The posterior median fissure penetrates deeply into the substance of the cord, quite down to the gray matter, so that the posterior columns appear entirely separated from each other in a transverse section ; while the anterior median fissure is more shal- low and stops short of the gray matter, so that the anterior columns are connected with each other by the white commissure above men- tioned. By the encephalon we mean the whole of that portion of the cerebro-spinal system which is contained in the cranial cavity. It is divided into three principal parts, viz., the cerebrum, cerebellum, and medulla oblongata. The anatomy of these parts, though some- what complicated, can be readily understood if it be recollected that they are simply a double series of nervous ganglia, connected luith each other and with the spinal cord by transverse and longitudinal 21 322 GENERAL STRUCTURE AND FUNCTIONS commissures. The number and relative size of these ganglia, in different kinds of animals, depend upon the perfection of the bodily organization in general, and more especially on that of the intelli- gence and the special senses. They are most readily described by commencing with the simpler forms and terminating with the more complex. The brain of the Alligator (Fig. 128) consists of five pair of ganglia, ranged one behind the other in the interior of the cranium. The first of these are two rounded masses (i), lying just above and behind the nasal cavities, which distribute their nerves upon the Schneiderian mucous mem- brane. These are the olfactory ganglia. They are connected with the rest of the brain by two long and slender commis- sures, the "olfactory commissures." The next pair (2) are somewhat larger and of a triangular shape, when viewed from above downward. They are termed the " cerebral ganglia," or the hemispheres. Imm.ediately following them are two quadrangular masses (3) which give ori- gin to the optic nerves, and are called therefore the optic ganglia. They are termed also the "optic tubercles;" and in some of the higher animals, where they present an imperfect division into four nearly equal parts, they are known as the " tubercula quadri- gemina." Behind them, we have a single triangular collection of nervous matter (4), which is called the cerebellum. Finally, the upper portion of the cord, just behind and beneath the cerebellum, is seen to be enlarged and spread out laterally, so as to form a broad oblong mass (^), the medulla oblongata. It is from this latter portion of the brain that the pneumogastric or respiratory nerves originate, and its ganglia are therefore sometimes termed the "pneu- mogastric" or " respiratory" ganglia. It will be seen that the posterior columns of the cord, as they diverge laterally, in order to form the medulla oblongata, leave be- tween them an open space, which is continuous with the posterior median fissure of the cord. This space is known as the " fourth ventricle." It is partially covered in by the backward projection Brain of Allioator. — 1. Ol- factory ganglia. 2. Hemisplieres. 3. Optic tubercles. 4. Cerebellum. 5. Medulla oblongata. OF THE NERVOUS SYSTEM. 323 of the cerebellum, but in the alligator is still somewhat open pos- teriorly, presenting a kind of chasm or gap between the two lateral halves of the medulla oblongata. The chain of ganglia which compose the brain, being arranged in pairs as above described, are separated from each other on the two sides by a longitudinal median fissure, which is continuous with the posterior median fissure of the cord. In the brain of the alligator this fissure appears to be interrupted at the cerebellum ; but in the higher classes, where the lateral portions of the cere- bellum are more highly developed, as in the human subject (Fig. 126), they are also separated from each other posteriorly on the median line, and the longitudinal median fissure is complete througbout. In hirds^ the hemispheres are of much larger size than in rep- tiles, and partially conceal th.e optic ganglia. The cerebellum, also, is very well developed iu this class, and presents on its sur- face a number of transverse foldings or convolutions, by which the quantity of gray matter which it contains is considerably in- creased. The cerebellum bere extends so far backward as almost completely to conceal the medulla oblongata and the fourth ven- tricle. In the quadrupeds^ the hemispheres and cerebellum attain a still greater size in proportion to the remaining parts of tbe brain. There are also two other pairs of ganglia, situated beneath the hemispheres, and between them and thetubercula quadrigemina. These are the corpora striata in front and the optic thalami behind. In Fig. 129 is shown the brain of the rabbit, with the hemispheres laid open and turned aside, so as to show the internal parts in their natural situation. The olfactory ganglia are seen in front (i) con- nected with the remaining parts by the olfactory commissures. The separation of the hemispheres (2,2) shows the corpora striata (3) and the optic thalami (4). Then come the tubercula quadrigemina (5), which are here composed, as Brain of Rabbit, vieived from above. — 1. Olfactory ganglia. 2. Hemispheres, turned aside. 3. Corpora striata. 4. Optic thalami. 5. Tubercula quadrigemina. 6. Cerebellum. 324 GENERAL STRUCTURE AND FUNCTIONS above mentioned, of four rounded masses, nearly similar in size. The cerebellum (e) is considerably enlarged by the development of its lateral portions, and shows an abundance of transverse convolutions. It conceals from view the fourth ventricle and most of the medulla oblongata. In other species of quadrupeds the hemispheres increase in size so as to project entirely over the olfactory ganglia in front, and to cover in the tubercula quadrigemina and the cerebellum behind. The surface of the hemispheres also becomes covered with numer- ous convolutions, which are curvilinear and somewhat irregular in form and direction, instead of being transverse, like those of the cerebellum. In man, the development of the hemispheres reaches its highest point; so that they preponderate altogether in size over the rest of the ganglia constituting the brain. In the human brain, accordingly, when viewed from above downward, there is nothing to be seen but the convex surfaces of the hemispheres ; and even in a posterior view, as seen in Fig. 126, they conceal everything but a portion of the cerebellum. All the remaining parts, how- ever, exist even here, and have the same connections and relative situation as in other instances. They may be best studied in the following order. As the spinal cord, in the human subject, passes upward into the cranial cavity, it enlarges into the medulla oblongata as already described. The medulla oblongata presents on each side three pro- jections, two anterior and one posterior. The middle projections on its anterior surface (Fig. 130, i, i), which are called the anterior pyramds^ are the con- tinuation of the anterior columns of the cord. They pass onward, underneath the transverse fibres of the pons Varolii, run upward to the corpora striata, pass through these bodies, and radiate upward and outward from their exter- nal surface, to terminate in the gray matter of the hemispheres. The projections immedi- ately on the outside of the anterior pyramids, in the medulla oblongata, are the olivary bodies (2, 2). They contain in their interior a thin layer of gray matter folded upon itself, the functions and connections of which are but little understood, and are not, apparently, of very great importance. Fis. 130. Medclla Oblongata OF Human Brain, ante- rior view. — 1, 1. Anterior py- ramids. 2, 2. Olivary bodies. 3, 3. Kestiform bodies. 4. De- cussation of tlie anterior co- lamns. The medulla oblong- ata is seen terminated above by the transverse fibres of the pons Varolii. OF THE NERVOUS SYSTEM. 325 The anterior columns of the cord present, at the lower part of the medulla oblongata, a remarkable interchange or crossing of their fibres (4). The fibres of the left anterior column pass across the median line at this spot, and becoming continuous with the right anterior pyramid, are finally distributed to the right side of the cerebrum; while the fibres of the right anterior column, passing over to the left anterior pyramid, are distributed to the left side of the cerebrum. This interchange or crossing of the nervous fibres is known as the decussation of the anterior columns of the cord. The posterior columns of the cord, as they diverge on eacli side the fourth ventricle, form the posterior and lateral projections of the medulla oblongata (3, 3). They are sometimes called the "resti- form bodies," and are extremely important parts of the brain. They consist in great measure of the longitudinal filaments of the posterior columns, which pass upward and outward, and are distributed partly to the gray matter of the cerebellum. The remainder then pass forward, underneath the tubercula quadri- gemina, into and through the optic thalami; and radiating thence upward and outward, are distributed, like the continuation of the anterior columns, to the gray matter of the cerebrum. The resti- form bodies, however, in passing upward to the cerebellum, are supplied with some fibres from the anterior columns of the cord, which, leaving the lower portion of the anterior pyramids, join the restiform bodies, and are distributed with them to the cerebellum. From this description it will be seen that both the cerebrum and the cerebellum are supplied with filaments from both the anterior and posterior columns of the cord. In the substance of each restiform body, moreover, there is im- bedded a ganglion which gives origin to the pneumogastric nerve, and presides over the functions of respiration. This ganglion is surrounded and covered by the longitudinal fibres passing upward from the cord to the cerebellum, but may be discovered by cutting into the substance of the restiform body, in which it is buried. It is the first important ganglion met with, in dissecting the brain from below upward. While the anterior columns are passing beneath the pons Varolii, they form, together with the continuation of the posterior columns and the transverse fibres of the pons itself, a rounded prominence or tuberosity, which is known by the name of the tuher annulare. In the deeper portions of this protuberance there is situated, among the longitudinal fibres, another collection of gray matter, which. 326 GENEEAL STRUCTURE AND FUNCTIONS though not of large size, has very important functions and connec- tions. This is known as the ganglion of the tuher annulare. Situated almost immediately above these parts we have the cor- pora striata in front, and the optic thalami behind, nearly equal in size, and giving passage, as above described, to the fibres of the anterior and posterior columns. Behind them still, and on a little lower level, are the tubercula quadrigemina, giving origin to the optic nerves. The olfactory ganglia rest upon the cribriform plate of the ethmoid bone, and send the olfactory filaments through the perforations in this plate, to be distributed upon the mucous mem- brane of the upper and middle turbinated bones. The cerebellum covers in the fourth ventricle and the posterior surface of the medulla oblongata ; and finally the cerebrum, which has attained 'the size of the largest ganglion in the cranial cavity, extends so far in all directions, forward, backward, and laterally, as to form a con- voluted arch or vault, completely covering all the remaining parts of the encephalon. The entire brain may therefore be regarded as a connected series of ganglia, the arrangement of which is shown in the accompany- ing diagram. (Fig. 131.) These ^ig-l^l- ganglia occur in the following order, counting from before back- ward: 1st. The olfactory gan- glia. 2d. The cerebrum or hemi- spheres. 3d. The corpora striata. 4th. The optic thalami. 5th. The tubercula quadrigemina. 6th. The cerebellum. 7th. The gan- glion of the tuber annulare. And 8th. The ganglion of the medulla oblongata. Of these ganglia, only the hemispheres and cere- bellum are convoluted, while the remainder are smooth and round- ed or somewhat irregular in shape. The course of the fibres coming from the anterior and posterior columns of the cord is also to be seen in the accompany- ing figure. A portion of the anterior fibres, we have already ob- served, pass upward and backward, with the restiform bodies, to the cerebellum ; while the remainder run forward through the tuber Diagram of Human Brain, in vertical sec- tion ; ijhowi;ig the situation of the different gan- glia, and the course of the fihres. 1 Olfactory g:inglion. 2. Hemisphere. 3. Corpus striatum. 4. ijptic thalamus. .'5. Tubercula quadrigemina. 6. Cerebellum. 7. Ganglion of tuber annulare. 8. Ganglion of medulla oblongata. OF THE NERVOUS SYSTEM. 327 annulare and the corpus striatum, and then radiate to the gray- matter of the cerebrum. The posterior fibres, constituting the res- tiform body, are distributed partly to the cerebellum, and then pass forward, as previously described, underneath the tubercula quadri- gemina to the optic thalami, whence they are also finally distributed to the gray matter of the cerebrum. The cerebrum and cerebellum, each of which is divided into two lateral halves or " lobes," by the great longitudinal fissure, are both provided with transverse commissures, by which a connection is established between their right and left sides. The great trans- verse commissure of the cerebrum is that layer of white substance which is situated at the bottom of the longitudinal fissure, and which is generally known by the name of the " corpus callosum." Tt consists of nervous filaments, which originate from the gray matter of one hemisphere, converge to the centre where they be- come parallel, cross the median line, and are finally distributed to the corresponding parts of the hemisphere upon the opposite side. The transverse commissure of the cerebellum is the pons Varolii. Its fibres converge from the gray matter of the cerebellum on one side, and pass across to the opposite ; encircling the tuber annulare with a band of parallel curved fibres, to which the name of " pons Varolii" has been given from their resemblance to an arched bridge. The cerebro-spinal system, therefore, consists of a series of gan- glia situated in the cranio-spinal cavities, connected with each other by transverse and longitudinal commissures, and sending out nerves to the corresponding parts of the body. The spinal cord supplies the integument and muscles of the neck, trunk, and extremities ; while the ganglia of the brain, beside supplying the corresponding parts of the head, preside also over the organs of special sense, and perform various other functions of a purely nervous character. 328 OF NERVOUS IRRITABILITY CHAPTETl II. OF NERVOUS IRRITABILITY AND ITS MODE OF ACTION. We have already mentioned, in a previous chapter, that every organ in the body is endowed with the property o^ irritability ; that is, the property of reacting in some peculiar manner when subjected to the action of a direct stimulus. Thus the irritability of a gland shows itself by increased secretion, that of the capillary vessels by congestion, that of the muscles by contraction. Now the irritability of the muscles, indicated as above by their contraction, is extremely serviceable as a means of studying and exhibiting nervous pheno- mena. We shall therefore commence this chapter by a study of some of the more important facts relating to muscular irritability. The irritability of the muscles is a 'property inherent in the mvscular fibre itself. The existence of muscular irritability cannot be ex- plained on any known physical or chemical laws, so far as they relate to inorganic substances. It must be regarded simply as a peculiar property, directly dependent on the structure and consti- tution of the muscular fibre; just as the property of emitting light belongs to phosphorus, or that of combining with metals to oxygen. This property may be called into action by various kinds of stimu- lus; as by pinching the muscular fibre, or pricking it with the point of a needle, the application of an acid or alkaline solution, or the discharge of a galvanic battery. All these irritating applications are immediately followed by contraction of the muscular fibre. This contraction will even take place under the microscope, when the fibre is entirely isolated, and removed from contact with any other tissue ; showing that the properties of contraction and irrita- bility reside in the fibre itself, and are not communicated to it by other parts. Muscular irritability continues for a certain time after death. The stoppage of respiration and circulation does not at once destroy the vital properties of the tissues, but nearly all of them retain these properties to a certain extent for some time afterward. It is only when the constitution of the tissues has become altered by AND ITS MODE OF ACTION. 329 being deprived of blood, and by the consequent derangement of the nutritive process, that their characteristic properties are finally lost. Thus, in the muscles, irritability and contractility may be easily shown to exist for a short time after death by applying to the exposed muscular fibre the same kind of stimulus that we have already found to affect it during life. It is easy to see, in the muscles of the ox, after the animal has been killed, flayed, and eviscerated, different bundles of muscular fiibres contracting irregu- larly for a long time, where they are exposed to the contact of the air. Even in the human subject the same phenomenon may be seen in cases of amputation; the exposed muscles of the amputated limb frequently twitching and quivering for many minutes after their separation from the body. The duration of muscular irritability, after death, varies consi- derably in different classes of animals. It disappears most rapidly in those whose circulation and respiration are naturally the most active; while it continues for a longer time in those whose circula- tion and respiration are sluggish. Thus the muscular irritability in birds continues only a few minutes after the death of the ani- mal. That of quadrupeds lasts somewhat longer ; while in reptiles it remains, under favorable circumstances, for many hours. The cause of this difference is probably that in birds and quadrupeds, the tissues being very vascular, and the molecular changes of nu- trition going on with rapidity, the constitution of the muscular fibre becomes so rapidl}'^ altered after the circula- tion has ceased, that its irritability soon disap- pears. In reptiles, on the other hand, the tissues are less vascular than in birds and quadrupeds, and all the nutritive changes go on more slowly. Eespiration and circulation can therefore be dis- pensed with for a longer period, before the consti- tution of the tissues becomes so much altered as to destroy altogether their vital properties. Owing to this peculiarity of the cold-blooded animals, their tissues may be used with great ad- vantage for purposes of experiment. If a frog's leg, for example, be separated from the body of the animal (Fig. 132), the skin removed, and the poles of a galvanic apparatus applied to the sur- frog's leo, face of the muscle (a, b), a contraction takes place "^''"^ po'<^^ °^ s'^'- 1 . . . 1 1 1 T ^ vanic battery applied every time the circuit is completed and a discharge to the muscles at a, 6. Fig. 132. 330 OF NERVOUS IRRITABILITY passed through the tissues of the limb. The leg of the frog, pre- pared in this way, may be employed for a long time for the pur- pose of exhibiting the effect of various kinds of stimulus upon the muscles. All the mechanical and chemical irritants which we have mentioned, pricking, pinching, cauterizing, galvanism, &c., act with more or less energy and promptitude, though the most efficient of all is the electric discharge. Continued irritation exhausts the irritability of the muscles. It is found that the irritability of the muscles wears out after death more rapidly if they be artificially excited, than if they be allowed to remain at rest. During life, the only habitual excitement of mus- cular contraction is the peculiar stimulus conveyed by the nerves. After death this stimulus may be replaced or imitated, to a certain extent, by other irritants ; but their application gradually exhausts the contractility of the muscle and hastens its final disappearance. Under ordinary circumstances, the post-mortem irritability of the muscle remains until the commencement of cadaveric rigidity. When this has become fairly established, the muscles will no longer contract under the application of an artificial stimulus. Certain poisonous substances have the power of destroying the irritability of the muscles by a direct action upon their tissue. Sulpho-cyanide of potassium, for example, introduced into the cir- culation in sufficient quantity to cause death, destroys entirely the muscular irritability, so that no contraction can afterward be pro- duced by the application of an external stimulant. Nervous Irritability. — The irritability of the nerves is the pro- perty by which they may be excited by an external stimulus, so as to be called into activity and excite in their turn other organs to which their filaments are distributed. When a nerve is irritated, therefore, its power of reaction, or its irritability, can only be esti- mated by the degree of excitement produced in the organ to which the nerve is distributed. A nerve running from the integument to the brain produces, when irritated, a painful sensation ; one distributed to a glandular organ produces increased secretion ; one distributed to a muscle produces contraction. Of all these effects, muscular contraction is found to be the best test and measure of nervous irritability, for purposes of experiment. Sensation cannot of course be relied on for this purpose, since both consciousness and volition are abolished at the time of death. The activity of the glandular organs, owing to the stoppage of the circulation, disappears also very rapidly, or at least cannot readily be demonstrated. The AND ITS MODE OF ACTION. 331 Fig. 133. M contractility of the muscles, however, lasts, as we have seen, for a considerable time after death, and may accordingly be employed with great readiness as a test of nervous irritability. The manner of its employment is as follows: — The leg of a frog is separated from the body and stripped of its integument; the sciatic nerve having been previously dissected out and cut off at its point of emergence from the spinal canal, so that a considerable portion of it remains in connection with the separated limb. (Fig, 133.) If the two poles of a galvanic appa- ratus be now placed in contact with different points (a b) of the exposed nerve, and a discharge allowed to pass between them, at the moment of discharge a sudden contraction takes place in the muscles below. It will be seen that this ex- periment is altogether different from the one re- presented in Fig. 182. In that experiment the galvanic discharge is passed througli the muscles themselves, and acts upon them by direct stim- ulus. Here, however, the discharge passes only from a to & through the tissues of the nerve, and acts directly upon the nerve alone ; while the nerve, acting upon the muscles by its own pecu- liar agency, causes in this way a muscular con- traction. It is evident that in order to produce this effect, two conditions are equally essential: 1st. The irritability of the muscles ; and 2d. The irri- tability of the nerves. So long, therefore, as the muscles are in a healthy condition, their contraction, under the influence of a stimulus applied to the nerve, demonstrates the irritability of the latter, and may be used as a convenient measure of its intensity. The irritdbility of the nerve continues after death. The knowledge of this fact follows from what has just been said with regard to ex- perimenting upon the frog's leg, prepared as above. The irrita- bility of the nerve, like that of the muscle, depends directly upon its anatomical structure and constitution ; and so long as these re- main unimpaired, the nerve will retain its vital properties, though respiration and circulation may have ceased. For the same reason, also, as that given above with regard to the muscles, nervous irri- tability lasts much longer after death in the cold-blooded than in Fror' s Leg, with sciatic nerve (N) at- tached. — a b. Poles of galvanic battery, ap- plied to nerve. 332 OF NERVOUS IRRITABILITY the warm-blooded animals. Various artificial irritants may be em- ployed to call it into activity. Pinching or pricking the exposed nerve with steel instruments, the application of caustic liquids, and the passage of galvanic discharges, all have this effect. The electric current, however, is much the best means to employ for this pur- pose, since it is more delicate in its operation than the others, and will continue to succeed for a longer time. The nerve is, indeed, so exceedingly sensitive to the electric cur- rent, that it will respond to it when insensible to all other kinds of stimulus. A frog's leg freshly prepared with the nerve attached, as in Fig. 133, will react so readily whenever a discharge is passed through the nerve, that it forms an extremely delicate instrument for detecting the presence of electric currents of low intensity, and has even been used for this purpose by Matteucci, under the name of the " galvanoscopic frog." It is only necessary to introduce the nerve as part of the electric circuit; and if even a very feeble cur- rent be present, it is at once betrayed by a muscular contraction. The superiority of electricity over other means of exciting nerv- ous action, such as mechanical violence or chemical agents, pro- bably depends upon the fact that the latter necessarily alter and disintegrate more or -less the substance of the nerve, so that its irritability soon disappears. The electric current, on the other hand, excites the nervous irritability without any marked injury to the substance of the nervous fibre. Its action may, therefore, be continued for a longer period. Nervous irritability^ like that of the muscles, is exhausted by repeated excitement. If a frog's leg be prepared as above, with the sciatic nerve attached, and allowed to remain at rest in a damp and cool place, where its tissue will not become altered by desiccation, the nerve will remain irritable for many hours ; but if it be excited, soon after its separation from the body, by repeated galvanic shocks, it soon begins to react with diminished energy, and becomes gra- dually less and less irritable, until it at last ceases to exhibit any further excitability. If it be now allowed to remain for a time at rest, its irritability will be partially restored ; and muscular con- tractions will again ensue on the application of a stimulus to the nerve. Exhausted a second time, and a second time allowed to re- pose, it will again recover itself; and this maybe even repeated several times in succession. At each repetition, however, the re- covery of nervous irritability is less complete, until it finally dis- appears altogether, and can no longer be recalled. AND ITS MODE OF ACTION. 383 Yarious accidental circumstances tend to diminisTi or destroy nervous irritability. The action of the woorara poison, for example, destroys at once the irritabilit}'- of the nerves; so that in animals killed by this substance, no muscular contraction takes place on irritatinor the nervous trunk. Severe and sudden mechanical in- juries often have the same effect; as where death is produced by violent and extensive crushing or laceration of the body or limbs. Such an injury produces a general disturbance, or shock as it is called, which affects the entire nervous system, and destroys or suspends its irritability. The effects of such a nervous shock may frequently be seen in the human subject after railroad accidents, where the patient, though very extensively injured, may remain for some hours without feeling the pain of his wounds. It is only after reaction has taken place, and the activity of the nerves has been restored, that the patient begins to be sensible of pain. It will often be found, on preparing the frog's leg for experiment as above, that immediately after the limb has been separated from the body and the integument removed, the nerve is destitute of irritability. Its vitality has been suspended by the violence in- flicted in the preparatory operation. In a few moments, however, if kept under favorable conditions, it recovers from the shock, and regains its natural irritability. The action. of the galvanic current upon the nerve, as first shown by the experiments of Matteucci, is in many respects peculiar. If the current be made to traverse the nerve in the natural direction of its fibres, viz., from its origin toward its distribution, as from a to b in Fig. 138, it is called the direct current. If it be made to pass in the contrary direction, as from b to a, it is called the inverse current. When the nerve is fresh and exceedingly irritable, a muscular contraction takes place at both the commencement and termination of the current, whether it be direct or inverse. But very soon afterward, when the activity of the nerve has become somewhat diminished, it will be found that contraction takes place only at the commencement of the direct and at the termination of the inverse current. This may readily be shown by preparing the two legs of the same frog in such a manner that they remain connected with each other by the sciatic nerves and that portion of the spinal column from which these nerves take their origin. The two legs, so prepared, should be placed each in a vessel of water, with the nervous connection hanging between (Fig. 184). If the positive pole, a, of the battery be now placed in the vessel which holds leg 334 OF NERVOUS IRRITABILITY No. 1, and the negative pole, J, in that containing leg No. 2, it will be seen that the galvanic current will traverse the two legs in op- posite directions. In No. 1 it will pass in a direction contrary to the course of its nervous fibres, that is, it will be for this leg an Fig. 134. inverse current; while in No. 2 it will pass in the same direction with that of the nervous fibres, that is, it will be for this leg a direct current. It will now be found that at the moment when the cir- cuit is completed, a contraction takes place in No. 2 by the direct current, while No. 1 remains at rest ; but at the time the circuit is broken, a contraction is produced in No. 1 by the inverse current, but no movement takes place in No. 2. A succession of alternate contractions may thus be produced in the two legs by repeatedly closing and opening the circuit. If the position of the poles, a, i, be reversed, the effects of the current will be changed in a corres- ponding manner. After a nerve has become exhausted by the direct current, it is still sensitive to the inverse; and after exhaustion by the inverse, it is still sensitive to the direct. It has even been found by Mat- teucci that after a nerve has been exhausted for the time by the direct current, the return of its irritability is hastened by the subsequent passage of the inverse current; so that it will become again sensi- tive to the direct current sooner than if allowed to remain at rest. Nothing, accordingly, is so exciting to a nerve as the passage of direct and inverse currents, alternating with each other in rapid succession. Such a mode of applying the electric stimulus is that usually adopted in the galvanic machines used in medical practice, for the treatment of certain paralytic affections. In these machines, AND ITS MODE OF ACTION. 335 tbe electric current is alternately formed and broken witli great rapidity, thus producing the greatest effect upon the nerves with the smallest expenditure of electricity. Such alternating currents, however, if very powerful, exhaust the nervous irritability more rapidly and completely than any other kind of irritation ; and in an animal killed by the action of a battery used in this manner, the nerves may be found to be entirely destitute of irritability from the moment of death. The irrilabiUty of the nerves is distinct from that of the muscles; and the two may be destroyed or suspended independently of each other. When the frog's leg has been prepared and separated from the body, with the sciatic nerve attached, the muscles contract, as we have seen, whenever the nerve is irritated. The irritability of the nerve, therefore, is manifested in this instance only through that of the muscle, and that of the muscle is called into action only through that of the nerve. The two properties may be separated from each other, however, by the action of tcoorara, which has the power, as first pointed out by Bernard, of destroying the irritability of the nerve without affecting that of the muscles. If a frog be poisoned by this substance, and the leg prepared as above, the poles of a galvanic battery applied to the nerve will produce no effect ; show- ing that the nervous irritability has ceased to exist. But if the galvanic discharge be passed directly through the muscles, contrac- tion at once takes place. The muscular irritability has survived that of the nerves, and must therefore be regarded as essentiallv distinct from it. It will be recollected, on the other hand, that in cases of death from the action of sulphocyanide of potassium, the muscular irri- tability is itself destroyed ; so that no contractions occur, even when the galvanic discharge is made to traverse the muscular tissue. There are, therefore, two kinds of paralysis : first, a muscular paralysis, in which the muscular fibres themselves are directly affected ; and second, a nervous paralysis, in which the affection is confined to the nervous filaments, the muscles retaining their natural properties, and being still capable of contracting under the influence of a direct stimulus. Nature of the Nervous Force. — It will readily be seen that the nervous force, or the agency by which the nerve acts upon a muscle and causes its contraction, is entirely a peculiar one, and cannot be regarded as either chemical or mechanical in its nature. The force which is exerted by a nerve in a state of activity is not directly 336 OF NERVOUS IRRITABILITY appreciable in any way by the senses, and can be judged of only by its efifect in causing muscular contraction. This peculiar vitality of the nerve, or, as it is sometimes called, the " nervous force," does not precisely resemble in its operation any of the known physical forces. It has, however, a partial resemblance in some respects to electricity; and this has been sufficient to lead some writers into the error of regarding the two as identical, and of supposing electricity to be really the force acting in the nerves, and operating through them upon the muscles. The principal points of resemblance existing between the two forces, and which have been used in support of the above opinion, are the following : — 1st. The identity of their effects upon the muscular fibre. 2d. The rapidity and peculiarity of their action, by which the force is transmitted almost instantaneously to a distant point, with- out producing any visible effect on the intervening parts. 3d. The extreme sensibility of nerves to the electric current; and 4th. The phenomena of electrical fishes. As these considerations are of some importance in settling the question which now occupies us, we shall examine them in succes- sion. 1st. The Identity of their Effects upon the Muscular Fibre. — It is very true that the muscular fibre contracts under the influence of electricity, as it does under that of the nervous force. This fact, however, does not show the identity of the two forces, but only that they are both capable of producing one particular phenomenon; or that electricity may replace or imitate the nervous force in its action on the muscles. But there are various other agents, as we have already seen, both mechanical and chemical, which will pro- duce the same effect, when applied to the muscular tissue. Elec- tricity, therefore, is only one among several physical forces which resemble each other in this respect, but which are not on that account to be regarded as identical. 2d. The Rapidity and Peculiarity of their Action^ by which the force is transmitted almost instantaneously to a distant pointy without producing any visible effect on the intervening parts, — This is a very remarkable and important character, both of the nervous force and of electricity. In neither case is there any visible effect produced on the nervous or metallic fibre which acts as a conducting medium ; but the final action is exerted upon the substances or organs with which it is in connection. No definite conclusion, however, can be properly derived from the rapidity of their transmission, since AND ITS MODE OF ACTION. 337 this rapidity has never been accurately measured in either instance. We know that light and sound both travel with much greater rapidity than most other physical forces, and that electricity is more rapid in its transmission than either ; but there is no evidence that the velocity of the latter and that of the nervous force are the same. We can only say that in both instances the velocity is very great, without being able to compare them together with any degree of accuracy. The mode of transmission, moreover, alluded to above, is not peculiar to the two forces which are supposed to be identical. Light, for example, is transmitted like them through conducting media, without producing in its passage any sensible effect until it meets with a body capable of reflecting it. In the interval, there- fore, between the luminous body and the reflecting one, there is the same apparent want of action as in the nerve, between the point at which the irritation is applied and its termination in the mus- cular tissue. 3d. The extreme Sensibility of Nerves to the Electric Current. — It has already been mentioned that the electric current is the most delicate of all the means of irritation that may be applied to the nerve after death ; and that it may be used with less deleterious effect than any other. The evident reason for this, however, has already been given. Electricity is one among several physical agents by which the nerve may be artificially excited after death. It is less destructive to the nervous texture than any other, and consequently exhausts its vitality less rapidly. All these agents vary in the delicacy of their operation; and though the electric current happens to be the most efficient of all, it is still simply an artificial irritant, like the rest, capable of imitating, in its own way, the natural stimulus of the nerve. 4th. The Phenomena of Electrical Fishes. — It has been fully demon- strated that certain fish (gymnotus and torpedo) have the power of generating electricity, and of producing electric discharges, which are often sufficiently powerful to kill small animals that may come within their reach. That the force generated by these animals is in reality electricity, is beyond a doubt. It is conducted by the same bodies which serve as conductors for electricity, and is stopped by those which are non-conductors of the same. All the ordinary phenomena produced by the electric current, viz : the heating and melting of a fine conducting wire, the induction of secondary currents and of magnetism, the decomposition of saline solutions, and even the electric spark, have all been produced by the force 22 33S OF NERVOUS IRRITABILITY generated by these animals. There is accordingly, no room for doubt as to its nature. This fact, however, is very far from demonstrating the electric character of the nervous force in general. It is, on the contrary, directly opposed to such a supposition ; since the gymnotus and torpedo are capable of generating electricity simply because they have a special organ destined for this purpose. This organ, which is termed the " electrical organ," is peculiar to these fish, and where it is absent, the power of generating electricity is absent also. The electrical organs of the gymnotus and torpedo occupy a considerable portion of the body, and are largely supplied with nerves which regulate their function. If these nerves be divided, tied, or injured in any way, the electrical organ is weakened or paralyzed, just as the muscles would be if the nerves distributed to them were sub- jected to a similar violence. The electricity produced by these animals is not supplied by the nerves, but by a special generating organ, the action of which is regulated by nervous influence. The reasons quoted above, therefore, are quite insufficient for establishing any relation of identity between the nervous force and electricity. There are, moreover, certain well authenticated facts directly opposed to such a supposition, the most important of which are the following : — The first is, that no electrical current has been actually found to exist in an irritated nerve. The most conclusive experiments on this point are those which were made by Longet and Matteucci, in company with each other, at the veterinary school of Alfort.^ The galvano- meter employed in these investigations was constructed under the personal direction of the experimenters, and was of extreme delicacy. The oscillating needle was surrounded by 2500 turns of conducting wire, and the poles were each armed with a platinum plate, having an exposed surface of one sixth of a square inch. When the poles of the apparatus had been repeatedly immersed in spring water, so that no further variation was produced from this source, the instru- ment was considered as ready for use. The sciatic nerve of a liv- ing horse was then exposed, and the poles of the galvanometer placed in contact with it, in various positions, both diagonally and longitudinally, and at various depths in its interior. The examina- tion was continued for a quarter of an hour, during which time the painful sensations of the animal were testified by constant strug- gling movements of the limbs; showing that both the motor and ' Longet, Traite de Physiologic. Paris, 1850, vol ii. p. 130. AND ITS MODE OF ACTION. 339 sensitive filaments of the nerve were in a high state of activity. The conclusion, however, to which the experimenters were con- ducted was the following, viz: that "there was no constant and re- liable evidence of the existence of an electric current in the nerve." Secondly. The mode of conduction of the nervous force is different from that of electricity/. The latter force, in order to exert its charac- teristic effects, must be transmitted through isolated conductors, so arranged as to form a complete circuit. No such circuit has ever been shown to exist in the nervous system ; and the nerves them- selves, the only tissues capable of conducting the nervous force, are not particularly good conductors of electricity; no better, for exam- ple, than the muscles or the areolar tissue. We know of nothing, therefore, which should prevent an electric current, passing through a nerve, from being dispersed and lost among the adjacent tissues. This is not the case, however, with the natural stimulus conveyed by the nervous filament. Moreover the nerve, in order to conduct its own peculiar force, must be in a state of complete integrity. If a ligature be applied to it, or if it be pinched or lacerated, the muscles to which it is dis- tributed are paralyzed for all voluntary motion, and yet it transmits the electric current as readily as before. If the nerve be divided, and its divided extremities replaced in apposition with each other, it will still act perfectly well as a conductor of electricity, though it is needless to say that its natural function is at once destroyed. The difference in the mode of conduction between the two forces may be shown in a still more striking manner, as follows. Let the nerve connected with a frog's leg be divided, and its two extremi- ties joined to each other by a piece of moist cotton thread. If the galvanic current be now passed through the detached portion of the nerve, no contraction will take place ; because the nervous force, excited in the detached portion, cannot be transmitted through the cotton thread to the remainder. But if one of the galvanic poles be applied above, and the other below the point of division, a con- traction is immediately produced ; since the electric current is readily transmitted by the cotton thread, and excites the lower portion of the nerve, which is still in connection with the muscles. The nervous force, therefore, while it has some points of resem- blance with electricity, presents also certain features of dissimilarity which are equally important. It must be regarded accordingly as distinct in its nature from other known physical forces, and as altogether peculiar to the nervous tissue in which it originates. 840 THE SPINAL CORD. CHAPTER III. THE SPINAL CORD. We have already seen that the spinal cord is a long ganglion, covered with longitudinal bundles of nervous filaments, and occu- pying the cavity of the spinal canal. It sends out nerves which supply the muscles and integument of at least nine-tenths of the whole body, viz., those of the neck, trunk, and extremities. All these parts of the body are endowed with two very remarkable properties, the exercise of which depends, directly or indirectly, upon the integrity and activity of the spinal cord, viz., the power of sensation and the power of motion. Both these properties are said to reside in the nervous system, because they are so readily influenced by its condition, and are so closely connected with its physiological action. "We shall therefore commence the study of the spinal cord with an examination of these two functions, and of the situation which they occupy in the nervous system. / DISTINCT SEAT OF SENSATION AND MOTION IN THE NERVOUS SYSTEM. Sensation and motion are usually the first functions which suffer by any injury inflicted on the nervous system. As a general rule, they are both suspended or impaired at the same time, and in a nearly equal degree. In a fainting fit, an attack of apoplexy, con- cussion or compression of the brain or spinal cord, or a wound of any kind involving the nerves or nervous centres, insensibility and loss of motion usually appear simultaneously. It is difficult, there- fore, under ordinary conditions, to trace out the separate action of these two functions, or to ascertain the precise situation occupied by each. This difficulty, however, may be removed by examining sepa- rately different parts of the nervous system. In the instances mentioned above, the injury which is inflicted is comparatively an DISTINCT SEAT OF SENSATION AND MOTION. 341 extensive one, and involves at the same tinne many adjacent parts. T5ut instances sometimes occur in which the two functions, sensa- tion and motion, are affected independently of each other, owing to the peculiar character and situation of the injury inflicted. Sensa- tion may be impaired without loss of motion, and loss of motion may occur without injury to sensation. In tic douloureux, for example, we have an exceedingly painful affection of the sensitive parts of the face, without any impairment of its power of motion • and in facial paralysis we often see a complete loss of motion affect- ing one side of the face, while the sensibility of the part remains altogether unimpaired. The above facts first gave rise to the belief that sensation and motion might occupy distinct parts of the nervous system ; since it would otherwise be difficult to understand how the two could be affected independently of each other by anatomical lesions. It has accordingly been fully established, by the labors of Sir Charles Bell, Miiller, Panizza, and Longet, that the two functions do in reality occupy distinct parts of the nervous system. If any one of the spinal nerves, in the living animal, after being exposed at any part of its course outside the spinal canal, be divided, ligatured, bruised, or otherwise seriously injured, paralysis of motion and loss of sensation are immediately produced in that part of the body to which the nerve is distributed. If, on the other hand, the same nerve be pricked, galvanized, or otherwise gently irritated, a painful sensation and convulsive movements are produced in the same parts. The nerve is therefore said to be both sensitive and excitable; sensitive, because irritation of its fibres produces a pain- ful sensation, and excitable, because the same irritation causes mus- cular contraction in the parts below. The result of the experiment, however, will be different if it be tried upon the parts situated inside the spinal canal, and particularly upon the anterior and posterior roots of the spinal nerves. If an irritation be applied, for example, to the anterior root of a spinal nerve, in the living animal, convulsive movements are produced in the parts below, but there is no painful sensation. The anterior root accordingly is said to be excitable, but not sensitive. If the posterior root, on the other hand, be irritated, acute pain is pro- duced, but no convulsive movements. The posterior root is there- fore sensitive, but not excitable. A similar result is obtained by a complete division of the two roots. Division of the anterior root produces paralysis of motion, but no insensibility ; division of the 3ri2 THE SPINAL CORD. posterior root produces complete loss of sensibility, but no muscular paralysis. We have here, then, a separate localization of sensation and motion in the nervous system ; and it is accordingly easy to under- stand how one may be impaired without injury to the other, or how both may be simultaneously affected, according to the situation and extent of the anatomical lesion. The two roots of a spinal nerve differ from each other, further- more, in their mode of transmitting the nervous impulse. If the posterior root be divided (E'ig. 135) at «, b, and an irritation applied Fig. 135. Diagi-am of SpiXAL Cord and Nekves. The posterior root is seeu divided at n, h. tlie anterior at c, d. to the separated extremity (a), no effect will be produced ; but if the irritation be applied to the attached extremity {h\ a painful sensation is immediately the result. The nervous force, therefore, travels in the posterior root from without inward, but cannot pass from within outward. If the anterior root, on the other hand, be divided at c, d, and its attached extremity [d) irritated, no effect follows ; but if the separated extremity (c) be irritated, convulsive movements instantly take place. The nervous force, consequently, travels in the anterior root from within outward, but cannot pass from without inward. The same thing is true with regard to the transmission of sensa- tion and motion in the spinal nerves outside the spinal canal. If one of these nerves be divided in the living animal, and its attached extremity irritated, pain is produced, but no convulsive motion; if SENSIBILITY AND EXCITABILITY IN SPINAL CORD. 343 the irritation be applied to its separated extremity, muscular con- tractions follow, but no painful sensation. There are, therefore, two kinds of filaments in the spinal nerves, not distinguishable by the eye, but entirely distinct in their character and function, viz., the "sensitive" filaments, or those which convey sensation, and the " motor" filaments, or those which excite move- ment. These filaments are never confounded with each other in their action, nor can they perform each other's functions. The sen- sitive filaments convey the nervous force only in a centripetal, the motor only in a centrifugal direction, The former preside over sensation, and have nothing to do with motion ; the latter preside over motion, and have nothing to do with sensation. Within the spinal canal the two kinds of filaments are separated from each other, constituting the anterior and posterior roots of each spinal nerve; but externally they are mingled together in a common trunk. While the anterior and posterior roots, therefore, are ex- clusively sensitive or exclusively motor, the spinal nerves beyond the junction of the roots are called mixed nerves^ because they con- tain at the same time motor and sensitive filaments. The mixed nerves accordingly preside at the same time over the functions of movement and of sensation. DISTINCT SEAT OF SENSIBILITY AND EXCITABILITY IN THE SPINAL CORD. Various experimenters have demonstrated the fact that different ])arts of the spinal cord, like the two roots of the spinal nerves, are separately endowed with sensibility and excitability. The anterior columns of the cord, like the anterior roots of the spinal nerves, are excitable but not sensitive ; the posterior columns, like the posterior roots of the spinal nerves, are sensitive but not excitable. Accordingly, when the spinal canal is opened in the living animal, an irritation applied to the anterior columns of the cord produces immediately convulsions in the limbs below ; but there is no indi- cation of pain. On the other hand, signs of acute pain become manifest whenever the irritation is applied to the posterior column ; but no muscular contractions follow, other than those of a voluntary character. Longet has found'that if the spinal cord be exposed in the lumbar region and completely divided at that part by trans- ' Traite de Physiologie, voL ii. part 2, p. 8. 344 THE SPINAL COED. verse section, the application of any irritant to tbe anterior surface of the separated portion produces at once convulsions below ; while if applied to the posterior columns behind the point of division, it has no sensible effect whatever. The anterior and posterior columns of the cord are accordingly, so far, analogous in their properties to the anterior and posterior roots of the spinal nerves, and are plainly composed, to a greater or less extent, of a continuation of their filaments. These filaments, derived from the anterior and posterior roots of the spinal nerves, pass upward through the spinal cord toward the brain. An irritation applied to any part of the integument is then conveyed, along the sensitive filaments of the nerve and its posterior root, to the spinal cord; then upward, along the longitudinal fibres of the cord to the brain, where it produces a sensation corresponding in character with the original irritation. A motor impulse, on the other hand, originating in the brain, is transmitted downward, along the longitudinal fibres of the cord, passes outward by the anterior root of the spinal nerve, and follow- ing the motor filaments of the nerve through its trunk and branches, produces at last a muscular contraction at the point of its final distribution. CEOSSED ACTION OF THE SPINAL COED. As the anterior columns of the cord pass upward to join the medulla oblongata, a decussation takes place between them, as we have already mentioned in Chapter I. The fibres of the right anterior column pass over to the left side of the medulla oblongata, and so upward to the left side of the brain ; while the fibres of the left anterior column pass over to the right side of the medulla oblongata, and so upward to the right side of the brain. This decussation may be readily shown (as in Fig. 130) by gently separating the anterior columns from each other, at the lower ex- tremity of the medulla oblongata, where the decussating bundles may be seen crossing obliquely from side to side, at the bottom of the anterior median fissure. Below this point, the anterior columns remain distinct from each other on each side, and do not communi- cate by any further decussation. If the anterior columns of the spinal cord, therefore, be wounded at any point in the cervical, dorsal, or lumbar region, a paralysis CROSSED ACTION OF THE SPINAL CORD. 345 of voluntary motion is produced in the limbs below, on the same side with the injury. But if a similar lesion occur in the brain, the paralysis which results is on the opposite side of the body. Thus it has long been known that an abscess or an apoplectic hemorrhage on the right side of the brain will produce paralysis of the left side of the body; and injury of the left side of the brain will be followed by paralysis of the right side of the body. The spinal cord has also a crossed action in transmitting sensi- tive as well as motor impulses. It has been recently demonstrated by Dr. Brown-Sdquard,' that the crossing of the sensitive fibres in the spinal cord does not take place, like that of the motor fibres, at its upper portion only, but throughout its entire length ; so that the sensitive fibres of the right spinal nerves, very soon after their entrance into the cord, pass over to the left side, and those of the left spinal nerves pass over to the right side. For if one lateral half of the spinal cord of a dog be divided in the dorsal region, the power of sensation remains upon the corresponding side of the body, but is lost upon the opposite side. It has been shown, fur- thermore, by the same observer,^ that the sensitive fibres of the spinal nerves, when they first enter the cord join the posterior columns, which are everywhere extremely sensitive ; but that they very soon leave the posterior columns, and, passing through the central parts of the cord, run upward to the opposite side of the brain. If the posterior columns, accordingly, be alone divided at any part of the spinal cord, sensibility is not destroyed in all the nerves behind the seat of injury, but only in those which enter the cord at the point of section ; since the posterior columns consist of different nervous filaments, joining them constantly on one side from below, and leaving them on the other to pass upward toward the brain. """"^ The spinal cord has therefore a crossed action, both for sensa- tion and motion ; but the crossing of the motor filaments occurs only at the medulla oblongata, while that of the sensitive filaments takes place throughout the entire length of the cord itself. There are certain important facts which still remain to be noticed, regarding the mode of action of the spinal cord and its nerves. They are as follows : — ' Experimental Researches applied to Physiology and Pathology. New York, 1853. 2 Memoirs sur la Physiologie de la Moelle epiniere ; Gazette Medicale de Paris, 1855. 3.46 THE SPINAL CORD. 1. An {rrilation a2'>plied to a spinal nerve at the middle of its course "produces the same effect as if it traversed its entire length. Thus, if the sciatic or median nerve be irritated at any part of its course, con- traction is produced in the muscles to which these nerves are dis- tributed, just as if the impulse had originated as usual from the brain. This fact depends upon the character of the nervous fila- ments, as simple conductors. Wherever the impulse may originate, the final effect is manifested only at the termination of the nerve. As the impulse in the motor nerves travels always in an outward direction, the effect is always produced at the muscular termination of the filaments, no matter how small or how large a portion of their length may have been engaged in transmitting the stimulus. If the irritation, again, be applied to a sensitive nerve in the middle of its course, the painful sensation is felt, not at the point of the nerve directly irritated, but in that portion of the integu- ment to which its filaments are distributed. Thus, if the ulnar nerve be accidentally struck at the point where it lies behind the inner condyle of the humerus, a sensation of tingling and numb- ness is produced in the last two fingers of the corresponding hand. It is common to hear patients who have suffered amputation com- plain of painful sensations in the amputated limb, for weeks or months, and sometimes even for years after the operation. They assert that they can feel the separated parts as distinctly as if they were still attached to the body. This sensation, which is a real one and not fictitious, is owing to some irritation operating upon the divided extremities of the nerves in the cicatrized wound. Such an irritation, conveyed to the brain by the sensitive fibres, will pro- duce precisely the same sensation as if the amputated parts were still present, and the irritation actually applied to them. It is on this account also that division of the trifacial nerve is not always effectual in the cure of tic douloureux. If the cause of the difficulty be seated upon the trunk of the nerve, between its point of emergence from the bones and its origin in the brain, it is evident that division of the nerve upon the face will be of no avail; since the cause of irritation will still exist behind the point of section, and the same painful sensations will still be produced in the brain. 2. The irritability of the motor filaments disapjyears from within oiit- ward, that of the sensitive filaments from without inward. Immedi- ately after the separation of the frog's leg from the body, irritation of the nerve at any point produces muscular contraction in the INDEPENDENCE OF NEEVOUS FILAMENTS. 347 limb below. As time elapses, however, and the irritability of the nerve diminishes, the galvanic current, in order to produce con- traction, must be applied at a point nearer its termination. Subse- quently, the irritability of the nerve is entirely lost in its upper portions, but is retained in the parts situated lower down, from which it also, in turn, afterward disappears; receding in this man- ner farther and farther toward the terminal distribution of the nerve, where it finally disappears altogether. On the other hand, sensibility disappears, at the time of death, first in the extremities. From them the numbness gradually creeps upward, invading successively the middle and upper portions of the limbs, and the more distant portions of the trunk. The central parts are the last to become insensible. 3. Each nervous filament acts independently of the rest throughout its entire lengthy and does not communicate its irritation to those which are in proximity with it. It is evident that this is true with regard to the nerves of sensation, from the fact that if the integument be touched with the point of a needle, the sensation is referred to that spot alone. Since the nervous filaments coming from it and the adjacent parts are all bound together in parallel bundles, to form the trunk of the nerve, if any irritation were communicated from one sensitive filament to another, the sensation produced would be indefinite and diffused, whereas it is really confined to the spot irri- tated. If a frog's leg, furthermore, be prepared, with the sciatic nerve attached, a few of the fibres separated laterally from the nervous trunk for a portion of its length, and the poles of a galvanic battery applied to the separated portion, the contractions which follow in the leg will not be general, but will be confined to those muscles in which the galvanized nervous fibres especially have their distribution. There are also various instances, in the body, of antagonistic muscles, which must act independently of each other, but which are supplied with nerves from a common trunk. The superior and inferior straight muscles of the eyeball, for example, are both supplied by the motor oculi communis nerve. Extensor and flexor muscles, as, for example, those of the fingers, are often supplied by the same nerve, and yet act alternately with- out mutual interference. It is easy to see that if this were not the case, confusion would constantly arise, both in the perception of sensations, and in the execution of movements, 4. There are certain sensations which are excited simultaneously by the same causes, and which are termed associated sensations ; and 348 THE SPINAL COED. there are also certain movements which take place simultaneously, and are called associated movements. In the former instance, one of the associated sensations is called up immediately upon the percep- tion of the other, without requiring any direct impulse of its own. Thus, tickling the soles of the feet produces a peculiar sensation at the epigastrium. Nausea is occasioned by certain disagreeable odors, or by rapid rotation of the body, so that the landscape seems to turn round. A striking example of associated movements, on the other hand, may be found in the action of the muscles of the eyeball. The eyeballs always accompany each other in their lateral motions, turning to the right or the left side simultaneously. It is evident, however, that in producing this correspondence of motion, the left internal rectus muscle must contract and relax together with the right external; while a similar harmony of action must exist between the right internal and the left external. The explana- tion of such singular correspondences cannot be found in the anato- mical arrangement of the muscles themselves, nor in that of the nervous filaments by which they are directly supplied, but must be looked for in some special endowment of the nervous centres from which they originate. EEFLEX ACTION OF THE SPINAL CORD. The spinal cord, as we have thus far examined it, may be re- garded simply as a great nerve ; that is, as a bundle of motor and sensitive filaments, connecting the muscles and integument below with the brain above, and assisting, in this capacity, in the produc- tion of conscious sensation and voluntary motion. Beside its nerv- ous filaments, however, it contains also a large quantity of gray matter, and is, therefore, itself a ganglionic centre, and capable of independent action as such. We shall now proceed to study it in its second capacity, as a distinct nervous centre. If a frog be decapitated, and the body allowed to remain at rest for a few moments, so as to recover from the depressing effects of shock upon the nervous system, it will be found that, although sen- sation and consciousness are destroyed, the power of motion still remains. If the skin of one of the feet be irritated by pinching it with a pair of forceps, the leg is immediately drawn up toward the body, as if to escape the cause of irritation. If the irritation applied to the foot be of slight intensity, the corresponding leg only will KEFLEX ACTION OF THE SPINAL CORD. 349 move; but if it be more severe in character, motions will often be produced in the posterior extremity of the opposite side, and even in the two fore-legs, at the same time. These motions, it is import- ant to observe, are never spontaneous. The decapitated frog remains perfectly quiescent if left to himself. It is only when some cause of irritation is applied externally, that movements occur as above described. It will be seen that the character of these phenomena indicates the active operation of some part of the nervous system, and par- ticularly of some ganglionic centre. The irritation is applied to the skin of the foot, and the muscles of the leg contract in conse- quence; showing evidently the intermediate action of a nervous connection between the two. The effect in question is due to the activity of the spinal cord, operating as a nervous centre. In order that the movements may take place as above, it is essential that both the integument and the muscles should be in communication with the spinal cord by nerv- ous filaments, and that the cord itself be in a state of integrity. If the sciatic nerve be divided in the upper part of the thigh, irrita- tion of the skin below is no longer followed by any muscular con- traction. If either the anterior or posterior roots of the nerve be divided, the same want of action results; and finally, if, the nerve and its roots remaining entire, the spinal cord itself be broken up by a needle introduced into the spinal canal, the integument may then be irritated or mutilated to any extent, without exciting the least muscular contraction. It is evident, therefore, that the spinal cord acts, in this case, as a nervous centre, through which the irritation applied to the skin is communicated to the muscles. The irritation first passes upward, as shown in the accompanying diagram (Fig. 136), along the sensitive fibres of the posterior root (a) to the gray matter of the cord, and is then reflected back, along the motor fibres of the anterior root, until it finally reaches the mus- cles, and produces a contraction. This action is known, accord ingly, as the reflex action of the spinal cord. Fig. 136. Diagram of Spinal Cord in Ver- tical Section, showing reflex action. — a Posterior root of spinal nerve. 6. An- terior root of spinal nerve. 350 THE SPIXAL CORD. It will be remembered that this reflex action of the cord is not accompanied by any volition, nor even by any conscious sensation. The function of the spinal cord as a nervous centre is simply to convert an impression, received from the skin, into a motor impulse which is sent out again to the muscles. There is absolutely no farther action than this ; no exercise of will, consciousness, or judg- ment. This action will therefore take place perfectly well after the brain has been removed, and after the entire sympathetic sys- tem has also been taken away, provided only that the spinal cord and its nerves remain in a state of integrity. The existence of this reflex action after death is accordingly an evidence of the continued activity of the spinal cord, just as con- tractility is an evidence of the activity of the muscles, and irrita- bility of that of the nerves. Like the two last-mentioned properties, also, it continues for a longer time after death in cold-blooded than in warm-blooded animals. It is for this reason that frogs and other reptiles are the most useful subjects for the study of these pheno- mena, as for that of most others belonging to the nervous system. The irritability of the spinal cord, as manifested by its reflex action, may be very much exaggerated by certain diseases, and by the operation of poisonous substances. Tetanus and poisoning b}^ strychnine both act in this way, by heightening the irritability of the spinal cord, and causing it to produce convulsive movements on the application of external stimulus. It has been observed that the convulsions in tetanus are rarely, if ever, spontaneous, but that they always require to be excited by some external cause, such as the accidental movement of the bedclothes, the shutting of a door, or the sudden passage of a current of air. Such slight causes of irritation, which would be entirely inadequate to excite involuntary movements in the healthy condition, act upon the spinal cord, when its irritability is heightened by disease, in such a manner as to pro- duce violent convulsions. Similar appearances are to be seen in animals poisoned by strych- nine. This substance acts upon the spinal cord and increases its irritability, without materially affecting the functions of the brain. Its effects will show themselves, consequently, without essential modification, after the head has been removed. If a decapitated frocr be poisoned with a moderate dose of strychnine, the body and limbs will remain quiescent so long as there is no external source of excitement; but the limbs are at once thrown into convulsions by the slightest irritation applied to the skin, as, for example, the KEFLEX ACTION OF THE SPINAL CORD. 351 contact of a hair or a feather, or even the jarring of the table on which the animal is placed. That the convulsions in cases of poisoning by strychnine are always of a reflex character, and never spontaneous, is shown by the following fact first noticed by Bernard,' viz., that if a frog be poisoned after division of the posterior roots of all the spinal nerves, while the anterior roots are left untouched, death takes place as usual but is not preceded by any convulsions. In this instance the convulsions are absent simply because, owing to the division of the posterior roots, external irritations cannot be communicated to the cord. The reflex action, above described, may be seen very distinctly in the human subject, in certain cases of disease of the spinal cord. If the upper portion of the cord be disintegrated by inflammatory softening, so that its middle and lower portions lose their natural connection with the brain, paralysis of voluntary motion and loss of sensation ensue in all parts of the body below the seat of the anatomical lesion. Under these conditions, the patient is incapable of making any muscular exertion in the paralyzed parts, and is unconscious of any injury done to the integument in the same region. Notwithstanding this, if the soles of the feet be gently irritated with a feather, or with the point of a needle, a convulsive twitching of the toes will often take place, and even retractile move- ments of the leg and thigh, altogether without the patient's know- ledge. Such movements may frequently be excited by simply allowing the cool air to come suddenly in contact with the lower extremities. We have repeatedly witnessed these phenomena, in a case of disease of the spinal cord where the paralysis and in- sensibility of the lower extremities were complete. Many other similar instances are reported by various authors^ The existence of this reflex action of the cord has enabled the physiologist to ascertain several other important facts concerning the mode of operation of the nervous system. M, Bernard has demonstrated,^ by a series of extremely ingenious experiments on the action of poisonous substances, 1st, that the irritability of the muscles may be destroyed, while that of the nerves remains unal- tered ; and 2d, that the motor and sensitive nervous filaments may each be paralyzed independently of each other. The above facts are shown by the three following experiments: — ' Lecjons sur les eflFets des Substances toxiques et medicamenteuses, Paris, 1857, p. 357. ^ Lecjons sur le.s effets des Substances toxiques et nielicanienteuses, Chaps. 23 and 24. 352 THE SPIXAL COBD. 1. In a living frog (Fig. 137), the sciatic nerve (xV) is exposed in the back part of the thigh, after which a ligature is passed underneath it and drawn tight around the ^^" bone and the remaining soft parts. In this way the circu- lation is entirely cut off" from the limb {d)^ which remains in connection with the trunk only by means of the sciatic nerve. A solution of sulpho- cyanide of potassium is then introduced beneath the skin of the back, at /, in sufficient quantity to produce its speci- fic effect. The poison is then absorbed, and is carried by the circulation throughout the trunk and the three extremi- ties a, 6, c; while it is prevented from entering the limb c?, by the ligature which has been placed about the thigh. Sul- phocyanide of potassium pro- duces paralysis, as we have previously mentioned, by act- ing directly upon the muscu- lar tissue. Accordingly, a gal- vanic discharge passed through the limbs a, 5, and c, produces no contraction in them, while the same stimulus, applied to d, is follow^ed by a strong and healthy reaction. But at the moment when the irritation is applied to the poisoned limbs a, h, and c, though no visible effect is produced in them, an active movement takes place in the healthy limb, d. This can only be owing to a reflex action of the spinal cord, originating in the integument of a, 6, and c, and transmitted, by sensitive and motor filaments, through the cord, to d. While the muscles of the poisoned limbs, therefore, have been directly paralyzed, the nerves of the same parts have retained their irritability. 2. If a frog be poisoned with woorara by simply placing the REFLEX ACTION OF THE SPINAL CORD. 353 poison under the skin, no reflex action of the spinal cord can be demonstrated after death. We have already shown, from experi- ments detailed in Chapter II., that this substance destroys the irrita- bility of the motor nerves, without affecting that of the muscles. In the above instance, therefore, where the reflex action is abolished, its loss may be owing to a paralysis of both motor and sensitive fila- ments, or to that of the motor filaments alone. The following experi- ment, however, shows that the motor filaments are the only ones affected. If a frog be prepared as in Fig. 137, and poisoned by the introduction of woorara at /, when the limb d is irritated its own muscles react, while no movement takes place in a, 5, or c; but if the irritation be applied to a, &, or c, reflex movements are imme- diately produced in d. In the poisoned limbs, there/ore, while the motor nerves have been paralyzed, the sensitive filaments have retained their irritability. 3. If a frog be poisoned with strychnine, introduced underneath the skin in sufficient quantity, death takes place after general con- vulsions, which are due, as we have seen above, to an unnatural excitability of the reflex action. This is followed, however, by a paralysis of sensibility, so that after death no reflex movements can be produced by irritating the skin or even the posterior roots of the spinal nerves. But if the anterior roots, or the motor nerves themselves be galvanized, contractions immediately take place in the corresponding muscles. In this case, therefore, the sensitive fila- ments have been paralyzed, while the motor filaments and the muscles have retained their irritability. We come now to investigate the reflex action of the spinal cord, as it takes place in a healthy condition during life. This action readily escapes notice, unless our attention be particularly directed to it, because the sensations which we are constantly receiving, and the many voluntary movements which are continually executed, serve naturally to mask those nervous phenomena which take place without our immediate knowledge, and over which we exert no voluntary control. Such phenomena, however, do constantly take place, and are of extreme physiological importance. If the surface of the skin, for example, be at any time unexpectedly brought in contact with a heated body, the injured part is often withdrawn by a rapid and convulsive movement, long before we feel the pain, or even understand fairly the cause of the involuntary act. If the body, by any accident, suddenly and unexpectedly lose its balance, the limbs are thrown into a position calculated to protect the ex- 23 354 THE SPIXAL CORD. posed parts and to break the fall, by a similar involuntary and in- stantaneous movement. The brain does not act in these cases, for there is no intentional character in the movement, nor even any complete consciousness of its object. Everything indicates that it is the immediate result of a simple reflex action of the spinal cord. The cord exerts also an important and constant influence upon the sphincter muscles. The sphincter ani is habitually in a state of contraction, so that the contents of the intestine are not allowed to escape. When any external irritation is applied to the anus, or whenever the feces present themselves internally, the sphincter contracts involuntarily, and the discharge of the feces is prevented. This habitual closure of the sphincter depends on the reflex action of the spinal cord. It is entirely an involuntary act, and will con- tinue, in the healthy condition, during profound sleep, as complete and efficient as in the waking state. When the rectum, however, has become filled by the accumula- tion of feces from above, the nervous action changes. Then the impression produced on the mucous membrane of the distended rectum, conveyed to the spinal cord, causes at the same time re- laxation of the sphincter and contraction of the rectum itself; so that a discharge of the feces consequently takes place. Now all these actions are to some extent under the control of sensation and volition. The distended state of the rectum is usually accompanied by a distinct sensation, and the resistance of the sphincter may be voluntarily prolonged for a certain period, just as the respiratory movements, which are usually involuntary, may be intentionally hastened or retarded, or even temporarily suspended. But this voluntary power over the sphincter and the rectum is limited. After a time the involuntary impulse, growing more and more urgent with the increased distension of the rectum, becomes irresistible; and the discharge finally takes place by the simple reflex action of the spinal cord. If the spinal cord be injured in its middle or upper portions, the sensibility and voluntary action of the sphincter is lost, because its connection with the brain has been destroyed. The evacuation then takes place at once, by the ordinary mechanism, as soon as the rectum is filled, but without any knowledge on the part of the patient. The discharges are then said to be " involuntary and un- conscious." If the irritability of the cord, on the other hand, be exaggerated by disease, while its connection with the brain remains entire, the KEFLEX ACTION OF THE SrilSTAL CORD. 355 distension of the rectum is announced by the usual sensation, but the reflex impulse to evacuation is so urgent that it cannot be controlled bj the will, and the patient is compelled to allow it to take place at once. The discharges are then said to be simply '* involuntary." Finally, if the substance of the spinal cord be extensively de- stroyed by accident or disease, the sphincter is permanently relaxed. The feces are then evacuated almost continuously, without any knowledge or control on the part of the patient, as fast as they descend into the rectum from the upper portions of the intestine. Injury of the spinal cord produces a somewhat different effect on the urinary bladder. Its muscular fibres are directly para- lyzed ; and the organ, being partially protected by elastic fibres, both at its own orifice and along the urethra, becomes gradually distended by urine from the kidneys. The urine then overcomes the elasticity of the protecting fibres, by simple force of accumula- tion, and afterward dribbles away as fast as it is excreted by the kidneys. Paralysis of the bladder, therefore, first causes a perma- nent distension of the organ, which is afterward followed by a continuous, passive, and incomplete discharge of its contents. Injury of the spinal cord produces also an important, though probably an indirect effect on nutrition, secretion, animal heat, &c., in the paralyzed parts. Diseases of the cord which result in its softening or disintegration, are notoriously accompanied by consti- pation, often of an extremely obstinate character. In complete paraplegia, also, the lower extremities become emaciated. The texture and consistency of the muscles are altered, and the animal temperature is considerably reduced. All such disturbances of nutrition, however, which follow almost invariably upon local para- lysis are no doubt immediately owing to the inactive condition of the muscles; a condition which naturally induces debility of the circulation, and consequently of all those functions which are dependent upon it. It is less easy to explain the connection between injury of the spinal cord and inflammation of the urinary passages. It is, how- ever, a matter of common observation among pathologists, that injury or disease of the cord, particularly in the dorsal and upper lumbar regions, is soon followed by catarrhal inflammation of the urinary passages. This gives rise to an abundant production of altered mucus, which in its turn, by causing an alkaline fermenta- tion in the urine contained in the bladder, converts it into an irri- 356 THE SPINAL CORD. tating and ammoniacal liquid, which reacts upon the nnucous membrane and aggravates the previous inflammation. We find, therefore, that the spinal cord, in its character of a nervous centre, exerts a general protective action over the whole body. It presides over the involuntary movements of the limbs and trunk ; it regulates the action of the sphincters, the rectum, and the bladder; while at the same time it exerts an indirect influ- ence on the nutritive changes in those parts which it supplies with nerves. THE BRAIN. 857 CHAPTER IV. THE BRAIN. By the brain, or encephalon, as it is sometimes called, we mean all that portion of the nervous system which is situated within the cavity of the cranium. It consists, as we have already shown, of a series of different ganglia, connected with each other by transverse and longitudinal commissures. Since we have found the functions of sensation and motion, or sensibility and excitability, so distinctly separated in the spinal cord, we should expect to find the same distinction in the interior of the brain. These two properties have indeed been found to be distinct from each other, so far as they exist at all, in the encephalic mass ; but it is a very remarkable fact that they are both confined to very small portions of the brain, in comparison with its entire bulk. According to the investigations of Longet, neither the olfactory ganglia, the corpora striata, the optic thalami, the tuber- cula quadrigemina, nor the white or gray substance of the cerebrum or the cerebellum, are in the least degree excitable. Mechanical irritation of these parts does not produce the slightest convulsive movement in the muscles below. The application of caustic liquids and the passage of galvanic currents are equally without effect. The only portions of the brain in which irritation is followed by convulsive movements are the anterior surface of the medulla ob- longata, the tuber annulare, and the lower part of the crura cerebri; that is, the lower and central parts of the brain, containing continu- ations of the anterior columns of the cord. On the other hand, neither the olfactory ganglia, the corpora striata, the tubercula quadrigemina nor the white or gray substance of the cerebrum or cerebellum, give rise, on being irritated, to any painful sensation. The only sensitive parts are the posterior surfiice of the medulla oblongata, the restiform bodies, the processus e cerebello ad testes, and the upper part of the crura cerebri ; that is, those portions of 358 THE BRAIN". the base of the brain which contain prolongations of the posterior columns of the cord The most central portions of the nervous system, therefore, and particularly the gray matter, are destitute of both excitability and sensibility. It is only those portions which serve to conduct sen- sations and nervous impulses that can be excited by mechanical irritation; not the ganglionic centres themselves, which receive and originate the nervous impressions. "We shall now study in succession the different ganglia of which the brain is composed. OLFACTORY GANGLIA. These ganglia, which in some of the lower animals are very large, corresponding in size with the extent of the Schneiderian mucous membrane and the acuteness of the sense of smell, are very small in the human subject. They are situated on the cribriform plate of the ethmoid bone, on each side of the crista galli, just be- neath the anterior lobes of the cerebrum. They send their nerves through the numerous perforations which exist in the ethmoid bone at this part, and are connected with the base of the brain by two longitudinal commissures. The olfactory ganglia with their com- missures are sometimes spoken of as the "olfactory nerves." They are not nerves, however, but ganglia, since they are mostly com- posed of gray matter ; and the term " olfactory nerves" can be properly applied only to the filaments which originate from them, and which are afterward spread out in the substance of the Schnei- derian mucous membrane. It has been found difficult to determine the function of these ganglia by direct experiment on the lower animals. They may be destroyed by means of a strong needle introduced through the bones of the cranium ; but the signs of the presence or absence of the sense of smell, after such an operation, are too indefinite to allow us to draw from them a decided conclusion. The anatomical distribu- tion of their nerves, however, and the evident correspondence which exists, in different species of animals, between their degree of de- velopment and that of the external olfactory organs, leaves no doubt as to their true function. They are the ganglia of the special sense of smell, and are not connected, in any appreciable degree, with OPTIC THALAMI, — CORPORA STRIATA. — HEMISPHERES. 859 ordinary sensibility, nor with the production of voluntary move- ments. OFTIC THALAMI. These bodies are not, as their name would imply, the ganglia of vision. Longet has found that the power of sight and the sensi- bility of the pupil both remain, in birds, after the optic thalami have been thoroughly disorganized ; and that artificial irritation of the same ganglia has no effect in producing either contraction or dilatation of the pupil. The optic thalami, however, according to the same observer, have a peculiar crossed action upon the volun- tary movements. If both hemispheres and both optic thalami be removed in the rabbit, the animal is still capable of standing and of "using his limbs in progression. But if the right optic thalamus alone be removed, the animal falls at once upon his left side; and if the left thalamus be destroyed, a similar debility is manifest on the right side of the body. In these instances there is no absolute paralysis of the side upon which the animal falls, but rather a simple want of balance between the two opposite sides. The exact mechanism of this peculiar functional disturbance is not well understood; and but little light has yet been thrown, either by direct experiment or by the facts of comparative anatomy, on the real function of the optic thalami. CORPORA STRIATA. The function of these ganglia is equally uncertain with that of the preceding. They are traversed, as we have already seen, by fibres coming from the anterior columns of the cord; and they are connected, by the continuation of these fibres, with the gray sub- stance of the hemispheres. They have, therefore, in all probability, like the optic thalami, some connection with sensation and volition; but the precise nature of this connection is at present altogether unknown. HEMISPHERES. The hemispheres, or the cerebral ganglia, constitute in the human subject about nine tenths of the whole mass of the brain. 360 THE BRAIN. Throughout their whole extent they are entirely destitute, as we have already mentioned, of both sensibility and excitability. Both the white and gray substance may be wounded, burned, lacerated, crushed, or galvanized in the living animal, without exciting any convulsive movement or any apparent sensation. In the human subject a similar insensibility has been observed when the sub- stance of the hemispheres has been exposed by accidental violence, or in the operation of trephining. Yery severe mechanical injuries may also be inflicted upon the hemispheres, even in the human subject, without producing any directly fatal result. One of the most remarkable instances of this fact is a case reported by Dr. William Detmold, of New York,' in which an abscess in the anterior lobe of the brain was opened by an incision passing through the cerebral substance, not only without any immediate bad effect, but with great temporary relief to the patient. This was the case of a laborer who was struck on the left side of the forehead by a piece of falling timber, which produced a compound fracture of the skull at this part. One or two pieces of bone afterward became separated and were removed, and the wound subsequently healed. Nine weeks after the accident, how- ever, headache and drowsiness came on ; and the latter symptom, becoming rapidly aggravated, soon terminated in complete stupor. At this time, the existence of an abscess being suspected, the cicatrix, together with the adherent portion of the dura mater, was dissected away, several pieces of fractured bone removed, and the surface of the brain exposed. A knife was then passed into the cerebral substance, making a wound one inch in length and half an inch in depth, when the abscess was reached and about liij of pus discharged. I^he patient immediately aroused from his coma- tose condition, so that he was able to speak; and in a few days recovered, to a very considerable extent, his cheerfulness, intelli- gence, and appetite. Subsequently, however, the collection of pus returned, accompanied by a renewal of the previous symptoms; and the patient finally died at the end of seven weeks from the time of opening the abscess. Another and still more striking instance of recovery from severe injury of the brain is reported by Prof. H. J. Bigelow in the American Journal of Medical Sciences for July, 1850. In this case, a pointed iron bar, three feet and a half in length, and one inch and a ' Am. Joiirn. of Med. Sci., January, 1850. HEMISPHERES. 361 quarter in diameter, was driven through the patient's head by the premature blasting of a rock. The bar entered the left side of the face, just in front of the angle of the jaw, and passed obliquely upward, inside the zygomatic arch and through the anterior part of the cranial cavity, emerging from the top of the frontal bone on the median line, just in front of the point of union of the coronal and sagittal sutures. The patient was at first stunned, but soon recovered himself so far as to be able to converse intelligently, rode home in a common cart, and with a little assistance walked up stairs to his room. He became delirious within two days after the acci- dent, and subsequently remained partly delirious and partly coma- tose for about three weeks. He then began to improve, and at the end of rather more than two months from the date of the injury, was able to walk about. At the end of sixteen months he was in perfect health, with the wounds healed, and with the mental and bodily functions entirely unimpaired, except that sight was perma- nently lost in the eye of the injured side. The hemispheres, furthermore, are not the seat of sensation or of volition, nor are they immediately essential to the continuance of life. In quadrupeds, the complete removal of the hemispheres is attended with so much hemorrhage that the operation is generally fatal from this cause within a few minutes. In birds, however, it may be performed without any immediate danger to life. Longet has removed the hemispheres in pigeons and fowls, and has kept these animals afterward for several days, with most of the organic functions unimpaired. We have frequently performed the same experiment upon pigeons, with a similarly favorable result. The effect of this mutilation is simply to plunge the animal into a state of profound stupor, in which he is almost entirely inatten- tive to surrounding objects. The bird remains sitting motionless upon his perch, or standing upon the ground, with the eyes closed, and the head sunk between the shoulders. (Fig. 138.) The plu- mage is smooth and glossy, but is uniformly expanded, by a kind of erection of the feathers, so that the body appears somewhat puffed out, and larger than natural. Occasionally the bird opens his eyes with a vacant stare, stretches his neck, perhaps shakes his bill once or twice, or smooths down the feathers' upon his shoulders, and then relapses into his former apathetic condition. This state of immobility, however, is not accompanied by the loss of sight, of hearing, or of ordinary sensibility. All these functions remain, as well as that of voluntary motion. If a pistol be discharged behind 362 THE BRAIN. the back of the animal, he at once opens his eyes, moves his head half round, and gives evident signs of having heard the report; but he immediately becomes quiet, again, and pays no farther attention to it. Sight is also retained, since the bird will sometimes fix its Fig. 138. PiGEOX, AFTER REMOVAL OF THE HEMISPHERES. eye on a particular object, and watch it for several seconds together. Longet has even found that by moving a lighted candle before the animal's eyes in a dark place, the head of the bird will often follow the movements of the candle from side to side or in a circle, showing that the impression of light is actually perceived by the sensoriura. Ordinary sensation also remains, after removal of the hemispheres, together with voluntary motion. If the foot be pinched with a pair of forceps, the bird becomes partially aroused, moves uneasily once or twice from side to side, and is evidently annoyed at the irritation. The animal is still capable, therefore, after removal of the hemi- spheres, of receiving sensations from external objects. But these sensations appear to make upon him no lasting impression. He is incapable of connecting with his perceptions any distinct succession of ideas. He hears, for example, the report of a pistol, but he is not alarmed by it ; for the sound, though distinctly enough perceived, does not suggest any idea of danger or injury. There is accord- ingly no power of forming mental associations, nor of perceiving the relation between external objects. The memory, more particu- larly, is altogether destroyed, and the recollection of sensations is not retained from one moment to another. The limbs and muscles are still under the control of the will ; but the will itself is inactive. HEMISPHERES. 363 because apparently it lacks its usual mental stimulus and direction. The powers whicli have been lost, therefore, by destruction of the cerebral hemispheres, are altogether of a mental or intellectual character; that is, the power of comparing with each other different ideas, and of perceiving the proper relation between them. The same result is well known to follow, in the human subject, from injury or disease of these parts. A disturbance of the mental powers has long been recognized as the ordinary consequence of lesions of the brain. In cases of impending apoplexy, for example, or of softening of the cerebral substance, among the earliest and most constant phenomena is a loss or impairment of the memory. The patient forgets the names of particular objects or of particular persons ; or he is unable to calculate numbers with his usual facility. His mental derangement is often shown in the undue estimate which he forms of passing events. He is no longer able to appreciate the true relation between different objects and different phenomena. Thus, he will show an exaggerated degree of solicitude about a trivial occurrence, and will pay no attention to other matters of real importance. As the difficulty increases, he becomes careless of the directions and advice of his attendants, and must be watched and managed like a child or an imbecile. After a certain period, he no longer appreciates the lapse of time, and even loses the dis- tinction between day and night. Finally, when the injury to the hemispheres is complete, the senses may still remain active and impressible, while the patient is completely deprived of intelligence, memory, and judgment. If we examine the comparative development of the hemispheres in different species of animals, and in different races of men, we shall find that the size of these ganglia corresponds very closely with the degree of intelligence possessed by the individual. We have already traced, in a preceding chapter, the gradual increase in size of the hemispheres in fish, reptiles, birds and quadrupeds: four classes of animals which may be arranged, with regard to the amount of intelligence possessed by each, in precisely the same order of succession. Among quadrupeds, the elephant has much the largest and most perfectly formed cerebrum, in proportion to the size of the entire body; and of all quadrupeds he is proverbially the most intelligent and the most teachable. It is important to observe, in this connection, that the kind of intelligence which characterizes the elephant and some other of the lower animals, and which most nearly resembles that of man, is a teachable intelli- 364 THE BRAIN. gence; a very different thing from the intelligence which depends upon instinct, such as that of insects, for example, or birds of pas- sage. Instinct is unvarying, and always does the same thing in the same manner, with endless repetition ; but intelligence is a power which adapts itself to new circumstances, and enables its possessor, by comprehending and retaining new ideas, to profit by experience. It is this quality which distinguishes the higher classes of animals from the lower; and which, in a very much greater degree, con- stitutes the intellectual superiority of man himself. The size of the cerebrum in man is accordingly very much greater, in pro- portion to that of the entire body, than in any of the lower animals; while other parts of the brain, on the contrary, such as the olfactory ganglia or the optic tubercles, are frequently smaller in him than in them. For while man is superior in general intelligence to all the lower animals, he is inferior to many of them in the acuteness of tlie special senses. As a general rule, also, the size of the cerebrum in different races and in different individuals corresponds with the grade of their intelligence. The size of the cranium, as compared with that of the face, is smallest in the savage negro and Indian tribes; larger in the civilized or semi-civilized Chinese, Malay, Arab, and Japan- ese; while it is largest of all in the enlightened European races. This difference in the development of the brain is not probably an effect of long-continued civilization or otherwise; but it is, on the contrary, the superiority in cerebral development which makes some races readily susceptible of civilization, while others are either altogether incapable of it, or can only advance in it to a certain limit. Although all races therefore may, perhaps, be said to start from the same level of absolute ignorance, yet after the lapse of a certain time one race will have advanced farther in civilization than another, owing to a superior capacity for improve- ment, dependent on original organization. The same thing is true with regard to different individuals. At birth, all men are equally ignorant ; and yet at the end of a certain period one will have acquired a very much greater intellectual power than another, even under similar conditions of training, education, &c. He has been able to accumulate more information from the same sources, and to use the same experience to better advantage than his associates ; and the result of this is a certain intellectual superiority, which becomes still greater by its own exercise. This superiority, it will be observed, lies not so much HEMISPHERES. 365 in tlie power of perceiving external objects and events, and of recognizing the connection between tliem, as in that of drawing conclusions from one fact to another, and of adapting to new com- binations the knowledge which has already been acquired. It is this particular kind of intellectual difference, existing in a marked degree between animals, races, and individuals, which cor- responds with the difference in development of the cerebral hemi- spheres. We have, therefore, evidence from three different sources that the cerebral hemispheres are the seat of the reasoning powers,. or of the intellectual faculties proper. First, when these ganglia are removed, in the lower animals, the intellectual faculties are the only ones which are lost. Secondly, injury to these ganglia, in the human subject, is followed by a corresponding impairment of the same faculties. Thirdly, in different species of animals, as well as in different races of men and in different individuals, the develop- ment of these faculties is in proportion to that of the cerebral hemispberes. When we say, however, that the hemispheres are the seat of the intellectual faculties, of memory, reason, judgment, and the like, we do not mean that these faculties are, strictly speaking, located in the substance of the hemispheres, or that they belong directly to the matter of which the hemispheres are composed. The hemi- spherical ganglia are simply the instruments through which the intellectual powers manifest themselves, and which are accordingly necessary to their operation. If these instruments be imperfect in structure, or be damaged in any manner by violence or disease, the manifestations of intelligence are affected in a corresponding degree. So far, therefore, as the mental faculties are the subject of physio- logical research and experiment, they are necessarily connected with the hemispherical ganglia; and the result of investigation shows this connection to be extremely intimate and important in its character. There are, however, various circumstances which modify, in particular cases, the general rule given above, viz., that the larger the cerebrum the greater the intellectual superiority. The func- tional activity of the brain is modified, no doubt, by its texture as well as by its size ; and an increased excitability may compensate, partially or wholly, for a deficiency in bulk. This fact is some- times illustrated in the case of idiots. There are instances where idiotic children with small brains are less imbecile and helpless than others with a larger development, owing to a certain vivacity 366 THE BRAIl^. and impressibility of organization which take the place, to a cer- tain extent, of the purely intellectual faculties. This was the case, in a marked degree, with a pair of dwarfed and idiotic Central American children, who were exhibited a few years ago in various parts of the United States, under the name of the "Aztec children." They were a bo}' and a girl, aged respectively about seven and five years. The boy was 2 feet 9| inches high, and weighed a little over 20 pounds. The girl was 2 feet 5| inches high, and weighed 17 pounds. Their bodies were tolerably well proportioned, but the cranial cavities, as shown by the accompan}^- ing portraits, were extremely small. Fig. 139. Aztec Children.— Taken from life. The antero-posterior diameter of the boy's head was only 4| inches, the transverse diameter less than 4 inches. The antero- posterior diameter of the girl's head was 4J inches, the transverse diameter only 3| inches. The habits of these children, so far as regards feeding and taking care of themselves, were those of chil- dren two or three years of age. They were incapable of learning to talk, and could only repeat a few isolated words. Notwithstand- ing, however, the extremely limited range of their intellectual powers, these children were remarkably vivacious and excitable. While awake they were in almost constant motion, and any new object or toy presented to them immediately attracted their atten- tion, and evidently awakened a lively curiosity. They were ac- cordingly easily influenced by proper management, and understood readily the meaning of those who addressed them, so far as this meaning could be conveyed by gesticulation and the tones of the voice. Their expression and general appearance, though decidedly idiotic, were not at all disagreeable or repulsive ; and they were HEMISPHEEES. 367 much less troublesome to the persons who had them in charge than is often the case with idiots possessing a larger cerebral development. It may also be observed that the purely intellectual or reasoning powers are not the only element in the mental superiority of certain races or of particular individuals over their associates. There is also a certain rapidity of perception and strength of will which may sometimes overbalance greater intellectual acquirements and more cultivated reasoning powers. These, however, are differ- ent faculties from the latter ; and occupy, as we shall hereafter see, different parts of the encephalon. A very remarkable physiological doctrine, dependent partly on the foregoing facts, was brought forward some years ago by Gall and Spurzheim, under the name of Phrenology. These observers recognized the fact that the intellectual powers are undoubtedly seated in the brain, and that the development of the brain is, as a general rule, in correspondence with the activity of these powers. They noticed also that in other parts of the nervous system, different functions occupy different situations ; and regarding the mind as made up of many distinct mental faculties, they conceived the idea that these different faculties might be seated in different parts of the cerebral mass. If so, each separate portion of the brain would undoubtedly be more or less developed in proportion to the activity of the mental trait or faculty residing in it. The shape of the head would then vary in different individuals, in accordance with their mental peculiarities; and the character and endowments of the individual might therefore be estimated from an examination of the elevations and depressions on the surface of the cranium. Accordingly, the authors of this doctrine endeavored, by examin- ing the heads of various individuals whose character was already known, to ascertain the location of the different mental faculties. In this manner they finally succeeded, as they supposed, in accom- plishing their object; after which they prepared a chart, in which the surface of the cranium was mapped out into some thirty or forty different regions, corresponding with as many different mental traits or faculties. With the assistance of this chart it was thought that phrenology might be practised as an art ; and that, by one skilled in its application, the character of a stranger might be discovered by simply examining the external conformation of his head. We shall not expend much time in discussing the claims of phre- nology to rank as a science or an art, since we believe that it has of late years been almost wholly discarded by scientific men, owing 368 THE BRAIN. to the very evident deficiencies of the basis upon which it was founded. Passing over, therefore, many minor details, we will merely point out, as matters of physiological interest, the principal defects which must always prevent the establishment of phrenology as a science, and its application as an art. First, though we have no reason for denying that different parts of the brain may be occupied by different intellectual faculties, there is no direct evidence which would show this to be the case. Phrenologists include, in those parts of the brain which they em- ploy for examination, both the cerebrum and cerebellum ; and they justly regard the external parts of these bodies, viz., the layer of gray matter which occupies their surface, as the ganglionic portion in which must reside more especially the nervous functions which they possess. But this layer of gray matter, in each principal por- tion of the brain, is continuous throughout. There is no anatomical division or limit between its different parts, as there are between the different ganglia in other portions of the nervous system ; and consequently such divisions of the cerebrum and cerebellum must be altogether arbitrary in character, and not dependent on any anatomical basis. Secondly, the only means of ascertaining the location of the different mental traits, supposing them to occupy different parts of the brain, would be that adopted by Gall and Spurzheim, viz., to make an accurate comparison, in a sufficient number of cases, of the form of the head in individuals of known character. But the prac- tical difficulty of accomplishing this is very great. It requires a long acquaintance and close observation to learn accurately the character of a single person; and it is in this kind of observation, more than in any other, that we are proverbially liable to mis- takes. It is extremely improbable, therefore, that either Gall or Spurzheim could, in a single lifetime, have accomplished this com- parison in so many instances as to furnish a reliable basis for the construction of a phrenological chart. A still more serious practical difficulty, however, is the following. The different intellectual faculties being supposed to reside in the layer of gray substance constituting the surfaces of the cerebrum and cerebellum, they must of course be distributed throughout this layer, wherever it exists. Gall and Spurzheim located all the mental faculties in those parts of the brain which are accessible to external exploration. An examination of different sections of the brain will show, however, that the greater portion of the gray substance is so placed, that its quantity cannot be estimated by an external HEMISPHEEES. 869 Fig. 140. examination througli the skull. The only portions which are ex- posed to such an examination are the upper and lateral portions of the convexities of the hemispheres, together with the posterior edge and part of the under surface of the cerebellum. (Fig. 140.) A verj extensive portion of the cerebral surface, however, remains concealed in such a manner that it cannot possibly be subjected to examination, viz., the entire base of the brain, with the under surface of the anterior and middle lobes (1,2 ); the upper surface of the cerebellum (3) and the inferior surface of the posterior lobe of the cerebrum which covers it (4); that portion of the cerebellum situated above the medulla oblongata (5); and the two opposite convoluted surfaces in the fissure of Sylvius (e, 7), where the ante- rior and middle lobes of the cerebrum lie in contact with each other. The whole extent, also, of the cerebral surfaces which are opposed to each other in the great longitudinal fissure (Fig. 141), throughout its entire length, are equally protected by their position, and concealed from external exa- mination. The whole of the convoluted surface of the brain must, however, be re- garded as of equal importance in the distri- bution of the mental qualities ; and yet it is evident that not more than one-third or one- quarter of this surface is so placed that it can be examined by external manipulation. It must furthermore be recollected that the gray matter of the cerebrum and cerebellum is everywhere convoluted, and that the convolutions penetrate to various depths in the substance of the brain. Even if we were able to feel, therefore, the external surface of the brain itself, it would not be the entire convolutions, but only their superficial edges, that we should really be able to examine. And yet the amount of gray matter contained in a given space depends quite as much upon the depth to which the convolutions penetrate, as upon the prominence of their edges. 24 Diagram of the B k A i n in situ, showing those portious which are ex- posed to examination. Fig. 141. Transverse section of B r a t x , showing depth of great longi- tudinal fissure, at a. 370 THE BRAIN. While phrenology, therefore, is partially founded upon acknow- ledged physiological facts, there are yet essential deficiencies in its scientific basis, as well as insurmountable difficulties in the way of its practical application. CEREBELLUM. The cerebellum is the second ganglion of the encephalon, in respect of size. If it be examined, moreover, in regard to the form and disposition of its convolutions, it will be seen that these are much more complicated and more numerous than in the cerebrum, and penetrate much deeper into its substance. Though the cerebel- lum therefore is smaller, as a whole, than the cerebrum, it contains, in proportion to its size, a much larger quantity of gray matter. In examining the comparative development of the brain, also, in different classes and species of animals, we find that the cerebellum nearly always keeps pace, in this respect, with the cerebrum. These facts would lead us to regard it as a ganglion hardly secondary in importance to the cerebrum itself. Physiologists, however, have thus far failed to demonstrate the nature of its function with the same degree of precision as that of many other parts of the brain. The opinion of Gall, which located in the cerebellum the sexual impulse and instincts, is at the present day generally abandoned; for the reason that it has not been found to be sufficiently supported by anatomical and experimental facts, many of which are indeed directly opposed to it. The opinion which has of late years been received with the most favor is that first advocated by Flourens, which attributes to the cerebellum the power of associating or "co-ordinating" the different voluntary movements. It is evident, indeed, that such a power does actually reside in some part of the nervous system. No movements are effected by the independent contraction of single muscles; but always by several muscles acting in harmony with each other. The number and complication of these associated movements vary in different classes of animals. In fish, for example, progression is accom- plished in the simplest possible manner, viz., by the lateral flexion and extension of the vertebral column. In serpents it is much the same. In frogs, lizards, and turtles, on the other hand, the four jointed extremities come into play, and the movements are some- CEKEBELLUM. 371 what complicated. They are still more so in birds and quadrupeds; and finally, in the human subject they become both varied and complicated in the highest degree. Even in maintaining the ordi- nary postures of standing and sitting, there are many different mus- cles acting together, in each of which the degree of contraction, in order to preserve the balance of the body, must be accurately pro- portioned to that of the others. In the motions of walking and running, or in the still more delicate movements of the hands and fingers, this harmony of muscular action becomes still more evident, and is seen also to be absolutely indispensable to the efficiency of the muscular apparatus. The opinion which locates the above harmonizing or associating power in the cerebellum was first suggested by the effects observed after experimentally injuring or destroying this part of the brain. If the cerebellum be exposed in a living pigeon, and a portion of its substance removed, the animal exhibits at once a peculiar un- certainty in his gait, and in the movement of his wings. If the injury be more extensive, he 'loses altogether the power of flight, and can walk, or even stand, only with great difficulty. This is not owing to any actual paralysis, for the movements of the limbs are exceedingly rapid and energetic; but is due to a peculiar want of control over the muscular contractions, precisely similar to that which is seen in a man in a state of intoxication. The movements of the legs and wings, though forcible and rapid, are confused and blundering; so that the animal cannot direct his steps to any par- ticular spot, nor support himself in the air by flight. He reels and tumbles, but can neither walk nor fly. The senses and intelligence at the same time are unimpaired. It is extremely curious, as first remarked by Longet, to compare the different phenomena produced by removal of the cerebrum and by that of the cerebellum. If we do these operations upon two dif- ferent pigeons, and place the animals side by side, it will be seen that the first pigeon, from whom the cerebrum only has been re- moved, remains standing firmly upon his feet, in a condition of complete repose ; and that when aroused and compelled to stir, he moves sluggishly and unwillingly, but acts otherwise in a perfectly natural manner. The second pigeon, on the other hand, from whom the cerebellum only has been taken away, is in a constant state of agitation. He is easily terrified, and endeavors, frequently with violent struggles, to escape the notice of those who are watching him ; but his movements are sprawling and unnatural. 372 THE BRAIN". and are evidently no longer under the effectual control of the will, (B'ig. 142.) If the entire cerebellum be destroyed, the animal is no longer capable of assuming or retaining any natural posture. His legs and wings are almost constantly agitated with ineffectual Fig. 142. PrOEON, AFTER REMOVAL OF THE CeREBELI, CM. struggles, which are evidently voluntary in character, but are at the same time altogether irregular and confused. Death generally takes place after this operation within twenty-four hours. The results of the above experiment are extremely constant and invariable, and by themselves would lead us to adopt, with a good degree of confidence, the opinion of Flourens. This opinion evi- dently has more direct evidence in its favor than any other theory which has yet been broached with regard to the function of the cerebellum. Many facts derived from comparative anatomy tend, also, to confi.rm the same opinion. If we compare different classes of animals with each other, as fish with reptiles, or birds with quad- rupeds, in which the development and activity of the entire nervous system vary extremely, the results of the comparison will be often contradictory; but if we compare different species belonging to the same class and order, in which the general structure and plan of organization are nearly the same, we often find the development of the cerebellum to correspond very closely with the perfection and variety of the muscular movements. The frog, for example, is an aquatic reptile provided with anterior and posterior extremi- ties ; but its movements, though rapid and vigorous, are exceed- ingly simple in character, consisting of little else than flexion and TUBERCULA QUADRIGEMINA. 373 extension of the posterior limbs. The cerebellum in this animal is exceedingly small, compared with the rest of the brain ; being nothing more than a thin, narrow ribbon of nervous matter, stretched across the upper part of the fourth ventricle. In the common turtle we have another aquatic reptile, where the move- ments of swimming, diving, progression, &c., are accomplished by the consentaneous action of both anterior and posterior extremities, and where tbe motions of the head and neck are also mucb more varied than in the frog. In this instance the cerebellum is very much more highly developed than in the former. In the alli- gator, again, a reptile whose motions, both of the head, limbs, and tail, approach very closely to those of the quadrupeds, the cerebellum is still larger in proportion to the remaining ganglia of the encephalon. In the above instances, therefore, an evident correspondence exists between the size of the cerebellum and the variety of move- ment of which the animal is capable. Still, this part of the subject has not yet been sufficiently investigated to enable us to say that such a correspondence exists in all cases. Morbid alterations of the cerebellum, furthermore, such as inflammations, abscess, tu- mors, &c., have not always been found to produce, in the human subject, symptoms connected with a loss of harmony in the volun- tary movements. The complete function of the cerebellum., ac- cordingly, cannot yet be regarded as positively ascertained ; but so far as we may rely on the results of direct experiment, and on the general facts of comparative anatomy, the most plausible opinion is that of Flourens, viz., that the cerebellum possesses the power of uniting and harmonizing the action of separate muscles, so that they may assist each other in the production of varied and com- plicated movements. TUBERCULA QUADRIGEMINA. These bodies, notwithstanding their small size, are very important in regard to their function. They give origin to the optic nerves, and preside, as ganglia, over the sense of sight; on which account they are also known by the name of the " optic ganglia." Their development corresponds very closely with that of the external organs of vision. Thus, they are large in fish, reptiles, and birds, in which the eyeball is for the most part very large in proportion 874 • THE BEAIN". to the entire head ; and are small in quadrupeds and in man, where the eyeball is, comparatively speaking, of insignificant size. Direct experiment also shows the close connection between the tubercula quadrigemina and the sense of sight. Section of the optic nerve at any point between the retina and the tubercles, pro- duces complete blindness; and destruction of the tubercles them- selves has the same effect. But if the division be made between the tubercles and the cerebrum, or if the cerebrum itself be taken away while the tubercles are left untouched, vision, as we have already seen, still remains. It is the tubercles, therefore, in which the impression of light is perceived. So long as these ganglia are uninjured and retain their connection with the eye, vision remains. As soon as this connection is cut off, or the ganglia themselves are injured, the power of vision is destroyed. The tubercula quadrigemina not only serve as nervous centres for the perception of light, but a reflex action also takes place through them, by which the quantity of light admitted to the eye is regulated to suit the sensibility of the pupil. In darkness and in twilight, or wherever the light is obscure and feeble, the pupil is enlarged by a relaxation of its circular fibres, so as to admit as large a quantity of light as possible. On first coming into a dark room, accordingly, everything is nearly invisible ; but gradually, aS the pupil dilates and as more light is admitted, objects begin to show themselves with greater distinctness, and at last we can see tolerably well in a place where we were at first unable to perceive a single object. On the other hand, when the eye is exposed to an unusually brilliant light, the pupil contracts and shuts out so much of it as would be injurious to the retina. The above is a reflex action, in which the impression received by the retina is transmitted along the optic nerve to the tubercula quadrigemina. From the tubercles, a motor impulse is then sent out through the motor nerves of the eye and the filaments dis- tributed to the iris, and a contraction of the pupil takes place in consequence. The optic nerves act here as sensitive fibres, which convey the impression from the retina to the ganglion; and if they be irritated in any part of their course with the point of a needle, the result is a contraction of the pupil. This influence is not communicated directly from the nerve to the iris, but is first sent inward to the tubercles, to be afterward reflected outward by the motor nerves. So long as the eyeball remains in connection with the brain, mechanical irritation of the optic nerve, as we have TUBEKCULA QUADRIGEMINA. 375 shown above, causes contraction of the pupil; but if the nerve be divided, and the extremity which remains in connection with the eyeball subjected to irritation, no effect upon the pupil is produced. The anatomical arrangement of the optic nerves, and the connec- tions of the optic tubercles, are modified in a remarkable degree in different animals, to correspond with the position of the two eyes. In fish, for example, the eyes are so placed, on opposite sides of the head, that their axes cannot be brought into parallelism with each other, and the two eyes can never be directed together to the same object. In these animals, the optic nerves cross each other at the base of the brain without any intermixture of their fibres; that from the right optic tubercle passing to the left eye, and that from the left optic tubercle passing to the right eye. (Fig. 148.) The two nervous cords are here totally distinct from each other throughout their entire length ; and are only connected, at the point of cross- ing, by intervening areolar tissue. Impressions made on the right eye must therefore be perceived on the left side of the brain; while those which enter the left eye are conveyed to the right side of the brain. Fig. 143. Fig. 144. Inferior Surface of Brain OF Cod. — 1. Eight optic nerve. 2. Left optic nerve. 3. Right optic tubercle. 4. Left optic tubercle. 6, 6. Hemispheres. 7. Medulla oblongata. Infkrior Surface of Brain op Fowl. — \. Right optic nerve. 2. Left optic nerre 3. Right optic tubercle. 4. Left optic tubercle. 6, 6. Hemispheres. 7. Me- dulla oblongata. In birds, also, the axes of the two eyes are so widely divergent that an object cannot be distinctly in focus for both of them at the 376 THE BRAIlSr. same time. The optic nerves are here united, and apparently sol- dered together, at their point of crossing; but the decussation of their fibres is nevertheless complete. (Fig. 144.) The nervous fila- ments coming from the left side pass altogether over to the right ; and those coming from the right side pass over to the left. The result of direct experiment on the crossed action of the tubercles in these animals corresponds with the anatomical arrangement of the nervous fibres. If one of the optic tubercles be destroyed in the pigeon, complete blindness is at once produced in the eye of the opposite side; but vision remains unimpaired in the eye of the same side with the injury. In the human subject, on the other hand, where the visual axes are parallel, and where both eyes are simultaneously directed to the same object, the optic nerves decussate with each other in such a manner as to form a connection between the two opposite sides, as well as between each tubercle and retina of the same side. (Fig. 145.) This decussation, which is somewhat complicated, takes place Fig. 145. CotTRSE OP Optic Nerves in Man. — 1, 2. Eight and left eyeballs. 3. Decussation of optic nerves. 4, i. Tubercula quadrigemina. in the following manner. From each optic tubercle three difi'erent bundles or " tracts" of nervous fibres are given ofi". One set passes TUBER ANNULARE. 377 across transversely at the point of decussation, and turning back- ward, terminates in the tubercle of the opposite side; another, cross ing diagonally, continues onward to the opposite eyeball; while a third passes directly forward to the eyeball of the same side. A fourth set of fibres, still, passes across, in front of the decussation, from the retina of one eye to that of the opposite side. We have, therefore, by this arrangement, the two retinae, as well as the two optic tubercles, connected with each other by commissural fibres ; while each tubercle is, at the same time, connected both with its own retina and with that of the opposite side. It is undoubtedly owing to these connections that when, in the human subject, the eyes are directed in their proper axes, the two retinae, as well as the two optic tubercles, act as a single organ. Vision is single, therefore, though there are two images upon the retinae. Double vision occurs only when the eyeballs are turned out of their proper direc- tion, so that the parallelism of their axes is lost, and the image no longer falls upon corresponding parts of the two retinae. TUBER ANNULARE. The collection of gray matter imbedded in the deeper portions of the tuber annulare occupies a situation near the central part of the brain, and lies directly in the course of the ascending fibres of the anterior and posterior columns of the cord. This ganglion is immediately connected with the functions of sensation and volun- tary motion. We have already seen that these functions are not destroyed by taking away the cerebrum, and that they also remain after removal of the cerebellum. According to the experiments of Longet, even after complete removal of the olfactory ganglia, the cerebrum, cerebellum, optic tubercles, corpora striata and optic thalami, and when nothing remains in the cavity of the cranium but the tuber annulare and the medulla oblongata, the animal is still sensitive to external impressions, and will still endeavor by volun- tary movements to escape from a painful irritation. The same observer has found, however, that as soon as the ganglion of the tuber annulare is broken up, all manifestations of sensation and volition cease, and even consciousness no longer appears to exist. The only movements which then follow external irritation are the occasional convulsive motions which are due to reflex action of the spinal cord, and which may be readily distinguished from those of a 378 THE BRAIN. voluntary character. The animal, under these circumstances, is to all appearances reduced to the condition of a dead body, except for the movements of respiration and circulation, which still go on for a certain time. The tuber annulare must therefore be regarded as the ganglion by which impressions, conveyed inward through the nerves, are first converted into conscious sensations; and in which the voluntary impulses originate, which stimulate the muscles to contraction. We must carefully distinguish, however, in this respect, a simple sensation from the ideas to which it gives origin in the mind, and the mere act of volition from the train of thought which leads to it. Both these purely mental operations take place, as we have seen, in the cerebrum ; for mere sensation and volition may exist independently of any intellectual action, as they may exist after the cerebrum has been destroyed. A sensation may be felt, for example, without our having the power of thoroughly appreciating it, or of referring it to its proper source. This condition is often experienced in a state of deep sleep, when, the body being exposed to cold, or accidentally placed in a constrained position, we feel a sense of suffering, without being able to understand its cause. We may even, under such circumstances, execute voluntary movements to escape the cause of annoyance ; but these movements, not being directed by any active intelligence, fail of accomplishing their ob- ject. We therefore remain in a state of discomfort until, on awak- ening, the activity of the reason and judgment is restored, when the offending cause is at once removed. We distinguish, then, between the simple power of sensation, and the power of fully appreciating a sensitive impression and of drawing a conclusion from it. We distinguish also between the intellectual process which leads us to decide upon a voluntary movement, and the act of volition itself. The former must precede, the latter must follow. The former takes place, so far as experi- ment can show, in tlie cerebral hemispheres ; the latter, in the gan- glion of the tuber annulare. MEDULLA OBLONGATA. The last remaining ganglion of the encephalon is that of the medulla oblongata. This ganglion, it will be remembered, is imbedded in the substance of the restiform body, occupying the MEDULLA OBLONGATA. 379 lateral and posterior portions of the medulla, at the point of origin of the pneumogastric nerves. This portion of the brain has long been known to be particularly essential to the preservation of life ; so that it has received the name of the " vital point," or the " vital knot." All the other parts of the brain may be injured or removed, as we have already seen, without the immediate and ne- cessary destruction of life ; but so soon as the medulla oblongata is broken up, and its ganglion destroyed, respiration ceases instanta- neously, and the circulation also soon comes to an end. Removal of the medulla oblongata produces, therefore, as its immediate and direct result, a stoppage of respiration; and death takes place prin- cipally as a consequence of this fact. Flourens and Longet have determined, with considerable accu- racy, the precise limits of this vital spot in the medulla oblongata. Flourens ascertained that in rabbits it extended from just above the origin of the pneumogastric nerve, to a level situated three lines and a half below this origin. In larger animals, its extent is pro- portionately increased. Longet ascertained, furthermore, that the properties of the medulla were not the same throughout its entire thickness ; but that its posterior and anterior parts might be de- stroyed with comparative impunity, the peculiarly vital spot being confined to the intermediate portions. This vital point accordingly is situated in the layer of gray matter, imbedded in the thickness of the restiform bodies, which has been previously spoken of as giving origin to the pneumogastric nerves. The precise nature of the connection between this ganglion and the function of respiration may be described as follows. The movements of respiration, which follow each other with incessant regularity through the whole period of life, are not voluntary movements. We may, to a certain extent, hasten or retard them at will, but our power over them, even in this respect, is extremely limited; and in point of fact they are performed, during the greater part of the time, in a perfectly quiet and regular manner, without our volition and even without our consciousness. They continue uninterrupted through the deepest slumber, and even in a condition of insensibility from accident or disease. These movements are the result of a reflex action taking place through the medulla oblongata. The impression which gives rise to them originates principally in the lungs, from the accumulation of carbonic acid in the pulmonary vessels and air-cells, is trans- mitted by the pneumogastric nerves to the medulla, and is thence 380 THE BRAIISr. reflected back along the motor nerves to the respiratory muscles. These muscles are then called into action, producing an expansion of the chest. The impression so conveyed to the medulla is usually unperceived by the consciousness. It is generally converted directly into a motor impulse, without attracting our attention or giving rise to any conscious sensation. Respiration, accordingly, goes on perfectly well without our interference and without our knowledge. The nervous impression, however, conveyed to the medulla, though usually imperceptible, may be made evident at any time by volun- tarily suspending the respiration. As the carbonic acid begins to accumulate in the blood and in the lungs, a peculiar sensation makes itself felt, which grows stronger and stronger with every moment, and impels us to recommence the movements of inspiration. This peculiar sensation, entirely different in character from any other, is desio'nated by the French under the name of " besoin de respirer." It becomes more urgent and distressing, the longer respiration is suspended, until finally the impulse to expand the chest can no longer be resisted by any effort of the will. During ordinary respiration, therefore, each inspiratory move- ment is excited by the partial vitiation of the air contained in the lungs. As soon as a new supply has been inhaled, the impulse to respire is satisfied, the muscles relax, and the chest collapses. In a few seconds the previous condition recurs and the same move- ments are repeated, producing in this way a regular alternation of inspirations and expirations. Since the movements of respiration are performed partly by the diaphragm and partly by the intercostal muscles, they will be differently modified by injuries of the nervous system, according to the spot at which the injury is inflicted. If the spinal cord, for example, be divided or compressed in the lower part of the neck, all the intercostal muscles will be necessarily paralyzed, and respi- ration will then be performed entirely by the diaphragm. The chest in these cases remaining motionless, and the abdomen alone rising and falling with the movements of the diaphragm, such respiration is called " abdominal" or " diaphragmatic" respiration. It is a common symptom of fracture of the spine in the lower cervical region. If the phrenic nerve, on the other hand, be divided, the diaphragm will be paralyzed, and respiration will then be performed altogether by the rising and falling of the ribs. It is then called "thoracic" or "costal" respiration. If the injury inflicted upon the spinal cord be above the origin of the second MEDULLA OBLONGATA. 381 and third cervical nerves, both the phrenic and intercostal nerves are at once paralyzed, and death necessarily takes place from suf; focation. The attempt at respiration, however, still continues in these cases, showing itself by ineffectaal inspiratory movements of the mouth and nostrils. Finally, if the medulla itself be broken up by a steel instrument introduced through the foramen magnum, so as to destroy the nervous centre in which the above reflex action takes place, both the power and the desire to breathe are at once taken away. No attempt is made at inspiration, there is no struggle, and no appearance of suffering. The animal dies simply by a want of aeration of the blood, which leads in a few moments to an arrest of the circulation. It is owing to the above action of the medulla oblongata that in- juries of this part are so promptly and constantly fatal. When the " neck is broken," as in hanging or by sudden falls upon the head, a rupture takes place of the transverse ligament of the atlas; the head, together with the first cervical vertebra, is allowed to slide forward, and the medulla is compressed between the odontoid process of the axis in front and the posterior part of the arch of the atlas behind. In cases of apoplexy, where any part of the hemispheres, corpora striata, or optic thalami, is the seat of the hemorrhage, the patient generally lives at least twelve hours ; but if the hemorrhage take place into the medulla itself, or at the base of the brain in its imme- diate neighborhood, so as to compress its substance, death follows instantaneously, and by the same mechanism as where the medulla is intentionally destroyed. An irregularity or want of correspondence in the movements of respiration is accordingly found to be one of the most threatening of all symptoms in affections of the brain. A disturbance or sus- pension of the intellectual powers does not indicate necessarily any immediate danger to life. Even sensation and volition may be im- paired without serious and direct injury to the organic functions. These symptoms only indicate the threatening progress of the dis- ease, and show that it is gradually approaching the vital centre. It is common to see, however, as the medulla itself begins to be impli- cated, a paralysis first showing itself in the respiratory movements of the nostrils and lips, while those of the chest and abdomen still go on as usual. The cheeks are then drawn in with every inspira- tion and puffed out sluggishly with every expiration, the nostrils themselves sometimes participating in these unnatural movements. A still more threatening symptom, and one which frequently pre- 382 THE BEAIN. cedes death, is an irregular, hesitating respiration, which sometimes attracts the attention of the physician, even before the remaining cerebral functions are seriously impaired. These phenomena de- pend on the connection between the respiratory movements and the reflex action of the medulla oblongata. We have now, in studying the functions of various parts of the cerebro-spinal system, become familiar with three difl'erent kinds of reflex action. The first is that of the spinal cord. Here, there is no proper sensation and no direct consciousness of the act which is performed. It is simply a nervous impression, coming from the integument, and transformed by the gray matter of the spinal cord into a motor impulse destined for the muscles. This action will take place after the removal of the hemispheres and the abolition of consciousness, as well as in the ordinary condition. The respiratory action of the medulla oblongata is of the same general character ; that is, it is not necessarily connected with either volition or consciousness. The only peculiarity in this instance is that the original nervous impression is of a special character, and its influence is finally exerted upon a special muscular apparatus. Actions of this nature are termed, par excellence, reflex actions. The second kind of reflex action takes place in the tuber annu- lare. Here the nervous impression, which is conveyed inward from the integument, instead of stopping,at the spinal cord, passes onward to the tuber annulare, where it first gives rise to a con- scious sensation ; and this sensation is immediately followed by a voluntary act. Thus, if a crumb of bread fall into the larynx, the sensation produced by it excites the movement of coughing. The sensations of hunger and thirst excite a desire for food and drink. The sexual impulse acts in precisely the same manner ; the percep- tion of particular objects giving rise immediately to special desires of a sexual character. It will be observed, in these instances, that in the first place, the nervous sensation must be actually perceived, in order to pro- duce its effect; and in the second place that the action which follows is wholly voluntary in character. But the most important peculiarity, in this respect, is that the voluntary impulse follows directly upon the receipt of the sensation. There is no intermediate reasoning or intellectual process. We do not cough because we know that this is the most effectual way to clear the larynx ; but simply because we are impelled to do so by the sensation which is MEDULLA OBLONGATA. 383 felt at the time. We do not take food or drink because we know- that they are necessary to support life, much less because we under- stand the mode in which they accomplish this object ; but merely because we desire them whenever we feel the sensations of hunger and thirst. All actions of this nature are termed insiinctive. They are volun- tary in character, but are performed blindly ; that is, without any idea of the ultimate object to be accomplished by them, and simply in consequence of the receipt of a particular sensation. Accord- ingly experience, judgment, and adaptation have nothing to do with these actions. Thus the bee builds his cell on the plan of a mathe- matical figure, without performing any mathematical calculation. The silkworm wraps himself in a cocoon of his own spinning, certainly without knowing that it is to afford him shelter during the period of his metamorphosis. The fowl incubates her eggs and keeps them at the proper temperature for development, simply because the sight of them creates in her a desire to do so. The habits of these animals, it is true, are so arranged by nature, that such instinctive actions are always calculated to accomplish an ultimate object. But this calculation is not made by the animal himself, and does not form any part of his mental operations. There is consequently no improvement in the mode of performing such actions, and but little deviation under a variety of circum- stances. The third kind of reflex action requires the co-operation of the hemispheres. Here, the nervous impression is not only conveyed to the tuber annulare and converted into a sensation, but, still following upward the course of the fibres to the cerebrum, it there gives rise to a special train of ideas. We understand then the external source of the sensation, the manner in which it is calcu- lated to affect us, and how by our actions we may turn it to our advantage or otherwise. The action which follows, therefore, in these cases, is not simply voluntary, but reasonable. It does not depend directly upon the external sensation, but upon an intellec- tual process which intervenes between the sensation and the voli- tion. These actions are distinguished, accordingly, by a character of definite contrivance, and a conscious adaptation of means to ends ; characteristics which do not belong to any other operations of the nervous system. The possession of this kind of intelligence and reasoning power is not confined to the human species. We have already seen that 384: THE BRAIN. there are many purely instinctive actions in man, as well as in animals. It is no less true that in the higher animals there is often the same exercise of reasoning power as in man. The degree of this power is much less in them than in him, but its nature is the same. Whenever, in an animal, we see any action performed with the evident intention of accomplishing a particular object, to which it is properly adapted, such an act is plainly the result of reason- ing powers, not essentially different from our own. The establish- ment of sentinels by gregarious animals, to warn the herd of the approach of danger, the recollection of punishment inflicted for a particular action, and the subsequent avoidance or concealment of that action, the teachability of many animals, and their capacity of forming new habits or of improving the old ones, are all instances of the same kind of intellectual power, and are quite different from instinct, strictly speaking. It is this faculty which especially pre- dominates over the others in the higher classes of animals, and which finally attains its maximum of development in the human species. CRANIAL NERVES. 385 CHAPTER V. THE CRANIAL NERVES. In examining the cranial nerves, we shall find that although they at first seem quite different in their distribution and properties from the spinal nerves, yet they are in reality arranged for the most part on the same plan, and may be studied in a similar manner. At the outset, however, we find that there are three^ of the cranial nerves, commonly so called, which must be arranged in a class by themselves ; since they have no character in common with the other nerves originating either from the brain or the spinal cord. These are the three nerves of special sense; viz., the Olfactory, Optic, and Auditory. They are, properly speaking, not so much nerves as commissures, connecting different parts of the encephalic mass with each other. They are neither sensitive nor motor, in the ordinary meaning of these terms; but are capable of conveying only the special sensation characteristic of the organ with which they are connected. Olfactory Nerves. — We have already described the so-called olfactory nerves as being in reality commissures, connecting the olfactory ganglia with the central parts of the brain. The masses situated upon the cribriform plate of the ethmoid bone are com- posed of gray matter; and even the filaments which they send outward to be distributed in the Schneiderian mucous membrane, are gray and gelatinous in their texture, and quite different from the fibres of ordinary nerves. The olfactory nerves are not very well adapted for direct experiment. It is, however, at least certain with regard to them that they serve to convey the special sensation of smell; that their mechanical irritation does not give rise to either pain or convulsions; and that finally their destruction, together with that of the olfactory ganglia, does not occasion any paralysis nor loss of ordinary sensibility. 25 386 THE CEANIAL NERVES. Optic Nerves. — We have already given some account of these nerves and their decussations, in connection with the history of the tubercula quadrigemina. They consist of rounded bundles of white fibres, running between the tubercles and the retinae. As the retinae themselves are membranous expansions consisting mostly of vesicular or cellular nervous matter, the optic nerves, or "tracts," must be regarded as commissures connecting the retinee with the tubercles. We have also seen that they serve, by some of their fibres, to connect the two retinas with each other, as well as the two tubercles with each other. The optic nerves convey only the special impression of light from without inward, and give origin to the reflex action of the optic tubercles, by which the pupil is made to contract. According to Lono-et, the optic nerves are absolutely insensible to pain throughout their entire length. When a galvanic current is passed through the eyeball, or when the retina is touched in operations upon the eye, the irritation has been found to produce the impression of luminous sparks and flashes, instead of an ordinary painful sensation. The impression of colored rings or spots may be easily produced by compressing the eye in particular directions; and a sudden stroke upon the eyeball will often give rise to an apparent dis- charge of brilliant sparks. Division of the optic nerves produces complete blindness, but does not destroy ordinary sensibility in any part of the eye, nor occasion any muscular paralysis. Auditory Nerves. — The nervous expansion in the cavity of the internal ear contains, like the retina, vesicles or cells as well as fibres ; and the auditory nerves are therefore to be regarded, like the optic and olfactory, as commissural in their character. They are also, like the preceding, destitute of ordinary sensibility. Ac- cording to Longet, they may be injured or destroyed without giving rise to any sensation of pain. They serve to convey to the brain the special sensation of sound, and seem incapable of transmitting any other. Longet' relates an experiment performed by Volta in which, by passing a galvanic current through the ears, the observer experienced the sensation of an interrupted hissing noise, so long as th.e connection of the wires was maintained. Inflammations within the ear, or in its neighborhood, are often accompanied by the perception of various noises, like the ringing of bells, the ' Traite de Physiologic, vol. ii. p. 286. THE CRANIAL NERVES. o»7 washing of the waves, the humming of insects; sounds which have no external existence, but which are simulated by the morbid irri- tation of the auditory nerve. It is evident from the facts detailed above that the nerves of special sense are neither motor nor sensitive, properly speaking; and that they are distinct in their nature from the ordinary spinal nerves. The remainder of the cranial nerves, however, have no such essential peculiarities. Some of them are exclusively motor in character, others exclusively sensitive; while most of them exhibit the two properties to a certain extent, as mixed nerves. They may be conveniently arranged in three pairs, according to the regions in which they are distributed, corresponding very closely with the motor and sensitive roots of the spinal nerves. According to such a plan, the arrangement of the cranial nerves would be as follows : — Ck vxiAL Nerves. Nerves of Special Sense. 1. Olfactory. 2. Optic. 3. Auditory. Motor Nerves. Distributed to Seasitive Nerves. Motor oculi com. ■ Patheticus Face. Large root of 5 th 1st PAIR. • Motor oc. externus Small root of 5th pai r L Facial , 2d PAIR. Sublingual Tongue. Glosso-pharyngeal 3d PAIR. Spinal accessory Neck, &c. Pneumogastric. The above arrangement of the cranial nerves is not absolutely perfect in all its details. Thus, while the sublingual supplies the muscles of the tongue alone, the glosso-pharyngeal sends part of its sensitive fibres to the tongue and part to the pharynx; and while the large root of the 5th pair is mostly distributed in the face, one of its branches, viz., the gustatory, is distributed to the tongue. Notwithstanding these irregularities, however, the above division of the cranial nerves is in the main correct, and will be found extremely useful as an assistant in the study of their func- tions. There is no impropriety, moreover, in regarding all the motor branches distributed upon the face as one nerve; since even the anterior roots of the spinal nerves originate from the spinal cord, each by several distinct filaments, which are associated into a single bundle only at a certain distance from their point of origin. The 388 THE CRANIAL NERVES. mere fact that two nerves leave the cavity of the cranium by the same foramen does not indicate that they have the same or even a similar function. Thus the facial and auditory both escape from the cavity of the cranium by the foramen auditorium internum, and yet we do not hesitate to regard them as entirely distinct in their nature and functions. It is the ultimate distribution of a nerve, and not its course through the bones of the skull, that indicates its physiological character and position. For while the ultimate distribution of any particular nerve is always the same, its arrange- ment as to trunks and branches may vary, in different species of animals, with the anatomical arrangement of the bones of the skull. This is well illustrated by a fact first pointed out by Dr. Jeffries Wyman^ in the anatomy of the nervous system of the bullfrog. In this animal, both the facial nerve and motor oculi externus, instead of arising as distinct nerves, are actually given off as branches of the 5th pair ; while their ultimate distribution is the same as in other animals. All the motor and sensitive nerves distributed to the face are accordingly to be regarded as so many different branches of the same trunk; varying sometimes in their course, but always the same in their ultimate distribution. MOTOR NERVES. The motor nerves of the head are in all respects identical in their properties with the anterior roots of the spinal nerves. For, in the first place, they are distributed to muscles, and not to the inteo-ument or to mucous membranes; secondly, their division causes muscular paralysis; and thirdly, mechanical irritation ap- plied at their origin produces muscular contraction in the parts to which they are distributed, but does not give rise to a painful sensation. In several instances, nevertheless, the motor nerves, though insensible at their origin, show a certain degree of sensibi- lity when irritated after their exit from the skull, owing to fibres of communication which they receive from the purely sensitive nerves. In this respect they resemble the spinal nerves, the motor and sensitive filaments of which are at first distinct in the anterior and posterior roots, but afterward mingle with each other, on leaving the cavity of the spinal canal. ' Nervous System of Rana pipiens ; published by the Smithsonian Institution. Washington, 1853. MOTOR CRANIAL NERVES. 389 Motor OcuLi Communis. — This nerve, which is sometimes known bj the more convenient name of the oculo-molorius, originates from the inner edge of the eras cerebri, passes into the cavity of the orbit by the sphenoidal fissure, and is distributed to the levator palpebra3 superioris, and to all the muscles moving the eyeball, except the external rectus and the superior oblique. Its irritation accordingly produces convulsive movements in these parts, and its division has the effect of paralyzing the muscles to which it is distributed. The superior eyelid falls down over the pupil, and cannot be raised, owing to the inaction of its levator muscle, so that the eye appears constantly half shut. This condition is known by the name of " ptosis." The movements of the eyeball are also nearly suspended, and permanent external strabismus takes place, owing to the paralysis of the internal rectus muscle, while the ex- ternal rectus, animated by a different nerve, preserves its activity. Patheticus. — This nerve, which supplies the superior oblique muscle of the eyeball, is similar in its general properties to the pre- ceding. Its section causes paralysis of the above muscle, without any loss of sensibility. Motor Externus. — This nerve, the sixth pair, according to the usual anatomical arrangement, is distributed to the external rectus muscle of the eyeball. Its division or injury by disease is followed by internal strabismus, owing to the unopposed action of the internal rectus muscle. Small Root of 5th Pair. — It will be remembered that the 5th pair of nerves arises by two roots, which run in close proximity to each other, as far as the level of the Casserian ganglion. The fibres of the smaller root, however, do not mingle at all with the substance of the ganglion, but pass underneath it, as a distinct bun- dle, and emerge afterward from the skull by the foramen ovale of the sphenoid bone, as a portion of the inferior maxillary branch of the 5th pair. While the remainder of the 5th pair is distributed to the integument and mucous membranes about the face, all the fibres derived from this smaller root are sent to the muscles con- cerned in mastication, viz., the great temporal, the masseter, the internal and external pterygoids, the digastric, and the mylo-hyoid. It is therefore sometimes known as the " masticator" nerve. It is exclusively the motor nerve of these muscles ; for while galvaniza- tion of the large root of the 5th pair, according to Longet, pro- 390 THE CRANIAL NERVES. duces no convulsive movement, but only a painful sensation, if the irritation be applied to the small root alone, the above muscles are immediately thrown into contraction, and the jaws violently brought together. Section of this nerve paralyzes the muscles of mastica- tion, without affecting the other muscles of the face. Facial. — This nerve was known to the older anatomists as the " portio dura of the seventh pair." Tt leaves the cavity of the cranium by the internal auditory foramen, in company with the auditory nerve; and as the latter is of a softer consistency than the facial, they have received the names respectively of the "portio mollis" and " portio dura" of the seventh pair. There is, however, no physiological connection between these two nerves; for while the auditory is spread out in the cavity of the internal ear, the facial passes onward through the petrous portion of the temporal bone, emerges at the stylo mastoid foramen, bends round beneath the external ear, and passes forward through the substance of the parotid gland, forming a plexus, called the " pes anserinus," by the abundant inosculation of its branches. Tt then sends its filaments forward in a diverging course, and is finally distributed to the superficial muscles of the face; those, namely, which are concerned in the production of expression. (Fig. 146.) The facial, consequently, is the motor nerve of the face. It has no- thing to do with transmitting sen- sitive impressions, since it has been frequently shown that after section of the 5th pair, the facial remaining entire, the sensibility of the face is completely lost; so that the integu- ment may be cut, pricked, burned, or lacerated, without any sign of pain being exhibited by the ani- mal. The facial, therefore, does not transmit sensation from these parts ; and its division, which was formerly resorted to in cases of tic doulou- reux, is accordingly altogether in- capable of relieving neuralgic pains. This nerve is, however, directly connected with muscular action. Mechanical or galvanic irritation of its fibres produces convulsive Fig. 146. Facial Nerve. MOTOR CRANIAL NERVES. 391 twitchiijgs in the nostrils, lips, cheeks, &c. Section of the facial on one side, its destruction by disease, or compression by a tumor, induces the extremely characteristic affection known as "facial paralysis." The affected side of the face in these cases, up to the median line, loses altogether its power of motion, and at the same time its natural expression. The corner of the mouth falls down- ward, and the whole lower part of the face is drawn over to the opposite side by the force of the antagonistic muscles. The lips are unable to retain the fluids of the mouth, and the saliva dribbles away from between them, giving to the face a remarkably vacant and helpless appearance. The lower eyelid also sinks downward from paralysis of the orbicularis muscle, and the eye cannot be completely closed. It will be observed that precisely opposite effects are produced upon the eyelids by paralysis of the oculo-motorius and of the facial. In the former instance, owing to paralysis of the levator palpebrse superioris, the eye is always partially closed ; in the latter, owing to paralysis of the orbicularis, it is always par- tially open. Though the facial, however, be essentially a motor nerve, yet its principal branches distributed to the face have a certain degree of sensibility ; that is, when irritated in the middle of their course, the animal immediately gives evidence of a painful sensation. Longet has shown, by an extremely ingenious mode of experiment,^ that this sensibility of the branches of the facial does not depend on any sensitive fibres of its own, but upon those which it derives from inosculation with the fifth pair. He exposes, for example, the facial nerve in the dog, and irritating its principal branches one after the other, at each application of the irritant there are evident signs of pain. He then divides the facial nerve at its point of exit from the stylo-mastoid foramen, and finds that, after this operation, the sensibility of its branches still remains. The fibres, accordingly, upon which this sensibility depends, do not pass out with the trunk of the nerve, but are derived from some other source. The experi- menter, then, upon another animal, divides the 5th pair within the skull, leaving the facial untouched ; and afterward, on irritating as before the exposed branches of the latter nerve, he finds that its sensibility has entirely disappeared. It is by filaments, accordingly, derived from the 5th pair, that a certain degree of sensibility is communicated to the branches of the facial. ' Traite de Physiologie, vol. ii. pp. 3.54-3.57. 392 THE CRANIAL NERVES. These facts account for the peculiar circumstance that, in cases of tic douloureux, the spasmodic pain sometimes follows exactly the course of the facial nerve, viz : from behind the ear forward upon the side of the face ; and yet the section of this nerve does not put an end to the neuralgia, but only causes paralysis of the facial muscles. Sublingual. — The sublingual nerve originates from the anterior and lateral portions of the medulla oblongata, and passing out by the anterior condyloid foramen, is distributed exclusively to the muscles of the tongue. Irritation of its fibres in any part of their course produces convulsive twitching in this organ. Its section paralyzes completely the movements of the tongue, without affecting directly the sensibility of its mucous membrane. If irritated at its origin, the sublingual nerve, according to the experiments of Longet, is entirely insensible ; but if the irritation be applied in the middle of its course, signs of pain are immediately manifested. Its sensi- bility, like that of the facial, is consequently derived from its inos- culating with other sensitive nerves after its emergence from the skull. Spinal Accessory. — This nerve originates by many filaments from the side of the medulla oblongata below the level of the pneu- mogastric, and also from the lateral portion of the spinal cord, between the anterior and posterior roots of the upper five or six cervical nerves. Its fibres pass upward, enter the cavity of the cra- nium by the foramen magnum, and again emerge from it by the posterior foramen lacerum, in company with the jugular vein and the glosso-pharyngeal and pneumogastric nerves, to the latter of which it gives off an important branch of communication. It is finally distributed to the sterno-mastoid and trapezius muscles. The above muscles, however, are also supplied by branches from the cervical and dorsal nerves ; and consequently, it has been found that division of the spinal accessory is not followed by their com- plete paralysis ; but only by a certain debility, owing to their having lost a part of their motor force. The sterno-mastoid and trapezius serve as accessory muscles of respiration, and come into play when the respiratory movements are unusually hurried or laborious. The spinal accessory was regarded by Sir Charles Bell as especially devoted to this function in the above muscles, and was, therefore^ called, by him, the "superior respiratory nerve." SENSITIVE CRANIAL NERVES. 393 The spinal accessory has been found by Bernard to be insensible at its origin, like the anterior roots of the spinal nerves; but if irri- tated after its exit from the skull, it gives signs of sensibility. This may be attributed to its receiving filaments of inosculation from the anterior branches of the first and second cervical nerves. The rea- son for the above anatomical fact, viz., that motor nerves are sup- plied daring their course with sensitive fibres by inosculation, becomes evident when we reflect that the muscles themselves pos- sess a certain degree of sensibility, though less than that which belongs to the integument and to some parts of the mucous mem- branes. This sensibility of the muscles is undoubtedly essential to the perfect performance of their function ; and, as the motor nerves are incapable by themselves of transmitting sensitive im- pressions, they are joined, soon after their origin, by other filaments which communicate to them the necessary power. SENSITIVE NERVES. The three sensitive nerves originating from the brain are the large root of the 5th pair, the glosso-pharyngeal, and the pneumo- gastric. It will be observed that, in all their essential properties, they correspond with the posterior roots of the spinal nerves. Like them they are inexcitable, but extremely sensitive. Irritated at their point of origin, they give rise to acutely painful sensations, but to no convulsive movements. Secondly, if divided at the same situation, the operation is followed by loss of sensibility in the parts to which they are distributed, without any disturbance of the motive power. Each of these nerves, furthermore, like the poste- rior root of a spinal nerve, is provided with a ganglion through which its fibres pass : the 5th pair, with the Casserian ganglion, situated near the inner extremity of the petrous portion of the tem- poral bone ; the glosso-pharyngeal, with the ganglion of Andersch, situated in the jugular fossa; while the pneumogastric presents, just before its passage through the jugular foramen, a ganglion known as the ganglion of the pneumogastric nerve. Finally, the sensitive fibres of all these nerves, beyond the situation of their ganglia, are intermingled with others of a motor origin. The large root of the 5th pair, wbich is exclusively sensitive, is accompanied by the fibres of the small root, which are exclusively motor. The glosso-pharyngeal receives motor filaments from the facial and spinal 394 THE CRANIAL NERVES. accessory, becoming consequently a mixed nerve outside the cra- nial cavity; while the pneumogastric is joined by fibres from the spinal accessory and various other nerves of a motor character. These nerves, accordingly, are exclusively sensitive only at their point of origin. Though they afterward retain the predominating character of sensitive nerves, they are yet found, if irritated in the middle of their course, to be intermingled with motor fibres, and to have consequently acquired, to a certain extent, the character of mixed nerves. The resemblance, therefore, between the cranial and spinal nerves is complete. Fifth Pair. — This is one of the most important and remarkable in its properties of all the cranial nerves. It is the sensitive nerve of the face, and of the adjoining mucous membranes. We have already described the course and distribution of the small root of this nerve. The large root, after emerging from the outer and under surface of the pons Varolii, passes forward over the inner extremity of the petrous portion of the temporal bone. It there expands into a crescentic-shaped swelling, containing a quantity of gray matter with which its fibres are intermingled, and which is known as the Casserian ganglion. The fibres of the smaller root, as already remarked, do not take any part in the formation of this ganglion, but may be seen passing beneath it as a distinct bundle, and continuing their course forward to the foramen ovale, through which they emerge from the skull. From the anterior and external border of the Casserian ganglion, the sensitive portion of the nerve emerges in three separate branches, viz., the ophthalmic, the su- perior maxillary, and the inferior maxillary. The first of these, viz., the ophthalmic, is so called because it passes through the orbit of the eye. It enters the sphenoidal fissure, and runs along the upper portion of the orbit, sending branches to the ophthalmic ganglion of the sympathetic, to the lachrymal gland, the conjunc- tiva, and the mucous membrane of the lachrymal sac. It also sends off a small branch (nasal branch) which penetrates into the nasal passages and supplies the Schneiderian mucous membrane. It then emerges upon the face by the supra-orbital foramen, and is dis- tributed to the integument of the forehead and side of the head as far back as the vertex. The second branch of this nerve, or the superior maxillary, passes out by the foramen rotund um, and runs along the longitudinal canal SENSITIVE NERVES. 395 Fis. 147. Distribution of Fifth Nerve UPON T H !•; F A c R . — n. Casserian ganglion. 1. Ophtlialmic branch. 2. Superior maxil- lary branch. 3. Inferior maxillary branch. in the floor of the orbit, giving oft' branches during its passage to the teeth of the upper jaw and to the mucous membrane of the antrum maxiHare. It finally emerges upon the middle of the face by the infra-orbital foramen, and is distributed to the lower eye- lid, the nose, the cheek and the upper lip. The third, or inferior maxillary branch of the fifth pair, leaves the cavity of the cranium by the fora- men ovale. It first sends off" a few branches to the integument of the temple and external ear, then a large and important branch, vi%., the "gustatory" or "lingual" branch, which is distributed to the mucous membrane of the anterior two-thirds of the tongue. The main trunk then enters the inferior dental canal, sends nerves to the teeth of the lower jaw, emerges at the mental foramen, and is finally distributed to the integument of the chin, lower lip, and inferior part of the face. This nerve is accordingly distributed to the sensitive surfaces, that is, the integument and mucous membranes, about the face. A few of its fibres are sent also to the muscles of the face ; but these fibres are sensitive in their character, and serve merely to impart to the muscles a certain degree of sensibility. It has been ascertained by Longet that if the ganglionic portion of this nerve be irritated by a galvanic current, no convulsive movements whatever are produced, even in those muscles which are supplied with filaments from its infra-orbital and mental branches; but if its smaller or non-ganglionic root be irritated in the same way, contractions instantly follow in the muscles of mastication. Irritation of the 5th pair, in any part of its course, as well as of its larger root behind the Casserian ganglion, produces intense pain. Its division is followed by complete loss of sensibility in the in- tegument of the face, the lips, the conjunctiva, and mucous mem- brane of the nares and mouth. The sense of taste is also destroyed throughout the anterior two-thirds of the tongue, owing to the paralysis of the lingual or gustatory nerve. The skin of the face 396 THE CRANIAL XERVES. may then be pricked, burned, cut or lacerated in any way, without producing pain, and even apparently without the knowledge of the animal upon whom the operation is performed. The 5th pair, beside supplying the sensibility of the integument of the face, has a peculiar and important influence on the organs of special sense. This influence appears to consist in some connec- tion between the action of the 5th pair and the processes of nutrition; so that when the former is injured, the latter become immediately deranged. For the perfect action of any one of the organs of special sense, two conditions are necessary : first, the sensibility of the special nerve belonging to it, and, secondly, the integrity of the component parts of the organ itself. Xow as the nutrition of the organ is, to a certain extent, under the control of the 5th pair, any serious injury to this nerve produces a derangement in the tissues of the organ, and consequently interferes with the due performance of its function. The mucous membrane of the nasal passages, for example, is supplied by two different nerves; first, the olfactory, distributed throughout its upper portion, by which it is endowed with the special sense of smell ; and, secondly, the nasal branch of the 5th pair, distributed throughout its middle and lower portions, by which it is supplied with ordinary sensibility. Although the Schneiderian mucous membrane, therefore, after destruction of the olfactory nerve, loses altogether its power of distinguisMng odors, properly so called, such as the odor of flowers, of turpentine, of sulphuretted hydrogen, and the like, it still remains sensitive to the action of vapors which are merely irritating in their character, such as those of ammonia and acetic acid. These substances act by exciting the ordinary sensibility of the mucous membrane, which is supplied by fibres of the 5th pair. They will also act in a similar manner upon the integument, or upon other mucous membranes endowed with general sensibility ; while true odors are perceptible by the olfactory organ alone. The 5th pair accordingly supplies general sensibilit}' to the nasal passages, and this property will remain after the special sense of smell has been destroyed. If, however, the 5th pair itself be divided, not only is general sensibility destroyed in the Schneider- ian mucous membrane, but, according to the experiments of Longet, a disturbance begins to take place in the nutrition of its tissue, by which it is gradually rendered unfit for the performance of its special function, and the power of smell is finally lost. The SENSITIVE CRANIAL NERVES. 397 mucous membrane, under these circumstances, becomes injected and swollen, assumes a fungous consistency, and is liable to bleed at the slightest touch. The effect of this alteration is to blunt or altogether destroy the sense of smell. It is owing to a similar unnatural condition of the mucous membrane that the power of smell is always more or less impaired in cases of coryza and influenza. The olfactory nerves become inactive in consequence of the morbid alteration in their mucous membrane, and in the secre- tions which cover it. The influence of this nerve over the organ of vision is still more remarkable. It has been known for many years that division of the 5th pair within the cranium, or of its ophthalmic branch, is fol- lowed by an inflammation of the corresponding eye which usually goes on to complete and permanent destruction of the organ. Immediately after the operation, the pupil becomes contracted and the conjunctiva loses its sensibility. At the end of twenty-four hours, the cornea begins to become opaline, and, by the second day, the conjunctiva is already inflamed and begins to discharge a purulent secretion. The inflammation, after commencing in the conjunctiva, increases in intensity and soon spreads to the iris, which becomes covered with a layer of inflammatory exudation. The cornea grows constantly more opaque, until it is at last altogether impermeable to light, and vision is consequently sus- pended. Blindness, therefore, does not result in these instances from any direct affection of the optic nerve or of the retina, but is owing simply to opacity of the cornea. Sometimes the diseased action goes on until it results in ulceration of the cornea and dis- charge of the humors of the eye ; sometimes, after the lapse of several days, the inflammatory appearances subside, and the eye is finally restored to its natural condition. It has been observed, however, that, although the above conse- quences always follow division of the 5th pair when performed at the level of the Casserian ganglion, or between it and the eyeball, they are either much diminished in intensity or altogether wanting when the division is made at a point posterior to the ganglion. This circumstance has led to the belief that the influence of the 5th pair on the nutrition of the eyeball does not reside in its own proper fibres, but in some filaments of the sympathetic which join the 5th pair at the level of the Casserian ganglion. If the section accordingly be made at this point, or in front of it, the fibres of the sympathetic will be divided with the others, and inflammation of 398 THE CRANIAL NERVES. the eye will result; but if the section be made behind the ganglion, the fibres of the sympathetic will escape division, and the injurious effects upon the eye will be wanting. Such is the explanation usually given of the above-mentioned facts; but the question has not as yet been determined in a direct manner. Division of the 5th pair destroys also the general sensibility of the external auditory meatus, the lining membrane of which is supplied by its filaments. Inflammation of this membrane and its consequent alterations, it is well known, interfere seriously with the sense of hearing. It is no uncommon occurrence for an accumulation of cerumen to take place after inflammation of this part, so as to block up the canal and produce partial or complete deafness. It has not been ascertained, however, whether division of the 5th pair is usually followed by similar changes in this part. The lingual branch of the 5th pair supplies the anterior ex- tremity and middle portion of the tongue both with general sensi- bility and with the power of taste. The sensibility of. the tongue is accordingly provided for by two difi'erent nerves; in its anterior two-thirds, by the lingual branch of the 5th pair ; in its posterior third, by fibres of the glosso-pharyngeal. The facial branches of the 5th pair are the ordinary seat of tic douloureux. This affection is not unfrequently confined to either the supra-orbital, the infra-orbital, or the mental branch ; and the pain may be accurately traced in the direction of their diverging fibres. It has already been mentioned that the painful sensations sometimes also follow the course of the facial, owing to some sen- sitive filaments which that nerve receives from the 5th pair. Glosso-Pharyngeal. — This nerve originates from the lateral portion of the medulla oblongata, passes outward, and enters the posterior foramen lacerum in company with the pneumogastric and spinal accessory. While in the jugular fossa it presents a gangliform enlargement, called the ganglion of Andersch, below the level of which it receives branches of communication from the facial and the spinal accessory. It then runs downward and forward and is dis- tributed to the mucous membrane of the base of the tongue, pillars of the fauces, soft palate, middle ear, and upper part of the pharynx. It also sends some branches to the constrictors of the pharynx and the neighboring muscles. Longet has found this nerve at its origin to be exclusively sensitive ; but below the level of its ganglion it has been found by various observers to be both sensitive and motor. PNEUMOGASTRIC NERVE. 399 owing to the fibres of communication received from the motor nerves mentioned above. Its final distribution is, however, as we have seen, principally to sensitive surfaces. The principal office of this nerve is to impart the sense of taste to those parts of the tongue to which it is distributed. It also presides over the general sensibility of this part of the tongue, as well as of the fauces and pharynx. There are certain reflex actions, furthermore, which take place through the medium of the glosso-pharyngeal nerve. After the food has been thoroughly masticated, it is carried, by the move- ments of the tongue and sides of the mouth, through the fauces, and brought in contact with the mucous membrane of the pharynx. This produces an impression which, conveyed to the medulla oblongata by the filaments of the glosso-pharyngeal, excites the muscles of the fauces and pharynx by reflex action. The food is consequently grasped by these muscles, without the concurrence of the will, and the process of deglutition is commenced. This action is not only involuntary, but it will frequently take place even in opposition to the will. The food, once past the isthmus of the fauces, is beyond the control of volition, and cannot be returned except by convulsive action, equally involuntary in its character. Natural stimulants, therefore, applied to the mucous membrane of the pharynx, excite deglutition ; unnatural stimulants, applied to the same part, excite vomiting. If the finger be introduced into the fauces and pharynx, or if the mucous membrane of these parts be irritated by prolonged tickling with the end of a feather, the sensation of nausea, conveyed through the glosso-pharyngeal nerve, is sometimes so great as to produce immediate and copious vomit- ing. This method may often be successfully employed in cases of poisoning, when it is desirable to excite vomiting rapidly, and when emetic medicines are not at hand. Pneumogastric. — Owing to the numerous connections of the pneumogastric with other nerves, its varied and extensive distribu- tion, and the important character of its functions, this is properly regarded as one of the most remarkable nerves in the whole body. Owing to the wandering course of its fibres, which are distributed to no less than four different vital organs, viz., the heart, lungs, stomach, and liver, as well as to several other parts of secondary importance, it has been often known by the name of the jmr vagum. The pneumogastric arises, by a number of separate filaments, from the lateral portion of the medulla oblongata, in the groove between 400 THE CRANIAL NERVES. the olivary and restiform bodies. These filaments unite into a single trunk, which emerges from the cranium by the jugular fora- men, where it is provided with a longitudinal ganglionic swelling, the "ganglion of the pneuraogastric nerve." Immediately below the level of this ganglion the nerve receives an important branch of communication from the spinal acces- sory, and afterward from the facial, the sublingual, and the anterior branches of the first and second cervicals. At its origin, the pneumogastric is ex- clusively a sensitive nerve. Irritated above the situation of its ganglion, it has been found to convey painful sensations alone ; but if the irritation be applied at a lower level, it causes at the same time muscular contractions, owing to the fila- ments which it has received from the above mentioned motor nerves. It be- comes, consequently, after emerging from the cranial cavity, a mixed nerve; and has accordingly, in nearly all its branches, a double distribution, viz., to the mucous membranes and the muscular coat of the organs to which it belongs. In passing down the neck this nerve sends branches to the mucous membranes and muscular coat of the pharynx, oeso- phagus, and respiratory passages. Among the most important of these are the two laryngeal nerves, viz., the superior and inferior. The superior laryngeal nerve, which is given off" from the trunk of the pneumogastric just after it has emerged from the cavity of the skull, passes down- ward and forward, penetrates the larynx by an opening in the side of the thyro- hyoid membrane, and is distributed to the raucous membrane of the larynx and glottis, and also to a single laryngeal muscle, viz., the crico-thyroid. This branch is therefore partly muscular, but mostly sensitive in its distribution. The infe- rior laryngeal branch is given off just after the pneumogastric has Diagram of Pneumogastric N E B V E, with its principal branches. — ^1. Pharyngeal branch. 2. Supe- rior laryngeal. 3. Inferior laryn- geal. 4. Pulmonary branches. 5. Stomach. 6. Liver. PNEUMOGASTRIC. 401 entered the cavity of the chest. It curves round the subclavian artery on the right side and the arch of the aorta on the left, and ascends in the groove between the trachea and oesophagus, to the larynx. It then enters the larynx between the cricoid cartilage and the posterior edge of the thyroid, and is distributed to all the mus- cles of the larynx, with the exception of the crico-thyroid. This branch is, therefore, exclusively muscular in its distribution. The trunk of the pneumogastric, after supplying the above branches, as well as sending numerous filaments in the neck to the trachea and oesophagus, gives off in the chest its pulmonary branches, which follow the bronchial tubes in the lungs to their minutest ramifications. It then passes into the abdomen and sup- plies the muscular and mucous layers of the stomach, ramifying over both the anterior and posterior surfaces of the organ; after which its fibres spread out and are distributed to the liver, spleen, pancreas, and gall-bladder. The functions of the pneumogastric will now be successively studied in the various organs to which it is distributed. Pharynx and (Esophagus. — The reflex action of deglutition, which has already been described as commencing in the upper part of the pharynx, by means of the glosso-pharyngeal, is con- tinued in the lower portion of the pharynx and throughout the oesophagus by the pneumogastric. As the food is compressed by the superior constrictor muscle of the pharynx and forced down- ward, it excites the mucous membrane with which it is brought in contact and gives rise to another contraction of the middle constric- tor. The lower constrictor is then brought into action in its turn in a similar manner; and a wave-like or peristaltic contraction is thence propagated throughout the entire length of the oesophagus, by which the food is carried rapidly from above downward, and conducted at last to the stomach. Each successive portion of the raucous membrane, in this instance, receives in turn the stimulus of the food, and excites instantly its own muscles to contraction ; so that the food passes rapidly from one end of the oesophagus to the other by an action which is wholly reflex in character and entirely withdrawn from the control of the will. Section of the pneumogastric, or of its pharyngeal and oesophageal branches, destroys therefore at the same time the sensibility and the motive power of these parts. The food is no longer conveyed readily to the stomach, but accumulates in the paralyzed oesophagus, into which it is forced by the voluntary movements of the mouth and 26 402 THE CRANIAL NERVES, fauces, and by the coDtinued action of the upper part of the pharynx. It must be remembered that the general sensibility of the oeso- phagus is very slight, as compared with that of the integument, or even of the mucous membranes near the exterior. It is a general rule, in fact, that the sensibility of the mucous membranes is most acute at the external orifices of their canals ; as, for example, at the lips, anterior nares, anus, orifice of the urethra, &c. It diminishes constantly from without inward, and disappears altogether at a certain distance from the surface. The sensibility of the pharynx is less acute than that of the mouth, but is still sufficient to enable us to perceive the contact of ordinary substances ; while in the oesophagus we are not usually sensible of the impression of the food as it passes from above downward. The reflex action takes place here without any assistance from the consciousness ; and it is only when substances of an unusually pungent or irritating nature are mingled with the food, that its passage through the oesophagus produces a distinct sensation. Larynx. — We have already described the course and distribution of the two laryngeal branches of the pneumogastric. The superior laryngeal nerve is principally the sensitive nerve of the larynx. Its division destroys sensibility in the mucous membrane of this organ, but paralyzes only one of its muscles, viz : the crico-thyroid. Galvanization of this nerve has also been found to induce con- tractions in the crico-thyroid, but in none of the other muscles belonging to the larynx. The inferior laryngeal, on the other hand, is a motor nerve. Its division paralyzes all the muscles of the larynx except the crico-thyroid, and irritation of its divided extremity produces contraction in the same muscles. The muscles and mucous membrane of the larynx are therefore supplied by two different branches of the same trunk, viz., the superior laryngeal nerve for the mucous membrane, and the inferior laryngeal nerve for the muscles. The larynx, in man and in all the higher animals, performs a double function ; one part of which is connected with the voice, the other with respiration. The formation of the voice in the larynx takes place as follows. If the glottis be exposed in the living animal, as already described in a previous chapter (Section I., Chap. XII.), it will be seen that so long as the vocal chords preserve their usual relaxed condition during expiration, no sound is heard, except the ordinary faint PNEUMOGASTRIC. 403 whisper of the air passing gently through the cavity of the larynx. When a vocal sound, however, is to be produced, the chorus are suddenly made tense and applied closely to each other, so as to diminish very considerabl}"- the size of the orifice; and the air, driven by an unusually forcible expiration through the narrow opening of the glottis, in passing between the vibrating vocal chords, is itself thrown into vibrations which produce the sound required. The tone, pitch, and intensity of this sound, vary with the conformation o the larynx, the degree of tension and approxi- mation of the vocal chords, and the force of the expiratory effort. The narrower the opening of the glottis, and the greater the tension of the chords, under ordinary circumstances, the more acute the sound; while a wider opening and a less degree of tension produce a graver note. The quality of the sound is also modified by the length of the column of air included between the glottis and the month, the tense or relaxed condition of the walls of the pharynx and fauces, and the state of dryness or moisture of the mucous membrane lining the aerial passages. Articulation, on the other hand, or the division of the vocal sound into vowels and consonants, is accomplished entirely by the lips, tongue, teeth, and fauces. These organs, however, are under the control of other nerves, and the mechanism of their action need not occupy us here. Since the production of a vocal sound, therefore, depends upon the tension and position of the vocal chords, as determined by the action of the laryngeal muscles, it is not surprising that division of the inferior laryngeal nerves, by paraljT'zing these muscles, should produce a loss of voice. It has been sometimes found that in very young animals the crico-thyroid muscles, which are the only ones not affected by division of the inferior laryngeal nerves, are still sufficient to give some degree of tension to the vocal chords, and to produce in this way an imperfect sound; but usually the voice is entirely lost after such an operation. It is a very remarkable fact, however, in this connection, that all the motor filaments of the pneumogastric, which are concerned in the formation of the voice, are derived from a single source. It will be remembered that the pneumogastric, itself originally a sensitive nerve, receives motor filaments, on leaving the cranial cavity, from no less than five different nerves. Of these filaments, however, those coming from the spinal accessory are the only ones necessary to the production of vocal sounds. For it has been 404 THE CRANIAL NEEVES. found by Bischofif and by Bernard^ that if all the roots of the spinal accessory be divided at their origin, or if the nerve itself be torn away at its exit from the skull, all the other cranial nerves remaining untouched, the voice is lost as completely as if the inferior laryngeal itself had been destroyed. All the motor fibres of the pneumogastric, therefore, which act in the formation of the voice are derived, by inosculation, from the spinal accessory nerve. In respiration^ again, the larynx performs another and still more important function. In the first place, it stands as a sort of guard, or sentinel, at the entrance of the respiratory passages, to prevent the intrusion of foreign substances. If a crumb of bread accidentally fall within the aryteno-epiglottidean folds, or upon the edges of the vocal chords, or upon the posterior surface of the epiglottis, the sensibility of these parts immediately excites a violent expulsive cough, by which the foreign body is dislodged. The impression, received and conveyed inward by the sensitive fibres of the supe- rior laryngeal nerve, is reflected back upon the expiratory muscles of the chest and abdomen, by whicb the instinctive movements of coughing are accomplished. Touching the above parts with the point of a needle, or pinching them with the blades of a forceps, will produce the same effect. This reaction is essentially dependent on the sensibility of the laryngeal mucous membrane ; and it can no longer be produced after section of the pneumogastric nerve, or of its superior laryngeal branch. In the second place, the respiratory movements of the glottis^ already described in a previous chapter, are of the greatest importance to the preservation of life. We have seen tbat at the moment of inspiration the vocal chords are separated from each other, and the glottis opened, by the action of the posterior crico arytenoid muscles; and tbat in expiration the muscles and the vocal chords are botb relaxed, and the air allowed to pass out readily througb the glottis. The opening of the glottis in inspiration therefore is an active movement, while its partial closure or collapse in expiration is a passive one. Furthermore, the opening of the glottis in inspiration is necessary in order to afford a sufficiently wide passage for the air, in its way to the trachea, bronchi, and pulmonary vesicles. Now we have found, as Budge and Longet had previously no- ticed, that if the inferior laryngeal nerve on the right side be divided while the glottis is exposed as above, the respiratory move- ' Reclierclies Experimentales sur les fonctions du nerf spinal. Paris, 1851. PNEUMOGASTEIC. 405 ments of the right vocal chord instantly cease, owing to the para- lysis of the posterior crico-arytenoid muscle on that side. If the inferior laryngeal nerve on the left side be also divided, the para- lysis of the glottis is then complete, and its respiratory movements cease altogether. A serious difficulty in respiration is the imme- diate consequence of this operation. For the vocal chords, being no longer stretched and separated from each other at the moment of inspiration, but remaining lax and flexible, act as a double valve, and are pressed inward by the column of inspired air ; thus par- tially blocking up the passage and impeding the access of air to the lungs. If the pneumogastrics be divided in the middle of the neck, the larynx is of course paralyzed precisely as after section of the inferior laryngeal nerves, since these nerves are given ofi" only after the main trunks have entered the cavity of the chest ; and the immediate effect of either of these operations is to produce a difficulty of inspiration, accompanied by a peculiar wheezing or sucking noise, evidently produced in the larynx and dependent on the falling together of the vocal chords. In very young animals, as for example in pups of a few days old, in whom the glottis is smaller and the larynx less rigid than in adult dogs, this difficulty is much more strongly marked. Legallois' has even seen a pup of two days old almost instantly suffocated after section of the two inferior laryngeal nerves. We have found that in pups of two weeks old, division of the inferior laryngeals is followed by death at the end of from thirty to forty hours, evidently from impeded respiration. The importance, therefore, of these movements of the glottis in respiration becomes very evident. They are, in fact, part and parcel of the general respiratory movements, and are necessary to a due performance of the function. It has been found, moreover, that the motor filaments concerned in this action are not derived, like those of the voice, from a single source. While the vocal movements of the larynx are arrested, as mentioned above, by division of the spinal accessory alone, those of respiration still go on ; and in order to put a stop to the latter, either the pneumo- gastrics themselves must be divided, or all five of the motor nerves from which their accessory filaments are derived. This fact has been noticed by Longet as showing that nature multiplies the safe- guards of a function in proportion to its importance ; for while the ' lu Longet's Traite de Physiologie. vol. ii. p. 324. 406 THE CEANIAL NERVES. spinal accessory, or any other one of the above mentioned nerves, might be affected by local accident or disease, it would be very improbable that any single injury should paralyze simultaneously the spinal acessory, the facial, the sublingual, and the first and second cervicals. The respiratory movements of the larynx are consequently much more thoroughly protected than those which are merely concerned in the formation of the voice. Lungs. — The influence of the pneumogastric upon the function of the lungs is exceedingly important. The nerve acts here, as in most other organs to which it is distributed, in a double or mixed capacity ; but it is principally as the sensitive nerve of the lungs that it has thus far received attention. It is this nerve which conveys from the lungs to the medulla oblongata that peculiar impression, termed hesoin de respirer, which excites by reflex action the diaphragm and intercostal muscles, and keeps up the play of the respiratory movements. As we have already shown, this action is an involuntary one, and will even take place when consciousness is entirely suspended. It may indeed be arrested for a time by an effort of the will; but the impression conveyed to the medulla soon becomes so strong, and the stimulus to inspiration so urgent, that it can no longer be resisted, and the muscles contract in spite of our attempts to restrain them. A very remarkable effect is accordingly produced on respiration by simultaneous division of both pneumogastric nerves. This experiment is best performed on adult dogs, which may be ether- ized, and the nerves exposed while the animal is in a condition of insensibility, avoiding, in this way, the disturbance of respiration, which would follow if the dissection were performed while the ani- mal was conscious and sensible to pain. After the effects of the etherization have entirely passed off, and respiration and circulation have both returned to a quiescent condition, the two nerves, which have been previously exposed and secured by a loose ligature, may be instantaneously divided, and the effects of the operation readily appreciated. Immediately after the division of the nerves, when performed in the above manner, the respiration is hurried and difStcult, owing to the sudden paralysis of the larynx and partial closure of the glottis by the vocal chords, as already described. This condition, how- ever, is of short continuance. In a few moments, the difficulty of breathing and the general agitation subside, the animal becomes perfectly quiet, and the only remaining visible effect of the opera- PNEUMOGASTRIC. 407 tion is a dimimshed fnquency in the movements of respiration. This diminution is frequently strongly marked from the first, the number of respirations falling at once to ten or fifteen per minute, and be- coming, in an hour or two, still farther reduced. The respirations are performed easily and quietly ; and the animal, if left undisturbed, remains usually crouched in a corner, without giving any special signs of discomfort. If he be aroused and compelled to move about, the frequency of the respiration is temporarily augmented ; but as soon as he is again quiet, it returns to its former standard. By the second or third day, the number of respirations is often reduced to five, four, or even three per minute; when this is the case, the animal usually appears very sluggish, and is roused with difficulty from his inactive condition. At this time, the respiration is not only diminished in frequency, but is also performed in a peculiar manner. The movement of inspiration is slow, easy, and silent, occupying several seconds in its accomplishment; expiration, on the contrary, is sudden and audible, and is accompanied by a well marked expulsive effort, which has the appearance of being, to a certain extent, voluntary in character. The intercostal spaces also sink inward during the lifting of the ribs; and the whole movement of respiration has an appearance of insufficiency, as if the lungs were not thoroughly filled with air. This insufficiency of respira- tion is undoubtedly owing to a peculiar alteration in the pulmonary texture, which has by this time already commenced. Death takes place at a period varying from one to six days after the operation, according to the age and strength of the animal. The only symptoms accompanying it are a steady failure of the respiration, with increased sluggishness and indisposition to be aroused. There are no convulsions, nor any evidences of pain. After death, the lungs are found in a peculiar state of solidification, which is almost exclusively a consequence of this operation, and which is entirely different from ordinary inflammatory hepatization. They are not swollen, but rather smaller than natural. They are of a dark purple color, leathery and resisting to the feel, destitute of crepitation, and infiltrated with blood. Pieces of the lung cut out sink in water. The pleural surfaces, at the same time, are bright and polished, and their cavity contains no effusion or exudation. The lungs, in a word, are simply engorged with blood and empty of air; their tissue having undergone no other alteration. These changes are not generally uniform over both lungs. The organs are usually mottled on their exterior; the variations in color 408 THE CRANIAL NERVES. corresponding with the different degrees of alteration exhibited by different parts. The explanation usually adopted of the above consequences fol- lowing division of the pneumogastrics is as follows: The nerves being divided, the impression which originates in the lungs from the accumulation of carbonic acid, and which is destined to excite the respiratory movements by reflex action, can no longer be trans- mitted to the medulla oblongata. The natural stimulus to respira- tion being wanting, it is, accordingly, less perfectly performed. The respiratory movements diminish in frequency, and, growing con- tinually slower and slower, finally cease altogether, and death is the result. The above explanation, however, is not altogether sufficient. It accounts very well for the diminished frequency of the respiration, but not for its partial continuance. For if the pneumogastric nerves be really the channel through which the stimulus to respiration is conveyed to the medulla, the difficulty is not to understand why respiration should be retarded after division of these nerves, but why it should continue at all. In point of fact, the respiratory movements, though diminished in frequency, continue often for some days after this operation. This cannot be owing to force of habit, or to any remains of nervous influence, as has been some- times suggested, since, when the medulla itself is destroyed, respira- tion, as we know, stops instantaneously, and no attempt at move- ment is made after the action of the nervous centre is suspended. It is evident, therefore, that the pneumogastric nerve, though the chief agent by which the respiratory stimulus is conveyed to the medulla, is not the only one. The lungs are undoubtedly the organs which are most sensitive to an accumulation of carbonic acid, and an imperfect arterialization of the blood ; and the sensa- tion which results from such an accumulation is accordingly first felt in them. There is reason to believe, however, that all the vas- cular organs are more or less capable of originating this impression, and that all the sensitive nerves are capable, to some extent, of trans- mitting it. Although the first disagreeable sensation on holding the breath makes itself felt in the lungs, yet, if we persist in sus- pending the respiration, we soon become conscious that the feeling of discomfort spreads to other parts ; and at last, when the accu- mulation of carbonic acid and the impurity of the blood have become excessive, all parts of the body suffer alike, and are per- vaded by a general feeling of derangement and distress. It is easy. PNEUMOGASTRIC. 409 therefore, to understand why respiration should be retarded, after section of the pneumogastrics, since the chief source of the stimulus to respiration is cut ofl"; but the movements still go on, though more slowly than before, because the other sensitive nerves, which con- tinue to act, are also capable, in an imperfect manner, of conveying the same impression. The immediate cause of death, after this operation, must no doubt be principally sought for in the altered condition of the lungs. These organs are evidently very imperfectly filled with air, for some time previous to death ; and their condition, as shown in post-mortem examination, is evidently incompatible with a due performance of the respiratory function. It is not at all certain, however, that these alterations in the pulmonary tissue are directly dependent on division of the pneumogastric nerves. It must be recollected that when the section of the pneumogastrics is performed in the middle of the neck, the filaments of the inferior laryngeal nerves are also divided, and the narrowing of the glottis, produced by their paralysis, must necessarily interfere with the free admission of air into the chest. This diflSculty, either alone or combined with the diminished frequency of respiration, must have a very con- siderable effect in impeding the pulmonary circulation, and bringing the lungs into such a condition as unfits them for maintaining life. In order to ascertain the comparative influence upon the lungs of division of the inferior laryngeals and that of the other filaments of the pneumogastrics, we have resorted to the following experi- ment. Two pups were taken, belonging to the same litter and of the same size and vigor, about two weeks old. In one of them (No. 1) the pneumogastrics were divided in the middle of the neck; and in the other (No. 2) a section was made at the same time of the inferior laryngeals, the trunk of the pneumogastrics being left un- touched. For the first few seconds after the operation, there was but little difi'erence in the condition of the two animals. There was the same obstruction to the breath (owing to closure of the glottis), the same gasping and sucking inspiration, and the same frothing at the mouth. Yery soon, however, in pup No. 1, the respiratory move- ments became quiescent, and at the same time much reduced in frequency, falling to ten, eight, and five respirations per minute, as usual after section of the pneumogastrics ; while in No. 2 the re- spiration continued frequent as well as laborious, and the general signs of agitation and discomfort were kept up for one or two hours. 410 THE CKANIAL NERVES. The animal, however, after that time became exhausted, cool, and partially insensible, like the other. They both died, between thirty and forty hours after the operation. On post-mortem inspection it was found that the peculiar congestion and solidification of the lungs, considered as characteristic of division of the pneumogastrics, existed to a similar extent in each instance ; and the only appre- ciable difference between the two bodies was that in No. 1 the blood was coagulated, and the abdominal organs natural, while in No. 2 the blood was fluid and the abdominal organs congested. We are led, accordingly, to the following conclusions with regard to the effect produced by division of this nerve. 1. After section of the pneumogastrics, death takes place by a pecu- liar congestion of the lungs. 2. This congestion is not directly produced by division of the nerves, but is caused by the imperfect admission of air into the chest. In adult dogs, the closure of the glottis from paralysis of the laryngeal muscles is less complete than in pups ; but it is still sufficient to exert a very decided influence on respiration, and to take an active part in the production of the subsequent morbid phenomena. We therefore regard the death which takes place after division of both pneuraogastric nerves, as produced in the following man- ner : — The glottis is first narrowed by paralysis of the laryngeal mus- cles, and an imperfect supply of air is consequently admitted, by each inspiration, into the trachea. Next, the stimulus to respiration being very much diminished, the respiratory movements take place more slowly than usual. From these two causes combined, the blood is imperfectly arterialized, and the usual consequence of such a condition then follows, viz., a partial stagnation of the pulmonary circulation. This stagnation still further impedes the action of the lungs; while it does not excite the respiratory muscles to increased activity as it would do in health, owing to the division of the pneu- mogastrics. At the same time, the accumulation of carbonic acid in the blood and the tissues begins to exert a narcotic effect, to diminish the sensibility of the nervous centres, and consequently to retard still more the movements of respiration. Thus all these causes react upon and aggravate each other ; because the connection naturally existing between imperfectly arterialized blood and the stimulus to respiration, is now destroyed. The narcotism and pul- PNEUMOGASTRIC. 411 monary engorgement, therefore, continue to increase, until the lungs are so seriously altered and engorged that they are no longer capable of transmitting the blood, and circulation and respiration come to an end at the same time. It must be remembered, also, that the pneumogastric nerve has other important distributions beside those to the larynx and the lungs; and the effect produced by its division upon these other organs has no doubt a certain share in producing the results which follow. Bearing in mind the very extensive distribution of the pneumogastric nerve and the complicated character of its func- tions, we may conclude that after section of this nerve death takes place from a combination of various causes ; the most active of which is a peculiar engorgement of the lungs and imperfect per- formance of the respiratory function. S(o7nach, and Digestive Function. — After division of the pneumo- gastric nerves, the sensations of hunger and thirst remain, and the secretion of gastric juice continues. Nevertheless the digestive function is disturbed in various ways, though not altogether abo- lished. The appetite is more or less diminished, as it would be after any serious operation, but it remains sufficiently active to show that its existence is not directly dependent on the integrity of the pneu- mogastric nerve. Digestion, however, very seldom actually takes place to any considerable extent, owing to the following circum- stances : The animal is frequently seen to take food and drink with considerable avidity; but in a few moments afterward the food and drink are suddenly rejected by a peculiar kind of regurgitation. This regurgitation does not resemble the action of vomiting, but the substances swallowed are again discharged so easily and instan- taneously as to lead to the belief that they had never passed into the stomach. Such, indeed, is actually the case, as any one may convince himself by watching the process, which is often repeated by the animal at short intervals. The food and drink, taken volun- tarily, pass down into the oesophagus, but owing to the paralysis of the muscular fibres of this canal, are not conveyed into the stomach. They accumulate consequently in the lower and middle part of the oesophagus ; and in a few moments are rejected by a sudden anti- staltic action of the parts, excited, apparently, through the influence of the great sympathetic. The muscular coat of the stomach is also paralyzed to a con- siderable extent by section of this nerve. Longet has shown, by introducing food artificially into the stomach, that gastric juice 412 THE CRANIAL NERVES. may be secreted and the food be actually digested and disappear, when introduced in small quantity. But when introduced in large quantity, it remains undigested, and is found after death with the exterior of the mass softened and permeated by gastric juice, while the central portions are unaltered, and do not even seem to have come in contact with the digestive fluid. This is undoubtedly owing both to the diminished sensibility of the mucous membrane of the stomach, and to the paralysis of its muscular fibres. The peristaltic action of the organ is very important in digestion, in order to bring successive portions of the food in contact with the mucous membrane, and to carry away such as are already softened or as are not capable of being digested in the stomach. This constant movement and agitation of the food is probably also one great stimulus to the continued secretion of the gastric juice. The digestive fluid will therefore be deficient in quantity after division of the pneumogastric nerves, at the same time that the peristaltic movements of the stomach are suspended. Under these circum- stances, the secretion of gastric juice may be sufficient to permeate and digest small quantities of food, while a larger mass may resist its action, and remain undigested. The effect produced by division of these nerves on the digestive, as on the respiratory organs, is therefore of a complicated character, and results from the combined action of several different causes, which influence and modify each other. The effect produced upon the liver by section of the pneumo- gastrics, as well as the influence usually exerted by these nerves upon the hepatic functions, has been so little studied that nothing definite has been ascertained in regard to it. We shall therefore pass over this portion of the subject in silence. That part of the nervous system which we have hitherto studied, viz., the cerebro-spinal system, consists of an apparatus of nerves and ganglia, destined to bring the individual into relation with the external world. By means of the special senses, he is made cognizant of sights, sounds, and odors, by which he is attracted or repelled, and which guide him in the pursuit and choice of food. By the general sensations of touch and the volun- tary movements, he is enabled to alter at will his position and location, and to adapt them to the varying conditions under which he may be placed. The great passages of entrance into the body, and of exit from it, are guarded also by the same portion of the nervous system. The introduction of food into the mouth, and its PNEUMOGASTRIC. 413 passage through the oesophagus to the stomach, are regulated by the same nervous apparatus ; and even the passage of air through the larynx, and its penetration into the lungs, are equally under the guidance of sensitive and motor nerves belonging to the cerebro-spinal system. It will be observed that the above functions relate altogether either to external phenomena or to the simple introduction into the body of food and air, which are destined to undergo nutritive changes in the interior of the frame. If we examine, however, the deeper regions of the body, we find located in them a series of internal phenomena, relating only to the substances and materials which have already penetrated into the frame, and which form or are forming a part of its structure. These are the purely vegetative functions, as they are called ; or those of growth, nutrition, secretion, excretion, and reproduction. These functions, and the organs to which they belong, are not under the direct influence of the cerebro-spinal nerves, but are regulated by another portion of the nervous system, viz., the "ganglionic system;" or, as it is more commonly called, the "sys- tem of the great sympathetic." 414 SYSTEM OF THE GREAT SYMPATHETIC. CHAPTER VI. SYSTEM OF THE GREAT SYMPATHETIC. The sympathetic system consists of a double chain of nervous ganglia, running from the anterior to the posterior extremity of the body, along the front and sides of the spinal column, and connected with each other by slender longitudinal filaments. Each ganglion is reinforced by a motor and sensitive filament derived from the cerebro-spinal system, and thus the organs under its influence are brought indirectly into communication with external objects and phenomena. The nerves of the great sympathetic are distributed to organs over which the consciousness and the will have no imme- diate control, as the intestine, kidneys, heart, liver, &c. The first sympathetic ganglion in the head is the ophthalmic gan- glion. This ganglion is situated within the orbit of the eye, on the outer aspect of the optic nerve. It communicates by slender fila- ments with the carotid plexus, which forms the continuation of the sympathetic system from below; and receives a motor root from the oculo-motorius nerve, and a sensitive root from the ophthalmic branch of the 5th pair. Its filaments of distribution, known as the "ciliary nerves," pass forward upon the eyeball, pierce the sclerotic, and finally terminate in the iris. The next division of the great sympathetic in the head is the sphenopalatine ganglion, situated in the spheno-maxillary fossa. It communicates, like the preceding, with the carotid plexus, and receives a motor root from the facial nerve, and a sensitive root from the superior maxillary branch of the 5th pair. Its filaments are distributed to the levator palati and azygos uvulae muscles, and to the mucous membrane about the posterior nares. The third sympathetic ganglion in the head is the submaxillary, situated upon the submaxillary gland. It communicates with the superior cervical ganglion of the sympathetic by filaments which accompany the facial and external carotid arteries. It derives its sensitive filaments from the lingual branch of the 5th pair, and its SYSTEM OF THE GREAT SYMPATHETIC. 415 Fiff. 149. motor filaments from the facial nerve, by means of the chorda tympani. Its branches of distribution pass to the sides of the tongue and to the submaxillary and sublingual glands. The last sympathetic ganglion in the head is the otic ganglion. It is situated just beneath the base of the skull, on the inner side of the third branch of the trifacial nerve. It sends fila- ments of communication to the carotid plexus; and re- ceives a motor root from the facial nerve, and a sensitive root from the inferior maxil- lary branch of the 5th pair. Its branches are sent to the internal muscle of the mal- leus in the middle ear (tensor tympani), and to the mucous membrane of the tympanum and Eustachian tube. The continuation of the sympathetic nerve in the neck consists of two and some- times of three ganglia, the superior, middle, and inferior. These ganglia communicate with each other, and also with the anterior branches of the cervical spinal nerves. Their filaments follow the course of the carotid artery and its branches, covering them with a network of inter- lacing fibres, and are finally distributed to the substance of the thyroid gland, and to the Course and distribution of the Great Sympa- thetic. walls of the larynx, trachea, pharynx, and oesophagus. By the superior, middle, and inferior cardiac nerves, they also supply sympathetic fibres to the cardiac plexuses and to the substance of the heart. In the chest, the ganglia of the sympathetic nerve are situated on 416 SYSTEM OF THE GREAT SYMPATHETIC. each side the spinal column, just over the heads of the ribs, with which they accordingly correspond in number. Their communi- cations with the intercostal nerves are double ; each sympathetic ganglion receiving two filaments from the intercostal nerve next above it. The filaments originating from the thoracic ganglia are distributed upon the thoracic aorta, and to the lungs and oesophagus. In the abdomen, the continuation of the sympathetic system con- sists principally of the aggregation of ganglionic enlargements situated upon the cceliac artery, known as the semilunar or coeliac ganglion. From this ganglion a multitude of radiating and inoscu- lating branches are sent out, which, from their diverging course and their common origin from a central mass, are termed the "solar plexus," From this, other diverging plexuses originate, which accompany the abdominal aorta and its branches, and are so dis- tributed to the stomach, small and large intestine, spleen, pancreas, liver, kidneys, and supra-renal capsules, and the internal organs of generation. Beside the above ganglia there are in the abdomen four other pairs, situated in front of the lumbar vertebrae, and having similar connections with those occupying the cavity of the chest. Their filaments join the plexuses radiating from the semilunar ganglion. In the pelvis, the sympathetic system is continued by four or five pairs of ganglia, situated on the anterior aspect of the sacrum, and terminating, at the lower extremity of the spinal column, in a single ganglion, the " ganglion impar," which is probably to be regarded as a fusion of two separate ganglia. The entire sympathetic series is in this way composed of nume- rous small ganglia which are connected throughout, first, with each other, secondly, with the cerebro-spinal system, and thirdly, with the internal viscera of the body. The properties and functions of the great sympathetic have been less successfully studied than those of the cerebro-spinal system, owing to the anatomical difficulties in the way of reaching and operating upon this nerve for purposes of experiment. The cerebro- spinal axis and its nerves are easily exposed and subjected to exami- nation. It is also easy to isolate particular portions, and to appreciate the disturbances of sensation and motion consequent upon local lesions or irritations. The phenomena, furthermore, which result from experiments upon this part of the nervous system, are promptly produced, are well-marked in character, and are, as a general rule, readily understood by the experimenter. On the other hand, the SYSTEM OP THE GREAT SYMPATHETIC. 417 principal part of the sympathetic system is situated in the interior of the chest and abdomen ; and the mere operation of opening these cavities, so as to reach the ganglionic centres, causes such a disturb- ance in the functions of vital organs, and such a shock to the system at large, that the results of these experiments have been always more or less confused and unsatisfactory. Furthermore, the con- nections of the sympathetic ganglia Avith each other and with the cerebro-spinal axis are so numerous and scattered, that these ganglia cannot be completely isolated without resorting to an operation still more mutilating and injurious in its character. And finally, the sensible phenomena which are obtained from experimenting on the great sympathetic are, in the majority of cases, slow in making their appearance, and not particularly striking or characteristic in their nature. Notwithstanding these difficulties, however, some facts have been ascertained with regard to this part of the nervous system, which give us a certain degree of insight into its character and functions. The great sympathetic is endowed both with sensibility and the power of exciting motion; but these properties are less active here than in the cerebro-spinal system, and are exercised in a dif- ferent manner. If we irritate, for example, a sensitive nerve in one of the extremities, or apply the galvanic current to the poste- rior root of a spinal nerve, the evidences of pain or of reflex action are acute and instantaneous. There is no appreciable inter- val between the application of the stimulus and the sensations which result from it. On the other hand, experimenters who have operated upon the sympathetic ganglia and nerves of the chest and abdomen find that evidences of sensibility are distinctly manifested here also, but less acutely and only after somewhat prolonged ap- plication of the irritating cause. These results correspond very closely with what we know of the vital properties of the organs which are supplied either principally or exclusively by the sympa- thetic; as the liver, intestine, kidneys, &c. These organs are insensible, or nearly so, to ordinary impressions. We are not con- scious of the changes and operations going on in them, so long as these changes and operations retain their normal character. But they are still capable of perceiving unusual or excessive irritations, and may even become exceedingly painful, when in a state of in- flammation. There is the same peculiar character in the action of the motor nerves belonging to the sympathetic. If the facial or sublingual, 27 418 SYSTEM OF THE GREAT SYMPATHETIC. or the anterior root of a spinal nerve be irritated, the convulsive movement which follows is instantaneous, violent, and only mo- mentary in its duration. But if the semilunar ganglion or its nerves be subjected to a similar experiment, no immediate effect is produced. It is only after a few seconds that a slow, vermicular, progressive contraction takes place in the corresponding part of the intestine, which continues for some time after the exciting cause has been removed. Morbid changes taking place in organs supplied by the sympa- thetic present a similar peculiarity in the mode of their produc- tion. If the body be exposed to cold and dampness, for example, congestion of the kidneys shows itself perhaps on the following (lay. Inflammation of any of the internal organs is very rarely established within twelve or twenty-four hours after the application of the exciting cause. The internal processes of nutrition together with their derangements, which are regarded as especially under the control of the great sympathetic, always require a longer time to be influenced by incidental causes, than those which are regulated by the nerves and ganglia of the cerebro-spinal system. In the head, the sympathetic has a close and important connec- tion with the exercise of the special senses. This is illustrated more particularly, in the case of the eye, by its influence over the alternate expansion and contraction of the pupil. The ophthalmic ganglion sends off a number of ciliary nerves, which are distributed to the iris. It is connected, as we have seen, with the remaining sympathetic ganglia in the head, and receives, beside, a sensitive root from the ophthalmic branch of the 5th pair, and a motor root from the oculo-motorius. The reflex action by which the pupil contracts under a strong light falling upon the retina, and expands under a diminution of light, takes place accordingly through this ganglion. The impression conveyed by the optic nerve to the tubercula quadrigemina, and reflected outward by the fibres of the oculo-motorius, is not transmitted directly by the last named nerve to the iris ; but passes first to the ophthalmic ganglion, and is thence conveyed to its destination by the ciliary nerves. The reflex movements of the iris exhibit consequently a some- what sluggish character, which indicates the intervention of a part of the sympathetic system. The changes in the size of the pupil do not take place instantaneously, with the variation in the amount of light, but always require an appreciable interval of time. If we pass suddenly from a brilliantly lighted apartment into a dark SYSTEM OF THE GREAT SYMPATHETIC. 419 room, we are unable to distinguish surrounding objects until a certain time has elapsed, and the expansion of the pupil has taken place; and vision even continues to grow more and more distinct for a considerable period afterward, as the expansion of the pupil becomes more complete. Again, if we cover the eyes of another person with the hand or a folded cloth, and then suddenly expose them to the light, we shall find that the pupil, which is at first dilated, contracts somewhat rapidly to a certain extent, and afterward con- tinues to diminish in size during several seconds, until the proper equilibrium is fairly established. Furthermore, if we pass sud denly from a dark room into the bright sunshine, we are immedi- ately conscious of a painful sensation in the eye, which lasts for a considerable time ; and which results from the inability of th^- pupil to contract with sufficient rapidity to shut out the excessive amount of light. All such exposures should be made gradually, so that the movements of the iris may keep pace with the varying quantity of stimulus, and so protect the eye from injurious impres- sions. The reflex movements of the iris, however, though accomplished through the medium of the ophthalmic ganglion, derive their original stimulus, through the motor root of this ganglion, from the oculo-motorius nerve. For it has been found that if the last mentioned nerve be divided between the brain and the eyeball, the pupil becomes immediately dilated, and will no longer contract under the influence of light. The motive power originally derived from the brain is, therefore, in the case of the iris, modified by passing through one of the sympathetic ganglia before it reache& its final destination. An extremely interesting fact in this connection is the following: Of the three organs of special sense in the head, viz., the eye, the nose, and the ear, each one is provided with two sets of muscles, superficial and deep, which together regulate the quantity of stimu- lus admitted to the organ. The superficial set of these muscles is animated by branches of the facial nerve ; the deep seated or in- ternal set, by filaments from a sympathetic ganglion. Thus, the front of the eyeball is protected by the orbicularis and levator palpebrae superioris muscles, which open or close the eye- lids at will, and allow a larger or smaller quantity of light to reach the cornea. These muscles are supplied by the oculo-motorius and facial nerves, and are for the most part voluntary in their action. The iris, on the other hand, is a more deeply-seated muscular 420 SYSTEM OF THE GREAT SYMPATHETIC. curtain, winch regulates the quantity of light admitted through the pupil. It is supplied, as we have seen, by filaments from the ophthalmic ganglion, and its movements are altogether involuntary in character. In the olfactory apparatus, the anterior or superficial set of muscles are the compressors and elevators of the alse nasi, which are animated by filaments of the facial nerve. By their action, odoriferous vapors, when faint and delicate in their character, are snuffed up and directed into the upper part of the nasal passages, where they come in contact with the most sensitive portions of the olfactory membrane ; or, if too pungent or disagreeable in flavor, are excluded from entrance. These muscles are not very im- portant or active in the human subject; but in many of the lower animals with a more active and powerful sense of smell, as for example the carnivora, they may be seen to play a very important part in the mechanism of olfaction. Furthermore, the levators and depressors of the velum palati, which are more deeply situated, serve to open or close the orifice of the posterior nares, and accom- plish a similar office with the muscles already named in front. The levator palati and azygos uvulae muscles, which, by their action, tend to close the posterior nares, are supplied by filaments from the spheno-palatine ganglion, and are involuntary in their character. The ear has two similar sets of muscles, similarly supplied. The first, or superficial set, are those moving the external ear, viz., the anterior, superior, and posterior auriculares. Like the muscles of the anterior nares, they are comparatively inactive in man, but in many of the lower animals are well developed and important. In the horse, the deer, the sheep, &c., they turn the ear in various directions so as to catch more distinctly faint and distant sounds, or to exclude those which are harsh and disagreeable. These muscles are supplied by filaments of the facial nerve, and are voluntary in their action. The deep seated set are the muscles of the middle ear. In order to understand their action, we must recollect that sounds are transmitted from the external to the middle ear through the membrane of the tympanum, which vibrates, like the head of a drum, on receiving sonorous impulses from without. Now it is well known that any resonant membrane or cord is capable of vibrating in unison with acute or grave sounds, according to its fctate of tension or relaxation. For any such membrane or cord, at SYSTEM OF THE GREAT SYMPATHETIC. 421 a given degree of tension, there is a limit both to the gravity and acuteness of the sounds which it is capable of transmitting. The greater its tension, the more acute the sounds which may be trans- mitted ; the lower its tension, the deeper the sounds to which it is capable of vibrating. Furthermore, any elastic membrane is more easily thrown into sonorous vibrations when in a tense con- dition, and is consequently more capable, when tightly strained, of receiving and transmitting sounds of feeble intensity. The membrane of the tympanum, accordingly, which is an elastic sheet stretched across the passage to the ear, may be made more or less sensitive to sonorous impressions by varying its con- dition of tension or relaxation. The handle of the malleus is attached to the membrana tympani in such a manner that when the internal muscle of the malleus (tensor tympani) is thrown into contraction, the tympanic membrane is drawn inward, and its tension increased. On the relaxation of this muscle, the chain of bones of the middle ear returns to its ordinary position by the elasticity of its ligaments, and restores the previous condition of the membrana tympani. It is undoubtedly by this mechanism that the sensibility of the hearing is increased or diminished according to circumstances. In listening attentively to a very faint sound, or in endeavoring to distinguish slight variations at a high pitch, a sense of exertion may be almost always perceived, which is proba- bly due in a great degree to the unusual tension of the membrana tympani. On the other hand, sounds of a very sharp and acute character are distressing to the ear, and may be diminished in apparent intensity by a relaxation of the same membrane. The internal muscle of the malleus, by which this action is accom- plished, corresponds therefore in its office with the muscular fibres of the iris, and to those which open and close the posterior nares. It is supplied by filaments from the otic ganglion, the fourth in the series of sympathetic ganglia situated in the head. In all these instances, the reflex action taking place in the deeper seated muscles, originates from a sensation which is con- veyed inward to the cerebro-spinal centres, and is then transmitted outward to its final destination through the medium of one of the sympathetic ganglia. Another very striking fact concerning the sympathetic relates to the changes produced by its division, in the nutritive processes of the parts supplied by it. One of the most important and remark- able of these changes is an elevation of temperalure in the affected 422 SYSTEM OF THE GREAT SYMPATHETIC. parts. If the sympathetic nerve be divided on one side of the neck, in the rabbit, cat, or dog, an elevation of temperature begins to be perceptible on the corresponding side of the head in a very short time. In the cat, we have found a very sensible difference in tem- perature between the two sides at the end of five or ten minutes ; and in the rabbit, at the end of half an hour. A vascular conges- tion of the parts also takes place, which may be seen to great advantage in the ear of the rabbit, when held up between the eye and the light. The elevation of temperature, in these cases, is very perceptible to the touch, and may be also measured by the thermo- meter. Bernard' has found it to reach 8° or 9° F. The elevation <)f temperature and congested state of the parts are sometimes found to be diminished by the next day, and afterward disappear rapidly. Occasionally, however, they last for a long time. Bernard {op. cit.) has seen the unnatural temperature of the affected parts remain in the rabbit for fifteen to eighteen days, and in the dog for two months. Where the superior cervical ganglion has been extir- pated, he has even found the above appearances to continue in the ^ 1 " fevl. ^ ■^^•s I '■^T^ -^":3 \ =.-•" = - ^-, \ @- ' \ '^^^rrroiri^ _ \ Single A h t i c u i, a t i o x of T .b n i a Crassicoi, I, IS, from small intestino of cat. — a, a, a. Ovary filled with eggs. b. Testicle, c. Geaital orifice. 444 SEXUAL GENERATION. upon itself, which opens by an external orifice (c) upon the lateral border of the articulation, about midway between its two ex- tremities. The spermatic fluid produced in the testicle is intro- duced into the female generative passage, which opens at the same spot, and, penetrating deeply into the interior, comes in contact with the eggs, which are thereby fecundated and rendered fertile. The fertile eggs are afterward set free by the rupture or decay of the articulation, and a vast number of young produced by their development. In snails, also, and in some other of the lower animals, the ovaries and testicles are both present in the same individual ; so that these animals are sometimes said to be " hermaphrodite," or of double sex. In realit}^, however, it appears that the male and female organs do not come to maturity at the same time; but the ovaries • are first developed and perform their function, after which the tes- ticles come into activity in their turn. The same individual, there- fore, is not both male and female at any one time ; but is first female and afterward male, exercising the two generative functions at different ages. In all the higher animals, however, the two sets of generative organs are located in separate individuals; and the species is consequently divided into two sexes, male and female. All that is absolutely requisite to constitute the two sexes is the existence of testicles in the one, and of ovaries in the other. Beside these, however, there are, in most instances, certain secondary or acces- sory organs of generation, which assist more or less in the accom- plishment of the process, and which occasion a greater difference in tlie anatomy of the two sexes. Suet are the uterus and mam- mary glands of the female, the vesiculse seminales and prostate of the male. The female naturally having the immediate care of the young after birth, and the male being occupied in providing food and protection for both, there are also corresponding differ- ences in the general structure of the body, which affect the whole external appearance of the two sexes, and which even show them- selves in their mental and moral, as well as in their physical characteristics. In some cases this difference is so excessive that the male and female would never be recognized as belonging to the same species, unless they were seen in company with each other. Not to mention some extreme instances of this among insects and other invertebrate animals, it will be sufficient to refer to the well known examples of the cock and the hen, the lion and lioness, the SEXUAL GENERATION". 445 buck and the doe. In the human species, also, the distinction between the sexes shows itself in the mental constitution, the dis- position, habits, and pursuits, as well as in the general conforma- tion of the body, and the peculiarities of external appearance. We shall now study more fully the character of the male and female organs of generation, together with their products, and the manner in which these are discharged from the body, and brought into relation with each other. 446 EGG AND FEMALE ORGANS OF GENERATION. CHAPTER III. ON THE EGG, AND THE FEMALE ORGANS OF GENERATION. The egg is a globular body which varies considerably in size in different classes of animals, according to the peculiar conditions under which its development is to take place. In the frog it mea- sures yig of an inch, in the lamprey g'g, in quadrupeds and in the human species y|(^. It consists, first, of a membranous external sac or envelope, the vitelline membrane; and secondly, of a spherical mass inclosed in its interior, called the vitellus. The vitelline membrane in birds and reptiles is very thin, measur- ing often not more than tsoct) of an inch in thickness, and is at the same time of a somewhat fibrous texture, ^^g- ^^^- In man and the higher animals, on the contrary, it is perfectly smooth, structure- less and transparent, and is about ^^^-q of an inch in thickness. Notwithstanding its delicate and transparent appearance, it has a considerable degree of resistance and elasticity. The egg of the human subject, for example, may be perceptibly flattened out under the microscope by pressing with the point of a needle upon the slip of glass which covers it; but it still remains unbroken, and when the pressure is removed, readily resumes its globular form. When the egg is somewhat flattened under the microscope in this way, by pressure of the glass slip, the apparent thickness of the vitelline membrane is increased, and it then appears (Fig. 159) as a rather wide, colorless, and pellucid border or zone, surrounding the granu- lar and opaque vitellus. Owing to this appearance, it has some- times received the name of the "zona pellucida." The name of vitelline membrane, however, is the one more generally adopted, and is also the more appropriate of the two. Human Ovum, magnified S.3 diameters, a. Virelline mpmbrane. h. Vitellus. c. Genniuative vesicle. d. Germinative spot. EGG AND FEMALE ORGANS OF GENERATION. 447 The vilellus {h), is a globular, semi-solid mass, contained within the vitelline membrane. It consists of a colorless albuminoid sub- stance, with an abundance of minute molecules and oleaginous granules scattered through it. These minute oleaginous masses give to the vitellus a partially opaque and granular aspect under the microscope. Imbedded in the vitellus, usually near its surface and almost immediately beneath the vitelline membrane, there is a clear, colorless, transparent vesicle (c) of a rounded form, known as the germinative vesicle. In the egg of the human subject and of the quadrupeds, this vesicle measures ggo to g J^ of an inch in diameter. It presents upon its surface a dark spot, like a nucleus {d), which is known ^'S* ^'^^' by the name of the germinative spot. The germinative vesicle, with its nucleus-like spot, is often partially concealed by the granules of the vitellus by which it is sur- rounded, but it may always be discovered by careful examination. If the egg be ruptured by excessive pres- human ovum, luptmed ly , , . , . ,, . pressure; showing the vitellus sure under the, microscope, the vitellus is partially expelled, the germlnu- seen to have a gelatinous consistency. It '*'<^ ''^^'c'e at «, and the smooth . fracture of the vitelline mem- is gradually expelled from the vitelline brane. cavity, but still retains the granules and oil globules entangled in its substance. (Fig. 160.) The edges of the fractured vitelline membrane, under these circumstances, present a smooth and nearly straight outline, without any appearance of laceration or of a fibrous structure. The membrane is, to all ap- pearance, perfectly homogeneous. The most essential constituent of the egg is the vitellus. It is from the vitellus that the body of the embryo will afterward be formed, and the organs of the new individual developed. The vitelline membrane is merely a protective inclosure, intended to jirotect the vitellus from injury, and enable it to retain its figure during the ea:rly periods of development. The egg, as above described, consists therefore of a simple vitellus of minute size, and a vitelline membrane inclosing it. It is such an egg which is found in the human subject, the quadru- peds, most aquatic reptiles, very many fish, and some invertebrate jinimals. In nearly all those species, in fact, where the fecundated eggs are deposited and hatched in the water, as well as those in which they are retained in the body of the female until the develop- 448 EGa AND FEMALE ORGANS OF GENERATION. merit of the young is completed, such an egg as above described is sufficient for the formation of the embryo; since during its develop- ment it can absorb freely, either from the water in which it floats, or from the mucous membrane of the female generative organs, the requisite supply of nutritious fluids. But in birds and in the terrestrial reptiles, such as lizards, tortoises, &c., where the eggs are expelled from the body of the female at an early period, and incubated on land, there is no external source of nutrition, to pro- vide for the support of the young animal during its development. In these instances accordingly the vitellus, or "yolk," as it is called, is of very large size; and the bulk of the egg is still further in- creased by the addition, within the female generative passages, of layers of albumen and various external fibrous and calcareous envelopes. The essential constituents of the egg, however, still remain the same in character, and the process of embryonic develop- ment follows the same general laws as in other cases. The eggs are produced in the interior of certain organs, situated in the abdominal cavity, called the ovaries. These organs consist of a number of globular sacs, or follicles, known as the "Graafian follicles," each one of which contains a single egg. The follicles are connected with each other by a quantity of vascular areolar tissue, which binds them together into a well defined and consistent mass, covered upon its exterior by a layer of peritoneum. The egg has sometimes been spoken of as a " product," or even as a " secretion" of the ovary. Nothing can be more inappropriate, however, than to compare the egg with a secretion, or to regard the ovary as in any respect resembling a glandular organ. The egg is simply an organized body, growing in the ovary like a tooth in its follicle, and forming a constituent part of the body of the female. It is destined to be finally separated from its attachments and thrown off; but until that time, it is, properly speaking, a part of the ovarian texture, and is nourished like any other portion of the female organism. The ovaries, accordingly, since they are directly concerned in the production of the eggs, are to be regarded as the essential parts of the female generative apparatus. Beside them, however, there are usually present certain other organs, which play a secon- dary or accessory part in the process of generation. The most important of these accessory organs are two symmetrical tubes, or oviducts, which are destined to receive the eggs at their internal extremity and convey them to the external generative orifice. The EGG AND FEMALE ORGANS OF GENERATION. 449 mucous membrane lining the oviducts is also intended to supply certain secretions during the passage of the egg, which are requi- site either to complete its structure, or to provide for the nutrition of the embryo. In the frog, for example, the oviduct commences at the upper part of the abdomen by a rather wide orifice, which communicates directly with the peritoneal cavity. It soon after contracts to a narrow tube, and pursues a zigzag course down the side of the abdomen (Fig. 161), folded upon itself in convolutions, like the small intestine, until it opens, near its fellow of the opposite side, into the "cloaca," or lower part of the intestinal canal. The oviducts present the same general characters with those described above, in nearly all species of reptiles and birds ; though there are some modi- fications, in particular instances, which do not require any special notice. The ovaries, as well as the egg which they contain, undergo at particular sea- sons a periodical development or increase in growth. If we examine the female frog iti the latter part of summer or the fall, we shall find the ovaries presenting the appearance of small clusters of minute and nearly colorless eggs, the smaller of which are perfectly transparent and not over y^o of an inch in diameter. But in the early spring, when the season of reproduction approaches, the ovaries will be found in- creased to four or five times their former size, and forming large lobulated masses, crowded with dark-colored opaque eggs, measur- ^i''g T-2 of ^^ ii^ch in diameter. At the approach of the generative season, in all the lower animals, a certain number of the eggs, which were previously in an imperfect and inactive condition, begin to in- crease in size and become somewhat altered in structure. The vitel- lus more especially, which was before colorless and transparent, becomes granular in texture as well as increased in volume ; and assumes at the same time, in many species of animals, a black, brown, yellow, or orange color. In the human subject, however, 29 Female Generative Or- gans OF Frog. — a, a Ovaries. b, b. Oviducts, e, c. Tlieir intertiiil oi-ifices. d. Cloaca, shovviug exter- nal orifices of oviducts. 450 EGG AND FEMALE ORGANS OF GENERATION. the change consists only in an increase of size and granulation, without any remarkable alteration of color. The eggs, as they ripen in this way, becoming enlarged and changed in texture, gradually distend the Graafian follicles and project from the surface of the ovary. At last, when fully ripe, they are discharged by a rupture of the walls of the follicles, and, passing into the oviducts, are conveyed by them to the external generative orifice, and there expelled. In this way, as successive seasons come round, successive crops of eggs enlarge, ripen, leave the ovaries, and are discharged. Those which are to be expelled at the next generative epoch may always be recognized by their greater degree of development; and in this way, in many animals, the eggs of no less than three different crops may be recognized in the ovary at once, viz., 1st, those which are perfectly mature and ready to be discharged ; 2d, those which are to ripen in the follow- ing season ; and 3d, those which are as yet altogether inactive and undeveloped. In most fish and reptiles, as well as in birds, this regular process of maturation and discharge of eggs takes place but once a year. In different species of quadrupeds it may take place annually, semi-annually, bimonthly, or even monthly; but in every instance it recurs at regular intervals, and exhibits accord- ingly, in a marked degree, the periodic character which we have seen to belong to most of the other vital phenomena. Action of the Oviducts and Female Generative Passages. — In frogs and lizards, the ripening and discharge of the eggs take place, as above mentioned, in the early spring. At the time of leaving the ovary, the eggs consist simply of the dark colored and granulnr vitellus, inclosed in the vitelline membrane. They are then received by the inner extremity of the oviducts, and carried downward by the peristaltic movement of these canals, aided by the more power- ful contraction of the abdominal mus- Fig. 162. cles. During the passage of the eggs, moreover, the mucous membrane of the oviduct secretes a colorless, viscid, %*• (® " * ^ -< albuminoid substance, which is depo- •® ^-7 ^-V**) sited in successive layers round each » %ti^ ggg^ forming a thick and tenacious Mature Frogs' eogs.-«. While coating or envclopc. (Fig. 162.) When «tiii in the ovary. 6. After passing ^^^ ^^^ finally discharged, this through the oviduct. OO . -^ o ' albuminoid matter absorbs the water in which the spawn is deposited, and swells up into a transparent EGG AND FEMALE ORGANS OF GENERATION. 451 gelatinous mass, in which the eggs are separately imbedded. This substance supplies, by its subsequent liquefaction and absorption, a certain amount of nutritious material, during the development and early growth of the embryo. In the terrestrial reptiles and in birds, the oviducts perform a still more important secretory function. In the common fowl, the ovary consists, as in the frog, of a large number of follicles, loosely connected by areolar tissue, in which the eggs can be seen in different stages of development. (Fig, ld3, a.) As the egg which is approach- ing maturity enlarges, it distends the cavity of its follicle and pro- jects farther from the general surface of the ovary; so that it hangs at last into the peritoneal cavity, retained only by the attenuated wall of the follicle, and a slender pedicle through which run the bloodvessels by which its circulation is supplied. A rupture of the follicle then occurs, at its most prominent part, and the egg is dis- charged from the lacerated opening. At the time of its leaving the ovary, the egg of the fowl consists of a large, globular, orange-colored vitellus, or " yolk," inclosed in a thin and transparent vitelline membrane. Immediately under- neath the vitelline membrane, at one point upon the surface of tlie vitellus, is a round white spot, consisting of a layer of minute granules, termed the ''cicatricula." It is in the central part of the cicatricula that the germinative vesicle is found imbedded, at an early stage of the development of the egg. At the time of its discharge from the ovary, the germinative vesicle has usually dis- appeared ; but the cicatricula is still a very striking and important part of the vitellus, as it is from this spot that the body of the chick begins afterward to be developed. At the same time that the egg protrudes from the surface of the ovary, it projects into the inner orifice of the oviduct ; so that; when discharged from its follicle, it is immediately embraced by the upper or fringed extremity of this tube, and commences its passage down- ward. In the fowl, the muscular coat of the oviduct is highly deve- loped, and its peristaltic contractions gently urge the egg from above downward, precisely as the oesophagus or the intestines transport the food in a similar direction. While passing through the first two or three inches of the oviduct (c, 7, c), measuring not less than ^\ of an 460 MALE ORGANS OF GENERATION. inch in length, about one-third of which is occupied by the head, or enlarged portion of the filament. The most remarkable peculiarity of the spermatozoa is their very singular and active movement, to which we have already alluded. If a drop of fresh seminal fluid be placed under the microscope, the numberless minute filaments with which it is crowded are seen to be in a state of incessant and agitated motion. This movement of the spermatozoa, in many species of animals, strongly resembles that of the tadpole; particularly when, as in the human subject, the rabbit, &c., the spermatozoa consist of a short and well defined head, followed by a long and slender tail. Here the tail-like filament keeps up a constant lateral or vibratory movement, by which the spermatozoon is driven from place to place in the spermatic fluid, just as the fish or the tadpole is pro- pelled through the water. In other instances, as for example in the water-lizard, and in some parasitic animals, the spermatozoa have a continuous writhing or spiral-like movement, which pre- sents a very peculiar and elegant appearance when large numbers of them are viewed together. It is the existence of this movement which first suggested the name of spermatozoa to designate the animated filaments of the spermatic fluid ; and which has led some writers to attribute to them an independent animal nature. This is, however, a very erroneous mode of regarding them ; since they cannot properly be considered as animals, notwithstanding the active character of their movement, and the striking resemblance which it sometimes pre- sents to a voluntary act. The spermatozoa are organic forms, which are produced in- the testicles, and constitute a part of their tissue ; just as the eggs, which are produced in the ovaries, natu- rally form a part of the texture of these organs. Like the egg, also, the spermatozoon is destined to be discharged from the organ where it grew, and to retain, for a certain length of time afterward, its vital properties. One of the most peculiar of these properties is its power of keeping in constant motion ; which does not, how- ever, mark it as a distinct animal, but only distinguishes it as a peculiar structure belonging to the parent organism. The motion of a spermatozoon is precisely analogous to that of a ciliated epi- thelium cell. The movement of the latter will continue for some hours after it has been separated from its mucous membrane, pro- vided its texture be not injured, nor the process of decomposition allowed to commence. In the same manner, the movement of the MALE ORGANS OF GENERATION. • 461 spermatozoa is a characteristic property belonging to them, which continues for a certain time, even after they have been separated from all connection with the rest of the body. In order to preserve their vitality, the spermatozoa must be kept at the ordinary temperature of the body, and preserved from the contact of the air or other unnatural fluids. In this way, they may be kept without difficulty many hours for purposes of examination. But if the fluid in which they are kept be allowed to dry, or if it be diluted by the addition of water, in tli'e case of birds and quadrupeds, or if it be subjected to extremes of heat or cold, the motion ceases, and the spermatozoa themselves soon begin to disintegrate. The spermatozoa are produced in certain glandular-looking organs, the testicles^ w^hich are characteristic of the male, as the ovaries are characteristic of the female. In man and all the higher animals, the testicles are solid, ovoid-shaped bodies, composed principally of numerous long, narrow, and convoluted tubes, the "seminiferous tubes," somewhat similar in their general anatomical characters to the tubuli uriniferi of the kidneys. These tubes lie for the most part closely in contact with each other, so that nothing intervenes between them except capillary bloodvessels and a little areolar tissue. They commence, by blind, rounded extremities, near the external surface of the testicle, and pursue an intricately con- voluted course toward its central and posterior part. They are not strongly adherent to each other, but may be readily unravelled by manipulation, and separated from each other. The formation of the spermatozoa, as it takes place in the substance of the testicle, has been fully investigated by Kolliker. According to his observations, as the age of puberty approaches, beside the ordinary pavement epithelium lining the seminiferous tubes, other cells or vesicles of larger size make their appearance in these tubes, each containing from one to fifteen or twenty nuclei, with nucleoli. It is in the interior of these vesicles that the spermatozoa are formed ; their number corresponding usually with that of the nuclei just mentioned. They are at first developed in bundles of ten to twenty, held together by the thin membranous substance which surrounds them, but are afterward set free by the liquefaction of the vesicle, and then fill nearly the entire cavity of the seminiferous ducts, mingled only with a YGvy minute quantity of transparent fluid. In the seminiferous tubes themselves, the spermatozoa are always 462 MALE ORGANS OF GENERATION, inclosed in the interior of their parent vesicles; they are liberated, and mingled promiscuously together, only after entering the rete testis and the head of the epididymis. Beside the testicles, which are, as above stated, the primary and essential parts of the male generative apparatus, there are certain secondary or accessory organs, by means of which the spermatic fluid is conveyed to the exterior, and mingled with certain secre- tions .which assist in the accomplishment of its functions. As the sperm leaves the testicle, it consists, as above mentioned, almost entirely of the spermatozoa, crowded together in an opaque, white, semi-fluid mass, which fills up the vasa eflferentia, and com- pletely distends their cavities. It then enters the single duct which forms the body and lower extremity of the epididymis, following the long and tortuous course of this tube, until it becomes continuous with the vas deferens; through which it is still conveyed onward to the point where this canal opens into the urethra. Throughout this course, it is mingled with a glairy, mucus-like fluid, secreted by the walls of the epididymis and vas deferens, in which the spermatozoa are enveloped. The mixture is then deposited in the vesiculee seminales, where it accumulates as fresh quantities are produced in the testicle and conveyed down- ward by the spermatic duct. It is probable that a second secretion is supplied also by the internal surface of the vesiculs seminales, and that the sperm, while retained in their cavities, is not only stored up for subsequent use, but is at the same time modified in its properties by the admixture of another fluid. At the time when the evacuation of the sperm takes place, it is driven out from the seminal vesicles by the muscular contraction of the surrounding parts, and meets in the urethra with the secre- tions of the prostate gland, the glands of Cowper, and the mucous follicles opening into the urethral passage. All these organs are at that time excited to an unusual activity of secretion, and pour out their different fluids in great abundance. The sperm, therefore, as it is discharged from the urethra, is an exceedingly mixed fluid, consisting of the spermatozoa derived from the testicles, together with the secretions of the epididymis and vas deferens, the prostate, Cowper's glands, and the mucous follicles of the urethra. Of all these ingredients, it is the sperma- tozoa which constitute the essential part of the seminal fluid. They are the true fecundating element of the sperm, while all the others are secondary in importance and perform only accessory functions. Spallanzani found that if frog's semen be passed through a sue- MALE OEGANS OF GENERATION. 463 cession of filters, so as to separate the spermatozoa from the liquid portions, the filtered fluid is destitute of any fecundating properties; while the spermatozoa remaining entangled in the filter, if mixed with a sufficient quantity of fluid of the requisite density for dilu- tion, may still be successfully used for the impregnation of eggs. It is well known, also, that animals or men from whom both testicles have been removed, are incapable of impregnating the female or her eggs ; while a removal or imperfection of any of the other generative organs does not necessarily prevent the accom- plishment of the function. In most of the lower orders of animals there is a periodical development of the testicles in the male, corresponding in time with that of the ovaries in the female. As the ovaries enlargje and the eggs ripen in the one sex, so in the other the testicles increase in size, as the season of reproduction approaches, and become turgid with spermatozoa. The accessory organs of generation, at the same time, share the unusual activity of the testicles, and become increased in vascularity and ready to perform their part in the reproductive function. In the fish, for example, where the testicles occupy a similar posi- tion in the abdomen as the ovaries in the opposite sex, these bodies enlarge, become distended with their contents, and project into the peritoneal cavity. Each of the two sexes is then at the same time under the influence of a corresponding excitement. The unusual development of the generative organs reacts upon the entire system, and produces a state of peculiar activity and excitability, known as the condition of "erythism." The female, distended with eggs, feels the impulse which leads to their expulsion ; while the male, bear- ing the weight of the enlarged testicles and the accumulation of newly-developed spermatozoa, is impelled by a similar sensation to the discharge of the spermatic fluid. The two sexes, accordingly, are led by instinct at this season to frequent the same situations. The female deposits her eggs in some spot favorable to the protec- tion and development of the young ; after which the male, appa- rently attracted and stimulated by the sight of the new-laid eggs, discharges the spermatic fluid upon them, and their impregnation is accomplished. In such instances as the above, where the male and female gene- rative products are discharged separately by the two sexes, the subsequent contact of the eggs with the spermatic fluid would seem to depend altogether on the occurrence of fortuitous circumstances, and their impregnation, therefore, be often liable to fail. In point 464 * MALE ORGANS OF GENERATION. of fact, however, the simultaneous functional excitement of the two sexes and the operation of corresponding instincts, leading them to ascend the same rivers and to frequent the same spots, provide with sufficient certainty for the impregnation of the eggs. In these animals, also, the number of eggs produced by the female is very large, the ovaries being often so distended as to fill nearly the whole of the abdominal cavity; so that, although many of the eggs may be accidentally lost, a sufficient number will still be im- pregnateid and developed to provide for the continuation of the species. In other instances, an actual contact takes place between the sexes at the time of reproduction. In the frog, for example, the male fastens himself upon the back of the female by the anterior extremities, which seem to retain their hold by a kind of spasmodic contraction. This continues for one or two days, during which time the mature eggs, which have been discharged from the ovary, are passing downward through the oviducts. At last they are ex- pelled from the anus, while at the same time the seminal fluid of the male is discharged upon them, and impregnation takes place. In the higher classes of animals, however, and in man, where the egg is to be retained in the body of the female parent during its development, the spermatic fluid is introduced into the female generative passages by sexual congi-ess, and meets the egg at or soon after its discharge from the ovary. The same correspondence, however, between the periods of sexual excitement in the male and female, is visible in many of these animals, as well as in fish and reptiles. This is the case in most species which produce young but once a year, and at a fixed period, as the deer and the wild hog. In other species, on the contrary, such as the dog, as well as the rabbit, the guinea pig, &c., where several broods of young are produced during the year, or where, as in the human subject, the generative epochs of the female recur at short intervals, so that the particular period of impregnation is comparatively indefinite, the generative apparatus of the male is almost constantly in a state of full deve- lopment ; and is excited to action at particular periods, apparently by some influence derived from the condition of the female. In the quadrupeds, accordingly, and in the human species, the contact of the sperm with the egg aud the fecundation of the latter take place in the generative passages of the female ; either in the uterus, the Fallopian tubes, or even upon the surface of the ovary ; in each of which situations the spermatozoa have been found, after the accomplishment of sexual intercourse. PERIODICAL OVULATION. 465 CHAPTER V. ON PERIODICAL OVULATION, AND THE FUNCTION OF MENSTRUATION. L PERIODICAL OVULATION. We have already spoken in general terms of the periodical ripen- ing of the eggs and their discharge from the generative organs of the female. This function is known by the name of " ovulation," and may be considered as the primary and most important act in the process of reproduction. We shall therefore enter more fully into the consideration of certain particulars in regard to it, by which its nature and conditions may be more clearly understood. 1st, Eggs exist originally in the ovaries of all animals^ as part of their natural structure. In describing the ovaries of iish and reptiles we have said that they consist of nothing more than Graafian vesi- cles, each vesicle containing an egg^ and united with each other by loose areolar tissue and a peritoneal investment. In the higher animals and in the human subject, the essential constitution of the ovary is the same ; only its fibrous tissue is more abundant, so that the texture of the entire organ is more dense, and its figure more compact. In all classes, however, without exception, the interior of each Graafian vesicle is occupied by an Qgg ; and it is from this egg that the young ofispring is afterward to be produced. The process of reproduction was formerly regarded as essentially different in the oviparous and viviparous animals. In the ovipa- rous classes, such as most fish, and all reptiles and birds, the young animal was well known to be formed from an egg produced by the female; while in the viviparous animals, or those which brino- forth their young alive, such as the quadrupeds and the human species, the embryo was supposed to originate in the body of the female, by some altogether peculiar and mysterious process, in consequence of sexual intercourse. As soon, however, as the microscope began to be used in the examination of the tissues, 30 466 OVULATION AND FUNCTION OF MENSTRUATION. the ovaries of quadrupeds were also found to contain eggs. These eggs had previously escaped observation on account of their simple structure and minute size ; but they were nevertheless found to possess all the most essential characters belonging to the larger eggs of the oviparous animals. The true difference in the process of reproduction, between the two classes, is therefore merely an apparent, not a fundamental one. In fish, reptiles, and birds, the egg is discharged by the female before or immediately after impregnation, and the embryo subse- quently developed and hatched externally. In the quadrupeds and the human species, on the other hand, the egg is retained within the body of the female until the embryo is developed ; when the membranes are ruptured and the young expelled at the same time. In all classes, however, viviparous as well as oviparous, the young is produced equally from an egg; and in all classes the egg, sometimes larger and sometimes smaller, but always consisting essentially of a vitellus and a vitelline membrane, is contained originally in the interior of an ovarian follicle. The egg is accordingl}'-, as we have already intimated, an integral part of the ovarian tissue. It may be found there long before the generative function is established, and during the earliest periods of life. It may be found without difficulty in the newly bora female infant, and may even be detected in the foetus before birth. Its growth and nutrition, also, are provided for in the same man- ner with that of other portions of the bodily structure. 2d. 27iese eggs become more fully developed at a certain age, ichen the generative function is about to be established. During the early periods of life, the ovaries and their contents, like many other organs, are imperfectly developed. They exist, but they are as yet inactive, and incapable of performing any function. In the yo^ng chick, for example, the ovary is of small size ; and the eggs, instead of presenting the voluminous, yellow, opaque vitellus which they afterward exhibit, are minute, transparent, and colorless. In the young quadrupeds, and in the human female during infancy and childhood, the ovaries are equally inactive. They are small, friable, and of a nearly homogeneous appearance to the naked eye ; presenting none of the enlarged follicles, filled with transparent fluid, which are afterward so readily distinguished. At this time, accordingly, the female is incapable of bearing young, because the ovaries are inactive, and the eggs which they contain immature. At a certain period, however, which varies in the time of its PERIODICAL OVULATION. 467 occurrence for different species of animals, the sexual apparatus begins to enter upon a state of activity. The ovaries increase in size, and their circulation becomes more active. The eggs, also, instead of remaining quiescent, take on a rapid growth, and the structure of the vitellus is completed by the abundant deposit of oleaginous granules in its interior. Arrived at this state, the eggs are ready for impregnation, and the female becomes capable of bearing young. She is then said to have arrived at tl^ state of " puberty," or that condition in which the generative organs are fully developed. This condition is accompanied by a visible alteration in the system at large, which indicates the complete development of the entire organism. In many birds, for example, the plumage assumes at this period more varied and brilliant colors; and in the common fowl the comb, or "crest," enlarges and becomes red and vascular. In the American deer (Cervus virginianus), the coat, which during the first year is mottled with white, becomes in the second year, of a uniform tawny or reddish tinge. In nearly all species, the limbs become more compact and the body more rounded ; and the whole external appearance is so altered, as to indicate that the animal has arrived at the period of puberty, and is capable of reproduction. 3d. Successive crops of eggs^ in the adult female, ripen and are discharged independently of sexual intercourse. It was formerly sup- posed, as we have mentioned above, that in the viviparous animals the germ was formed in the body of the female only as a conse- quence of sexual intercourse. Even after the important fact became known that eggs exist originally in the ovaries of these animals, and are only fecundated by the influence of the sperm- atic fluid, the opinion still prevailed that the occurrence of sexual intercourse was the cause of their being discharged from the ovary, and that the rupture of a Graafian vesicle in this organ was a certain indication that coitus had taken place. This opinion, however, was altogether unfounded. We already know that in fish and reptiles the mature eggs not only leave the ovary, but are actually discharged from the body of the female while still unimpregnated, and only subsequently come in contact with the spermatic fluid. In fowls, also, it is a matter of common observation that the hen will continue to lay fully-formed eggs, if well supplied with nourishment, without the presence of the cock ; only these eggs, being unimpregnated, are incapable of producing 468 OVULATION AND FUNCTION OF MENSTRUATION. chicks. In oviparous animals, therefore, the discharge of the egg, as well as its formation, is independent of sexual intercourse. Continued observation shows this to be the case, also, in the viviparous quadrupeds. The researches of Bischoff, Pouchet, and Coste have demonstrated that in the sheep, the pig, the bitch, the rabbit, &c., if the female be carefully kept from the male until after the period of puberty is established, and then killed, examination of the q«iries will show that Graafian vesicles have matured, rup- tured, and discharged their eggs, in the same manner as though sexual intercourse had taken place. Sometimes the vesicles are found distended and prominent upon the surface of the ovary ; sometimes recently ruptured and collapsed; and sometimes in vari- ous stages of cicatrization and atrophy, Bischoff',' in several in- stnnces of this kind, actually found the unimpregnated eggs in the oviduct, on their way to the cavity of the uterus. In those animals in which the ripening of the eggs takes place at short intervals, as, for example, the sheep, the pig, and the cow, it is very rare to exa- mine the ovaries in any instance where traces of a more or less recent rupture of the Graafian follicles are not distinctly visible. One of the most important facts, derived from the examination of such cases as the above, is that the ovarian eggs become deve- loped and are discharged in successive crops, which follow each other regularly at periodical intervals. If we examine the ovary of the fowl, for example (Fig. 163), we see at a glance how the eggs grow and ripen, one after tlie other, like fruit upon a vine. In this instance, the process of evolution is very rapid ; and it is easy to distinguish, at the same time, eggs which are almost microscopic in size, colorless, and transparent; those which are larger, firmer, somewhat opaline, and yellowish in hue ; and finally those which are fully developed, opaque, of a deep orange color, and just ready to leave the ovary. It will be observed that in this instance the difference between the undeveloped and mature eggs consists principally in the size of the vitellus, which is furthermore, for reasons previously given (Chap. III.), very much larger than in the quadrupeds. It is also seen that it is the increased size of the vitellus alone, by which the ovarian follicle is distended and ruptured, and the egg finally dis- charged. In the human species and the quadrupeds, on the other hand, ' Memoire sur la cliute perioflique de I'oeuf, &c., Annales fles Sciences Naturelles, Aoiit — Septembre, 1844. PERIODICAL OVULATION". 469 the microscopic egg never becomes large enough to distend the follicle by its own size. The rupture of the follicle and the libera- tion of the egg are accordingly provided for, in these instances, by a totally different mechanism. In the earlier periods of life, in man and the higher animals, the egg is contained in a Graafian follicle which closely embraces its exterior, and is consequently hardly larger than the egg itself. As puberty approaches, those follicles which are situated near the free surface of the ovary become enlarged by the accumulation of a colorless serous fluid in their cavity. We then find that the ovary, when cut open, shows a considerable number of globular, transpa- rent vesicles, readily perceptible by the eye, the smaller of which are deep seated, but which increase in size as they approach the free surface of the organ. These vesicles are the Graafian follicles, which, in consequence of the advancing maturity of the eggs con- tained in them, gradually enlarge as the period of generation ap- proaches. The Graafian follicle at this time consists of a closed globular sac or vesicle, the external wall of which, though quite translucent, has a fibrous texture under the microscope and is well supplied with bloodvessels. This fibrous and vascular wall is distinguished by the name of the " membrane of the vesicle." It is not very firm in texture, and if roughly handled is easily ruptured. The membrane of the vesicle is lined throughout by a thin layer of minute granular cells, which form for it a kind of epithelium, similar to the epithelium of the pleura, pericardium, and other serous membranes. This layer is termed the membrana granulosa. It adheres but slightly to the membrane of the vesicle, and may easily be detached by careless manipulation before the vesicle is opened, being then mingled, in the form of light flakes and shreds, with the serous fluid contained in the vesicle. At the most superficial part of the Graafian follicle, or that which is nearest the surface of the ovary, the membrana granulosa is thicker than elsewhere. Its cells are here accumulated, in a kind of mound or "heap," which has received the name of the cumulus proligervs. It is sometimes called the discus proligerus^ because the thickened mass, when viewed from above, has a some- what circular or disk-like form. In the centre of this thickened portion of the membrana granulosa the egg is imbedded. It is accordingly always situated at the most superficial portion of the follicle, and advances in this way toward the surface of the ovary. 470 OVULATION AND FUNCTION OF MENSTRUATION. As the period approaches at which the egg is destined to be dis- charged, the Graafian follicle becomes more vascular, and enlarges by an increased exudation of serum into its cavity. It then begins Fig. 168. Graafian Follicle, near the period of rupture. — a. Membrane of the vesicle. 6. Membrana granulosa, c. Cavity of follicle, d. Egg. e. Peritoneum. /. Tunica albuginea. g, g. Tissue of llie ovary. to project from the surface of the ovary, still covered by the albu- gineous tunic and the peritoneum, (Fig. 168.) The constant accu- mulation of fluid, however, in the follicle, exerts such a steady and increasing pressure from within outward, that the albugineous tunic and the peritoneum successively yield before it; until the Graafian follicle protrudes from the ovary as a tense, rounded, translucent vesicle, in which the sense of fluctuation can be readily perceived on applying the fingers to its surface. Finally, the process of effu- sion and distension still going on, the wall of the vesicle yields at its most prominent portion, the contained fluid is driven out with a o'ush, by the reaction and elasticity of the neighboring ovarian tissues, carrying with it the egg^ still entangled in the cells of the proligerous disc. The rupture of the Graafian vesicle is accompanied, in some instances, by an abundant hemor- rhage, which takes place from the internal surface of the congested follicle, and by which its cavity is filled with blood. This occurs in the human subject and in the pig, and to a certain extent, also, in other of the lower animals. Sometimes, as in the cow, where Fig. 169, Ovary with Graafian Follicle kdptured; at a, egg just discluirged with a portion of membraua granulosa. PERIODICAL OVULATION. 471 iio hemorrhage takes place, the Graafian vesicle when ruptured simply collapses; after which, a slight exudation, more or less tinged with blood, is poured out during the course of a few hours. Notwithstanding, however, these slight variations, the expulsion of the egg takes place, in the higher animals, always in the manner above described, viz., bj the accumulation of serous fluid in the cavity of the Graafian follicle, by which its walls are gradually dis- tended and finally ruptured. This process takes place in one or more Graafian follicles at a time, according to the number of young which the animal produces at a birth. In the bitch and the sow, where each litter consists of from six to twenty young ones, a similar number of eggs ripen and are discharged at each period. In the mare, the cow, and in the iiuman female, where there is usually but one foetus brought forth at a birth, the eggs are matured singly, and the Graafian vesicles luptured, one after another, at successive periods of ovulation. 4th. The ripening and discharge of the egg are accompanied by a pecu- liar condition of the entire system^ known as the " rutting'''' condition^ or '■'' (xstruationr The peculiar congestion and functional activity of the ovaries at each period of ovulation, act by sympathy upon the other generative organs, and produce in them a greater or less de- gree of excitement, according to the particular species of animal. Almost always there is a certain amount of congestion of the entire generative apparatus ; Fallopian tubes, uterus, vagina, and external organs. The secretions of the vagina and neighboring parts are more particularly affected, being usually increased in quantity and at the same time altered in quality. In the bitch, the vaginal raur cous membrane becomes red and tumefied, and pours out an abun- dant secretion which is often more or less tinged with blood. The secretions acquire also at this time a peculiar odor, which ap- ]iears to attract the male, and to excite in him the sexual impulse. An unusual tumefaction and redness of the vagina and vulva are also very perceptible in the rabbit ; and in some species of apes it has been observed that these periods are accompanied not only by a bloody discharge from the vulva, but also by an engorgement and infiltration of the neighboring parts, extending even to the skin of the buttocks, the thighs, and the under part of the tail.^ The system at large is also visibly affected by the process going on in the ovary. In the cow, for example, the approach of an ' Pouchet, Tlieorie positive de I'ovulation, &c. Paris, 1847, p. 230. 472 OVULATION AND FUNCTION OF MENSTRUATION. oestrual period is marked by an unusual restlessness and agitation, easily recognized by an ordinary observer. The animal partially loses her appetite. She frequently stops browsing, looks about un- easily, perhaps runs from one side of the field to the other, and then recommences feeding, to be disturbed again in a similar manner after a short interval. Her motions are rapid and nervous, and her hide often rough and disordered ; and the whole aspect of the ani- mal indicates the presence of some unusual excitement. After this condition is fully established, the vaginal secretions show them- selves in unusual abundance, and so continue for one or two days; after which the symptoms, both local and general, subside sponta- neously, and the animal returns to her usual condition. It is a remarkable fact, in this connection, that the female of these animals will allow the approaches of the male only during and immediately after the oestrual period ; that is, just when the egg is recently discharged and ready for impregnation. At other times, when sexual intercourse would be necessarily fruitless, the instinct of the animal leads her to avoid it ; and the concourse of the sexes is accordingly made to correspond in time with the maturity of the egg and its aptitude for fecundation. II, MENSTRUATION. In the human female, the return of the periods of ovulation is marked by a peculiar group of phenomena which are known as menstruation, and which are of sufficient importance to be described by themselves. During infancy and childhood the sexual system, as we have mentioned above, is inactive. No discharge of eggs takes place from the ovaries, and no external phenomena show themselves, connected with the reproductive function. At the age of fourteen or fifteen years, however, a change begins to manifest itself. The limbs become rounder, the breasts increase in size, and the entire aspect undergoes a peculiar alteration, which indicates the approaching condition of maturity. At the same time a discharge of blood takes place from the generative passages, accompanied by some disturbance of the general system, and the female is then known to have arrived at the period of puberty. Afterward, the bloody discharge just spoken of returns at regular MENSTRUATION. 473 intervals of four weeks; and, on account of this recurrence corres- ponding with the passage of successive lunar months, its phenomena are designated by the name of the "menses" or the "menstrual periods." The menses return with regularity, from the time of their first appearance, until the age of about forty-five years. During this period, the female is capable of bearing children, and sexual intercourse is liable to be followed by pregnancy. After the forty-fifth year, the periods first become irregular, and then cease altogether; and their final disappearance is an indication that the woman is no longer fertile, and that pregnancy cannot again take place. Even during the period above referred to, from the age of fifteen to forty-five, the regularity and completeness of the menstrual periods indicate to a great extent the aptitude of individual females for impregnation. It is well known that all those causes of ill health which derange menstruation are apt at the same time to interfere with pregnancy ; so that women whose menses are habi- tually regular and natural are much more likely to become preg- nant, after sexual intercourse, than those in whom the periods are absent or irregular. If pregnancy happen to take place, however, at any time during the child-bearing period, the menses are suspended during the con- tinuance of gestation, and usually remain absent after delivery, as long as the woman continues to nurse her child. They then re- commence, and subsequently continue to appear as before. The menstrual discharge consists of an abundant secretion of mucus mingled with blood. When the expected period is about to come on, the female is affected with a certain degree of discomfort and lassitude, a sense of weight in the pelvis, and more or less dis- inclination to society. These symptoms are in some instances slightly pronounced, in others more troublesome. An unusual discharge of vaginal mucus then begins to take place, which soon becomes yellowish or rusty brown in color, from the admixture of a certain proportion of blood; and by the second or third day the discharge has the appearance of nearly pure blood. The unpleasant sensations which were at first manifest then usually subside ; and the discharge, after continuing for a certain period, begins to grow more scanty. Its color changes from a pure red to a brownish or rusty tinge, until it finally disappears altogether, and the female returns to her ordinary condition. The menstrual epochs of the human female correspond with the 474 OVULATION AND FUNCTION OF MENSTEUATION. periods of oestruation in the lower animals. Their general resem- blance to these periods is too evident to require demonstration. Like them, they are absent in the immature female ; and begin to take place only at the period of puberty, when the aptitude for impregnation first commences. Like them, they recur during the child-bearing period at regular intervals ; and are liable to the same interruption bj' pregnancy and lactation. Finally, their dis- appearance corresponds with the cessation of fertility. The periods of oestruation, furthermore, in many of the lower animals, are accompanied, as we have already seen, with an unusual discharge from the generative passages; and this discharge is fre- quently more or less tinged with blood. In the human female the bloody discharge is more abundant than in other instances, but it is evidently a phenomenon differing only in degree from that which shows itself in many species of animals. The most complete evidence, however, that the period of men- struation is in realitv that of ovulation, is derived from the results of direct observation. A sufficient number of instances have now been observed to show that at the menstrual epoch a Graafian vesicle becomes enlarged, ruptures, and discharges its egg. Cruik- shank* noticed such a case so long ago as 1797. Negrier^ relates two instances, communicated to him by Dr. Ollivier d' Angers, in which, after sudden death during menstruation, a bloody and rup- tured Graafian vesicle was found in the ovary. Bischoff^ speaks of four similar cases in his own observation, in three of which the vesicle was just raptured, and in the fourth distended, prominent, and ready to burst. Coste^ has met with several of the same kind. Dr. Michel* found a vesicle ruptured and filled with blood in a woman who was executed for murder while the menses were pre- sent. We have also^ met with the same appearances in a case of death from acute disease, on the second day of menstruation. The process of ovulation, accordingly, in the human female, accompanies and forms a part of that of menstruation. As the menstrual period comes on, a congestion takes place in nearly the ' London Philosophical Transactions, 1797, p. 135. ^ Recherches sur les Ovaires, Paris, 1840, p. 78. '* Annales des Sciences Naturelles, August, 1844. '' Histoire du Developpement des Corps Organises, Paris, 1847, vol i. p. 221. 5 Am. .Tourn. Med. Sci., July, 1848. ^ Corpus Luteum of Menstruation and Pregnancy, in Transactions of American Medical Association, Philadelphia, 1851. MENSTRUATION. 475 whole of the generative apparatus; in the Fallopian tubes and the uterus, as well as in the ovaries and their contents. One of the Graafian follicles is more especially the seat of an unusual vascular excitement. It becomes distended by the fluid which accumulates in its cavity, projects from the surface of the ovary, and is finally ruptured, in the same manner as we have already described this process taking place in the lower animals. It is not quite certain at what particular period of the menstrual flow the rupture of the vesicle and discharge of the egg take place. It is the opinion of Bischoflf, Pouchet, and Kaciborski, that the regular time for this rupture and discharge is not at the commence- ment, but towards the termination of the period. Coste' has ascer- tained, from his observations, that the vesicle ruptures sometimes in the early part of the menstrual epoch, and sometimes later. So far as we can learn, therefore, the precise period of the discharge of the egg is not invariable. Like the menses themselves, it may take place apparently a little earlier or a little later, according to various accidental circumstances; but it always occurs at some time in connection with the menstrual flow, and constitutes the most essential and important part of the catamenial process. The egg, when discharged from the ovary, enters the fimbriated extremity of the Fallopian tube, and commences its passage toward the uterus. The mechanism by which it finds its way into and through the Fallopian tube is different, in the quadrupeds and the human species, and in birds and reptiles. In the latter, the bulk of the egg or mass of eggs discharged is so great as to fill entirely the wide extremity of the oviduct, and they are afterward conveyed down- ward by the peristaltic action of the muscular coat of this canal. In the higher classes, on the contrary, the egg is microscopic in size, and would be liable to be lost, were there not some further provision for its safety. The wide extremity of the Fallopian tube, accordingly, which is here directed toward the ovary, is lined with ciliated epithelium ; and the movement of the cilia, which is directed from the ovary toward the uterus, produces a kind of con- verging stream, or vortex, by which the egg is necessarily drawn toward the narrow portion of the tube, and subsequently conducted to the cavity of the uterus. Accidental causes, however, sometimes disturb this regular course or passage of the egg. The egg may be arrested, for example, ' Loc cit. 476 OVULATION AND FUNCTION OF MENSTRUATION. at the surfiice of the ovary, and so fail to enter the tube at all. If fecundated in this situation, it will then give rise to "ovarian pregnancy." It may escape from the fimbriated extremity into the peritoneal cavity, and form attachments to some one of the neigh- boring organs, causing "abdominal pregnancy;" or finally, it may stop at any part of the Fallopian tube, and so give origin to "tubal pregnancy." The egg, immediately upon its discharge from the ovary, is ready for impregnation. If sexual intercourse happen to take place about that time, the egg and the spermatic fluid meet in some part of the female generative passages, and fecundation is accomplished. It appears, from various observations of Bischoff, Coste, and others, that this contact may take place between the egg and the sperm, either in the uterus or any part of the Fallopian tubes, or even upon the surface of the ovary. If, on the other hand, coitus do not take place, the egg passes down to the uterus unimpregnated, loses its vitality after a short time, and is finally carried away with the uterine secretions. It is easily understood, therefore, why sexual intercourse should be more liable to be followed by pregnancy when it occurs about the menstrual epoch than at other times. This fact, which was long since established as a matter of observation by practical obstetri- cians, depends simply upon the coincidence in time between men- struation and the discharge of the egg. Before its discharge, the egg is immature, and unprepared for impregnation ; and after the menstrual period has passed, it gradually loses its freshness and vitality. The exact length of time, however, preceding and follow- ing the menses, during which impregnation is still possible, has not been ascertained. The spermatic fluid, on the one hand, retains its vitality for an unknown period after coition, and the egg for an unknown period after its discharge. Both these occurrences may, therefore, either precede or follow each other within certain limits, and impregnation be still possible ; but the precise extent of these limits is still uncertain, and is probably more or less variable in diffent individuals. The above facts indicate also the true explanation of certain exceptional cases, which have sometimes been observed, in which fertility exists without menstruation. Various authors (Churchill, Reid, Velpeau, &c.) have related instances of fruitful women in whom the menses were very scanty and irregular, or even entirely absent. The menstrual flow is, in fact, only the external sign and accompa- MENSTRUATION". 477 niment of a more important process taking place within. It is habitual]}'' scanty in some individuals, and abundant in others. Such variations depend upon the condition of vascular activity of the system at large, or of the uterine organs in particular; and though the bloody discharge is usually an index of the general aptitude of these organs for successful impregnation, it is not an absolute or necessary requisite. Provided a mature egg be dis- charged from the ovary at the appointed period, menstruation pro- perly speaking exists, and pregnancy is possible. The blood which escapes during the menstrual flow is supplied by the uterine mucous membrane. If the cavity of the uterus be examined after death during menstruation, its internal surface is seen to be smeared with a thickish bloody fluid, which may be traced through the uterine cervix and into the vagina. The Fallo- pian tubes themselves are sometimes found excessively congested, and filled with a similar bloody discharge. The menstrual blood has also been seen to exude from the uterine orifice in cases of pro- cidentia uteri, as well as in the natural condition by examination ■with the vaginal speculum. It is discharged by a kind of capillary hemorrhage, similar to that which takes place from the lungs in cases of hemoptysis, only less sudden and violent. The blood does not form any visible coagulum, owing to its being gradually exuded from many minute points, and mingled with a large quantity of mucus. When poured out, however, more rapidly or in larger quantity than usual, as in cases of menorrhagia, the menstrual blood coagulates in the same manner as if derived from any other source. The hemorrhage which supplies it comes from the whole extent of the mucous membrane of the body of the uterus, and is, at the same time, the consequence and the natural termination of the periodical congestion of the parts. 478 MENSTRUATION AND PREGNANCY. CHAPTER VI. ON THE CORPUS LUTEUM OF MENSTRUATION AND PREGNANCY. After the rupture of the Graafian vesicle at the menstrual period, a bloody cavity is left in the ovary which is subsequently obliterated by a kind of granulating process, somewhat similar in character to the healing of an abscess. For the Graafian vesicle is intended simply for the formation and growth of the egg. After the egg therefore has arrived at maturity and has been dis- charged, the Graafian follicle has no longer any function to per- form. It then only remains for it to pass through a process of obliteration and atrophy, as an organ which has become useless and obsolete. While undergoing this process, the Graafian vesicle is at one time converted into a peculiar, solid, globular body, which is called the corpus luteum ; a name given to it on account of the yellow color which it acquires at a certain period of its formation. We shall proceed to describe the corpus luteum in the human species; first, as it follows the ordinary course of development after menstruation ; and secondly, as it is modified in its growth and appearance during the existence of pregnancy. I. CORPUS luteum of menstruation. We have already described, in the preceding chapter, the man- ner in which a Graafian vesicle, at each menstrual epoch, swells, protrudes from the surface of the ovary, and at last ruptures and discharges its egg. At the moment of rupture, or immediately after it, an abundant hemorrhage takes place in the human sub- ject from the vessels of the follicle, by which its cavity is filled with blood. This blood coagulates soon after its exudation, as it would do if extravasated in any other part of the body, and the coagulum is retained in the interior of the Graafian follicle. CORPUS LUTEUM OF MENSTRUATION. 479 & - h Graafian Follicle receully ruplured during nienstniation, and filled with a bloody coaguliim ; shown in longitudinal sec- tion. — a. Tissue of the ovary, b. Membrane of the vesicle, c. Point of rupture. The opening by which the Qgg makes its escape is usually not an extensive laceration, but a minute rounded perforation, often not more than half a line in diameter. A small probe, introduced through this opening, passes directly into the cavity of the follicle. If the Graafian follicle be opened at this time by a longitudinal inci- sion (Fig. 170), it will be seen to form a globu- lar cavity, one-half to three-quarters of an inch in diameter, containing a soft, recent, dark colored coagulum. This coagulum has no organic connection with the walls of the follicle, but lies loose in its cavity and may be easily turned out with the handle of a knife. There is sometimes a slight mechanical adhe- sion of the clot to the eds^es of the lacerated opening, just as the coagulum in a recently ligatured artery is entangled by the divided edges of the internal and middle coats; but there is no continuity of substance between them, and the clot may be everywhere readily separated by careful manipulation. The membrane of the vesicle presents at this time a smooth, transparent, and vascular internal surface, without any alteration of color, consistency, or texture. An important change, however, soon begins to take place, both in the central coagulum and in the membrane of the vesicle. The clot, which is at first large, soft, and gelatinous, like any other mass of coagulated blood, begins to contract; and the serum separates from the coagulum proper. The serum, as fast as it separates from the coagulum, is absorbed by the neighboring parts; and the clot, accordingly, grows every day smaller and denser than before. At the same time the coloring matter of the blood under- goes the changes which usually take place in it after extravasation, and is partially reabsorbed together with the serum. This second change is somewhat less rapid than the former, but still a diminu- tion of color is very perceptible in the clot, at the expiration of two weeks. The membrane of the vesicle during this time is beginning to undergo a process of hypertrophy or development, by which it becomes thickened and convoluted, and tends to fill up partially the cavity of the follicle. This hypertrophy and convolution of the membrane just named commences and proceeds most rapidly 480 MENSTRUATION AND PREGNANCY. Fig. 171. at the deeper part of the follicle, directly opposite the situation of the superficial rupture. From this point it gradually becomes thinner and less convoluted as it approaches the surface of the ovary and the edges of the ruptured orifice. At the end of three weeks, this hypertrophy of the membrane of the vesicle has reached its maximum. The ruptured Graafian fol- licle has now become so completely solidified by the new growth above described, and by the condensation of its clot, that it receives the name of the corpus luteum. It forms a perceptible prominence upon the surface of the ovary, and may be felt between the fingers as a well-defined rounded tumor, which is nearly always somewhat flattened from side to side. It measures about three-quarters of an inch in length and half an inch in depth. On its surface may be seen a minute cicatrix of the peritoneum, occupying the spot of the original rupture. On cutting it open at this time (Fig. 171), the corpus luteum is seen to con- sist, as above described, of a central coagulum and a convoluted wall. The coagulum is semi-transparent, of a gray or light greenish color, more or less mottled with red. The con- voluted wall is about one-eighth of an inch thick at its deepest part, and of an indefinite yellowish or rosy hue, not very different in tinge from the rest of the ovarian tissue. The convoluted wall and the con- tained clot lie simply in contact with each other, as at first, without any intervening membrane or other organic connection ; and they may still be readily separated from each other by the handle of a knife or the flattened end of a probe. The corpus luteum at this time may also be stripped out, or enucleated entire, from the ovarian tissue, just as might have been done with the Graafian follicle pre- viously to its rupture. When enucleated in this way, the corpus luteum presents itself under the form of a solid globular or flat- tened tumor, with convolutions upon it somewhat similar in ap- pearance to those of the brain, and covered with the remains of the areolar tissue, by which it was previously connected with the substance of the ovary. . . Ovary cut open, showing corpus luteum divided longitudinally; three weeks after menstruation. From a girl dead of hsemoptysis. COEPUS LUTEUM OF MENSTRUATION. 481 Fig. 172. Ovary, showing corpus luteum four weeks after men- struation ; from a woman dead of apoplexy. After the third week from the close of menstruation, the corpus luteum passes into a retrograde condition. It diminishes percep- tibly in size, and the central coagulum continues to be absorbed and loses still farther its coloring matter. The whole body under- goes a process of partial atrophy ; and at the end of the fourth week it is not more than three-eighths of an inch in its longest diameter. (Fig. 172.) The external cicatrix may still usually be seea, as well as the point where the central coagulum comes in contact with the peritoneum. There is still no organic connection between the central coagulum and the convoluted wall ; but the partial condensation of the clot and the continued folding of the wall prevent the separation of the two being so easily accom- plished as before, though it may still be eflfected by careful management. The entire corpus luteum may also still be extracted from its bed in the ovarian tissue. The color of the convoluted wall, during the early part of this retrograde stage, instead of fading, like that of the fibrinous coagu- lum, becomes more strongly marked. From having a dull yellowish or rosy hue, as at first, it gradually assumes a brighter and more decided yellow. This change of color in the convoluted wall is produced in consequence of a kind of fatty degeneration which takes place in its texture ; a large quantity of oil-globules being deposited in it at this time, as may be readily recognized under the microscope. At the end of the fourth week, this alteration in hue is complete ; and the outer wall of the corpus luteum is then of a clear chrome-yellow color, by which it is readily distinguished from all the neighboring tissues. After this period, the process of atrophy and degeneration goes on rapidly. The clot becomes constantly more dense and shrivelled, and is soon converted into a minute, stellate, white, or reddish white cicatrix. The yellow wall becomes softer and more friable, as is the case with all 31 Fig. 173. ,4:^ OvART, showing corpus lu- teum, nine weeks after menstrua- tion ; from a girl dead of tuber- cular meningitis. 482 MENSTEUATION AND PEEGNANCY. tissues undergoing fatty degeneration, and shows less distinctly the markings of its convolutions. At the same time, its edges become confounded with the central coagulum on the one hand, and the neighboring tissues on the other, so that it is no longer possible to separate them fairly from each other. At the end of eight or nine weeks the whole body is reduced to the condition of an insignificant, yellowish, cicatrix-like spot, measuring less than a quarter of an inch in its longest diameter, in which the original texture of the corpus luteum can be recognized only by the pecu- liar folding and coloring of its constituent parts. Subsequently its atrophy goes on in a less active manner, and a period of seven or eight months sometimes elapses before its final and complete dis- appearance. The corpus luteum, accordingly, is a formation which results from the filling up and obliteration of a ruptured Graafian follicle. Under ordinary conditions, a corpus luteum is produced at every menstrual period ; and notwithstanding the rapidity with which it retrogrades and becomes atrophied, a new one is always formed before its predecessor has completely disappeared. When, therefore, we examine the ovaries of a healthy female, in whom the menses have recvirred with regularity for some time previous to death, several corpora lutea will be met with in different stages of formation and atrophy. Thus we have found, under such circumstances, four, five, six, and even eight corpora lutea in the ovaries at the same time, perfectly distinguishable by their texture, but very small, and most of them evidently in a state of advanced retrogression. They finally disappear altogether, and the number of those present in the ovary, therefore, no longer corresponds with that of the Graafian follicles which have been ruptured. II. CORPUS LUTEUM OF PREGNANCY. Since the process above described takes place at every menstrual period, it is independent of impregnation and even of sexual inter- course. The mere presence of a corpus luteum, therefore, is no indication that pregnancy has existed, but only that a Graafian follicle has been ruptured, and its contents discharged. We find, nevertheless, that when pregnancy does take place, the appearance of the corpus luteum becomes so much altered as to be readily dis- tinguished from that which simply follows the ordinary menstrual CORPUS LUTEUM OF PREGNANCY. 483 process. It is proper, therefore, to speak of two kinds of corpora lutea; one belonging to menstruation, the other to pregnancy. The difference between these two kinds of corpora lutea is not an essential or fundamental difference ; since they both originate in the same way, and are composed of the same structures. It is, properly speaking, only a difference in the degree and rapidity of their development. For while the corpus luteum of menstrua- tion passes rapidly through its different stages, and is very soon reduced to a condition of atrophy, that of pregnancy continues its development for a long time, attains a larger size and firmer organ- ization, and disappears finally only at a much later period. This variation in the development and history of the corpus luteum depends upon the unusually active condition of the pregnant uterus. This organ exerts a powerful sympathetic action, during pregnancy, upon many other parts of the system. The stomach becomes irritable, the appetite capricious, and even the mental faculties and the moral disposition are frequently more or less affected. The ovaries, however, feel the disturbing influences of gestation more certainly and decidedly than the other organs, since they are more closely connected with the uterus in the ordinary performance of their function. The moment that pregnancy takes place, the process of menstruation is arrested. No more eggs come to maturity and no more Graafian follicles are ruptured, during the whole period of gestation. It is not at all singular, therefore, that the growth of the corpus luteum should also be modified, by an influence which affects so profoundly the system at large, as well as the ovaries in particular. During the first three weeks of its formation, the growth of the corpus luteum is the same, in the impregnated, as in the unimpreg- nated condition. After that time, however, a difference becomes manifest. Instead of commencing a retrograde course during the fourth week, the corpus luteum of pregnancy continues its deve- lopment. The external wall grows thicker, and its convolutions more abundant. Its color alters in the same way as previously described, and becomes of a bright yellow by the deposit of fatty matter in microscopic globules and granules. By the end of the second month, the whole corpus luteum has in- creased in size to such an extent as to measure seven-eighths of an inch in length by half an inch in depth. (Fig. 174.) The central coagulum has by this time become almost entirely decolorized, so as 484 MENSTRUATION AND PREGNANCY. to present the appearance of a purely fibrinous deposit. Sometimes we find that a part of the serum, during its separation from the clot, has accumulated in the centre of the mass, as in Fig. 174, forming a little cavity containing a few drops of clear fluid and inclosed by a whitish, fibrinous layer, the remains of the solid portion of the clot. It is this fibrinous layer ^ig- l'''^- which has sometimes been mistaken for a distinct or- ganized membrane, lining the internal surface of the convoluted wall, and which has thus led to the belief that the yellow matter of the corpus luteum is nor- mally deposited outside the membrane of the Graafian follicle. Such,- however, is not its real structure. The convoluted wall of the corpus luteum is the membrane of the follicle itself, partially altered by hyper- trophy, as may be readily seen by examination in the earlier stages of its growth ; and the fibrinous layer, situated internally, is the original bloody coagulum, decolorized and condensed by continued absorption. The existence of a central cavity, containing serous fluid, is merely an occasional, not a constant phenomenon. More frequently, the fibrinous clot is solid throughout, the serum being gradually absorbed, as it separates spontaneously from the coagulum. During the third and fourth CoRPns Ll'TECM of pi'egnancy, at end of second mouth : from a woman dead from induced abortion. Fig. 175. Corpus Luteum of pregnancy, at end ■>( fourth month ; from a woman dead by poison. months, the enlargement of the corpus luteum continues; so that at the end of that time it may measure seven-eighths of an inch in length by three- quarters of an inch in depth. (Fig. 175.) The convoluted wall is still thicker and more highly developed than before, having a thickness, at its deep- est part, of three sixteenths of an inch. Its color, however, has already begun to fade, and is COKPUS LUTEUM OF PEEGNANCY. 485 Fig. 176. now of a dull yellow, instead of the bright, clear tinge which it previously exhibited. The central coagulum, perfectly colorless and fibrinous in appearance, is often so much flattened out, by the lateral compression of its mass, that it has hardly a line in thickness. The other relations of the different parts of the corpus luteum remain the same. The corpus luteum has now attained its maximum of develop- ment, and remains without any very perceptible alteration during the fifth and sixth months. It then begins to retrograde, diminish- ing constantly in size during the seventh and eighth months. Its external wall fades still more perceptibly in color, becoming of a faint yellowish white, not unlike that which it presented at the end of the third week. Its texture is thick, soft, and elastic, and it is still strongly convoluted. An abundance of fine red vessels can be seen penetrating from the exterior into the interstices of its con- volutions. The central coagulum is reduced by this time to the condition of a whitish, radiated cicatrix. The atrophy of the organ continues during the ninth month. At the termination of pregnancy, it is re- duced to the size of half an inch in length and three-eighths of an inch in depth. (Fig. 176.) It is then of a faint indefinite hue, but little contrasted with the remain- ing tissues of the ovary. The central cica- trix has become very small, and appears only as a thin whitish lamina with radiating processes which run in between the inter- stices of the convolutions. The whole mass, however, is still quite firm and resisting to the touch, and is readily distinguishable, both from its size and texture, as a pro- minent feature in the ovarian tissue, and a reliable indication of pregnancy. The con- voluted structure of its external wall is very perceptible, and the point of rupture, with its external peritoneal cicatrix, distinctly visible. After delivery, the corpus luteum retrogrades rapidly. At the end of eight or nine weeks, it has become so much altered that its color is no longer distinguishable, and only faint traces of its con- voluted structure are to be discovered by close examination. These Corpus Lpteum of preg- nancy, at term ; from a woman dead in delivery from rupture of the uterus. 486 MENSTRUATION AND PREGNANCY. traces may remain, however, for a long time afterward, more or less concealed in the ovarian tissue. We have distinguished them so late as nine and a half months after delivery. They finally disap- pear entirely, together with the external cicatrix which previously marked their situation. During the existence of gestation, the process of menstruation being suspended, no new follicles are ruptured, and no new corpora lutea produced ; and as the old ones, formed before the period of conception, gradually fade and disappear, the corpus luteum which marks the occurrence of pregnancy after a short time exists alone in the ovary, and is not accompanied by any others of older date. In twin pregnancies, we of course find two corporea lutea in the ovaries ; but these are precisely similar to each other, and, being evidently of the same date, will not give rise to any confusion. Where there is but a single foetus in the uterus, and the ovaries contain two corpora lutea of similar appearance, one of them belongs to an embryo which has been blighted by some accident in the early part of pregnancy. The remains of the blighted em- bryo may often be discovered, in such cases, in some part of the Fallopian tubes, where it has been arrested in its descent toward the uterus. After the process of lactation comes to an end, the ovaries again resume their ordinary function. The Graafian follicles mature and rupture in succession, as before, and new corpora lutea follow each other in alternate development and disappearance. We find, then, that the corpus luteum of menstruation differs from that of pregnancy in the extent of its development and the dura- tion of its existence. While the former passes through all the im- portant phases of its growth and decline in the period of two months, the latter lasts for from nine to ten months, and presents, during a great portion of the time, a larger size and a more solid organization. It will be observed that, even with the corpus luteum of pregnancy, the bright yellow color, which is so important a cha- racteristic, is only temporary in its duration ; not making its appear- ance till about the end of the fourth week, and disappearing after the sixth month. The following table contains, in a brief form, the characters of the corpus luteum, as belonging to the two different conditions of menstruation and pregnancy, corresponding with different periods of its development. CORPUS LUTEUM OF PREGNANCY. 487 At the end of three weeks One month Two months Six months Nine months CoKPus Ldteum of Menstruation. Corpus Luteum of Pregnancy. Three-quarters of an inch in diameter ; central clot reddish ; con- voluted wall pale. Smaller; convoluted wall bright yellow ; clot still reddish. Reduced to the condition of an insignificant cicatrix. Absent. Absent. Larger; convoluted wall bright yellow ; clot still reddish. Seven-eighths of an inch in dia- meter; convoluted wall bright yellow; clot perfectly decolor- ized. Still as large as at end of second month; clot fibrinous; convo- luted wall paler. One-half an inch in diameter ; central clot converted into a radiating cicatrix ; the external wall tolerably thick and convo- luted, but without any bright yellow color. 488 DEVELOPMENT OF THE IMPREGNATED EGG. CHAPTER VII. ON THE DEVELOPMENT OF THE IMPREGNATED EGG —SEGMENTATION OF THE VI T EL L US— BL A STODER- MIC MEMBRANE — FORMATION OF ORGANS IN THE FROG. We have seen, in the foregoing chapters, how the egg, produced in the ovarian follicle, becomes gradually developed and ripened, until it is ready to be discharged. The egg, accordingly, passes through several successive stages of formation, even while still con- tained within the ovary; and its vitellus becomes gradually com- pleted, by the formation of albuminous material and the deposit of molecular granulations. The last change which the egg undergoes, in this situation, and which marks its complete maturity, is the dis- appearance of the germinative vesicle. This vesicle, which is, in general, a prominent feature of the ovarian egg, disappears but a short time previous to its discharge, or even just at the period of its leaving the Graafian follicle. The egg, therefore, consisting simply of the mature vitellus and the vitelline membrane, comes in contact, after leaving the ovary, and while passing through the Fallopian tube, with the spermatic fluid, and thereby becomes fecundated. By the influence of fecun- dation, a new stimulus is imparted to its growth ; and while the vitality of the unimpregnated germ, arrived at this point, would have reached its termination, the fecundated egg, on the contrary, starts upon a new and more extensive course of development, by which it is finally converted into the body of the young animal. The egg, in the first place, as it passes down the Fallopian tube, becomes covered with an albuminous secretion. In the birds, as we have seen, this secretion is very abundant, and is deposited in suc- cessive layers around the vitellus. In the reptiles, it is also poured out in considerable quantity, and serves for the nourishment of the egg during its early growth. In quadrupeds, the albuminous matter is supplied in the same way, though in smaller quantity, by the SEGMENTATION OF THE VITELLUS. 489 mucous membrane of the Fallopian tubes, and envelopes the egg in a layer of nutritious material. A very remarkable change now takes place in the impregnated egg, which is known as the spontaneous division, or segyneiitation, of the vitellus. A furrow first shows itself, running round the globular mass of the vitellus in a vertical direction, which gradually deepens until it has divided the vitellus into two separate halves or hemispheres. (Fig. 177, a.) Almost at the same time another furrow, running at right angles with the first, penetrates also the substance of the vitellus and cuts it in a transverse direction. The vitellus is thus divided into four equal portions (Fig. 177, b), the edges and angles of which are rounded off, and which are still contained in the cavity of the vitelline membrane. The spaces between them and the internal surface of the vitelline membrane are occu- pied by a transparent fluid. The process thus commenced goes on by a successive formation of fur- rows and sections, in various direc- tions. The four vitelline segments already produced are thus subdivided into sixteen, the sixteen into sixty- four, and so on; until the whole vi- tellus is converted into a mulberry shaped mass, composed of minute, nearly spherical bodies, which are called the "vitelline spheres." (Fig. 177, c.) These vitelline spheres have a somewhat firmer consistency than the original substance of the vitellus; and this consistency appears to in- crease, as they successively multiply in numbers and diminish in size. At last they have become so abundant as to be closely crowded together, compressed into poly- gonal forms, and flattened against the internal surface of the vitel- Segmextation of the Vitellus. 490 DEVELOPMENT OF THE IMPREGNATED EGG. line membrane. (Fig. 177, d) They have by this time been con- verted into true animal cells; and these cells, adhering to each other by their adjacent edges, form a continuous organized membrane, which is termed the Blastodermic mewhrane. During the formation of this membrane, moreover, the egg, while passing through the Fallopian tubes into the uterus, has increased in size. The albuminous matter with which it was enveloped has liquefied; and, being absorbed by endosmosis through the vitelline membrane, has furnished the materials for the more solid and ex- tensive growth of the newly-formed structures. It may also be seen that a large quantity of this fluid has accumulated in the central cavity of the egg^ inclosed accordingly by the blastodermic membrane, with the original vitelline membrane still forming an external envelope round the whole. The next change which takes place, consists in the division or splitting of the blastodermic membrane into two layers, which are known as the external and internal layers of the blastodermic membrane. They are both still composed exclusively of cells ; but those of the external layer are usually smaller and more compact, while those of the internal are rather larger and looser in texture. The egg ihen presents the appearance of a globular sac, the walls of which consist of three concentric layers, lying in contact with and inclos- ing each other, viz., 1st, the structureless vitelline membrane on the outside ; 2d, the external layer of the blastodermic membrane, com- posed of cells ; and 3d, the internal layer of the blastodermic mem- brane, also composed of cells. The cavity of the egg is occupied by a transparent fluid, as above mentioned. This entire process of the segmentation of the vitellus and the formation of the blastodermic membrane is one of the most re- markable and important of all the changes which take place during the development of the egg. It is by this process that the simple globular mass of the vitellus, composed of an albuminous matter and oily granules, is converted into an organized structure. For the blastodermic membrane, though consisting only of cells nearly uniform in size and shape, is nevertheless a truly organized mem- brane, made up of fully formed anatomical elements. It is, more- over, the first sign of distinct organization which makes its appear- ance in the egg; and as soon as it is completed, the body of the new foetus is formed. The blastodermic membrane is, in fact, the body of the foetus. It is at this time, it is true, exceedingly simple in texture ; but we shall see hereafter that all the future organs BLASTODERMIC MEMBRANE. 491 of the body, however varied and complicated in structure, arise out of it, by modification and development of its different parts. The segmentation of the vitellus, moreover, and the formation of the blastodermic membrane, take place in essentially the same manner in all the different classes of animals. It is always in this way that the egg commences its development, whether it be des- tined to form afterward a fish or a reptile, a bird, a quadruped or a man. The peculiarities belonging to different species show them- selves afterward, by variations in the manner and extent of the development of different parts. In the higher animals and in the human subject the development of the egg becomes an exceedingly complicated process, owing to the formation of various accessory organs, which are made requisite by the peculiar conditions under which the development of the embryo takes place. It is, in fact, impossible to describe or understand properly the complex embry- ology of the quadrupeds, and more particularly that of the human subject, without first tracing the development of those species in which the process is more simple. We shall commence our descrip- tion, therefore, with the development of the egg of the frog, which is for many reasons particularly appropriate for our purpose. The egg of the frog, when discharged from the body of the female and fecundated by the spermatic fluid of the male, is deposited in the water, enveloped in a soft elastic cushion of albuminous sub- stance. It is therefore in a situation where it is freely exposed to the light, the air, and the moderate warmth of the sun's rays, and where it can absorb directly an abundance of moisture and of ap- propriate nutritious material. We find accordingly that under these circumstances the development of the egg is distinguished by a character of great simplicity ; since the whole of the vitellus is directly converted into the body of the embryo. There are no accessory organs required, and consequently no complication of the formative process. The two layers of the blastodermic membrane, above described, represent together the commencement of all the organs of the foetus. They are intended, however, for the production of two different systems ; and the entire process of their development may be ex- pressed as follows : The external layer of the blastodermic membrane produces the spinal column and all the organs of animal life; while the internal layer produces the intestinal canal, and all the organs of vege- tative life. The first sign of advancing organization in the external layer of 492 DEVELOPMENT OF THE IMPREGNATED EGG. the blastodermic membrane shows itself in a thickening and con- densation of its structure. This thickened portion has the form of an elongated oval-shaped spot, termed the "embryonic spot" (Fig. 178), the wide edges of which are somewhat more opaque than the rest of the blasto- dermic membrane. Inclosed within these opaque edges is a narrower color- less and transparent space, the "area pellucida," and in its centre is a delicate line, or furrow, running longitudinally from front to rear, which is called the " primitive trace." On each side of the primitive trace, 1.VPREGXATEI, E G r, , with com- ^^ thc arca pellucida, the substance of mencement of forniaiion of embryo: |;}]e blastodermic membrane riscs up in shoTving erabrvonic spot, area pellu- „ , cida, and primuive trace. such a manner as to torm two nearly parallel vertical plates or ridges, which approach each other over the dorsal aspect of the fcetus and are therefore called the "dorsal plates." They at last meet on the median line, so as to inclose the furrow above described and con- vert it into a canal. This afterward becomes the spinal canal, and in its cavity is formed the spinal cord, by a deposit of nervous matter upon its internal surface. At the anterior extremity of this canal, its cavity is large and rounded, to accommodate the brain and medulla oblongata; at its posterior extremity it is narrow and pointed, and contains the extremity of the spinal cord. In a transverse section of the egg at this stage (Fig. 179), the dorsal plates may be seen approaching each other above, on each side of the primitive furrow or "trace." At a more advanced period (Fig. 180) they may be seen fairly united with each other, so as to inclose the cavity of the spinal canal. At the same time, the edges of the thickened portion of the blastodermic membrane grow outward and downward, so as to spread out more and more over the lateral portions of the vitelline mass. These are called the "abdominal plates;" and as they increase in extent they tend to unite with each other below and inclose the abdominal cavity, just as the dorsal plates unite above, and inclose the spinal canal. At last the abdominal plates actually do unite with each other on the median line (at i. Fig. 180), embracing of course the whole internal layer of the blastodermic membrane (5), which incloses in FORMATION OF ORGANS. 498 its turn the remains of the original vitellus and the albuminous fluid which has accumulated in its cavity. Fig. 179. Fig. 180. Transverse section of Egg in an early stage of development — 1. External layer of blastodermic membrane. 2,2. Dorsal plates. 3. Internal layer of blastodermic membrane. 1 .MPREG N.A.TED EoQ, at a SOmeTvhHt more advanced period. — 1. Umbilicus, or point of union between abdominal plates. 2, 2. Dorsal plates united -with each other on the median line and inclosing the spinal canal. 3, 3. Abdominal plates. 4. Sec- tion of spinal column, with laminae and ribs. 5. Internal layer of blastodermic membrane. During this time, there is formed, in the thickness of the external blastodermic layer, immediately beneath the spinal canal, a longitu- dinal cartilaginous cord, called the "chorda dorsalis." Around the chorda dorsalis are afterward developed the bodies of the vertebrae (Fig. 180, 4), forming the chain of the vertebral column; and the oblique processes of the vertebrae run upward from this point into the dorsal plates ; while the transverse processes, and ribs, run outward and downward in the abdominal plates, to encircle more or less completely the corresponding portion of the body. If we now examine the egg in longitudinal section, while this process is going on, the thickened portion of the external blastoder mic layer may be seen in profile, as at i, Fig. 181. The anterior portion (2), which will form the head, is thicker than the posterior (3), Avhich will form the tail of the young animal. As the whole mass grows rapidly, both in the anterior and posterior direction, the head becomes very thick and voluminous, while the tail also begins to project backward, and the whole egg assumes a distinctly elongated form. (Fig. 182.) The abdominal plates at the same time- meet upon its under surface, and the point at which they finally 494 DEYELOPMENT OF THE IMPREGNATED EGG. unite forms the abdominal cicatrix or umbilicus. The internal blas- todermic layer is seen, of course, in the longitudinal section of the Fie. 181. Fig. 182. Diagram of Frog's Egg, in an parly E(jg of Frog, in process of develop- Rfage of development ; longitudinal sec- meui. tiou. — 1 Thickened portion of external blastodermic layer, forming body of foetus. 2. Anterior extremity of foetus. .3. Poste- rior extremity. 4. Internal layer of blas- todermic membrane. 5. Cavity of vitellus. egg, as well as in the transverse, embraced by the abdominal plates, and inclosing, as before, the renaains of the vitellus. As the development of the above parts goes on (Fig. 183), the head becomes still larger, and soon shows traces of the formation Fig. 183. OF Frog, farther advanced. of organs of special sense. The tail also increases in size, and pro- jects farther from the posterior extremity of the embryo. The spinal cord runs in a longitudinal direction from front to rear, and its anterior extremity enlarges, so as to form the brain and medulla oblongata. In the mean time, the internal blastodermic layer, which is subsequently to be converted into the intestinal canal, has been shut in by the abdominal walls, and still forms a perfectly closed sac, of a slightly elongated figure, without either inlet or outlet. Afterward, the mouth is formed by a process of atrophy and per- ibration, which takes place through both external and internal layers, at the anterior extremity, while a similar perforation, at the poste- rior extremity, results in the formation of the anus. FORMATION OF ORGANS. 495 All these parts, however, are as yet imperfect; and, being merely in the course of formation, are incapable of performing any active function. By a continuation of the same process, the different portions of the external blastodermic layer are further developed, so as to re- sult in the complete formation of the various parts of the skeleton, the integument, the organs of special sense, and the voluntary nerves and muscles. The tail at the same time acquires sufficient size and strength to be capable of acting as an organ of locomo- tion. (Fig. 184.) The intestinal canal, which has been formed from Fig. 184. T A B p o L E fully developed. the internal blastodermic layer, is at first a short, wide, and nearly straight tube, running directly from the mouth to the anus. It soon, however, begins to grow faster than the abdominal cavity which incloses it, becoming longer and narrower, and is at the same time thrown into numerous convolutions. It thus presents a larger internal surface for the performance of the digestive process. Arrived at this period, the young tadpole ruptures the vitelline membrane, by which he has heretofore been inclosed, and leaves the cavity of the egg. He at first fastens himself upon the remains of the albuminous matter deposited round the egg, and feeds upon it for a short period. He soon, however, acquires sufficient strength and activity to swim about freely in search of other food, propelling himself by means of his large, membranous, and muscular tail. The alimentary canal increases very rapidly in length and becomes spirally coiled up in the abdominal cavity, so as to attain a length from seven to eight times greater than that of the entire body. After a time, a change takes place in the external form of the young animal. Anterior and posterior extremities or limbs begin to show themselves, by budding or sprouting from the corresponding regions of the body. (Fig. 185.) At first these organs are very small, imperfect in structure, and altogether useless for purposes of 496 DEVELOPMENT OF THE IMPREGNATED EGG. locomotion. Thej soon, however, increase in size and strength ; and while thej keep pace with the increasing development of the whole body, the tail on the contrary ceases to grow, and becomes shrivelled and atrophied. The limbs, in fact, are destined finally to replace the tail as organs of locomotion ; and a time at last arrives (Fig. 186) when the tail has altogether disappeared, while Fig. 185. Fisj. 186. Tadpole, with limbs begiuoiug to be formed. Perfect Frog. the legs have become fully developed, muscular and powerful. Then the animal, which was before confined to an aquatic mode of life, becomes capable of living also upon land, and a trans- formation is effected from the tadpole into the perfect frog. During the same time, other changes of an equally important character have taken place in the internal organs. The tadpole at first breathes by gills; but these organs subsequently become atrophied and disappear, being finally replaced by well developed lungs. The structure of the mouth, also, of the integument, and of the circulatory system, is altered to correspond with the varying conditions and wants of the growing animal ; and all these changes, taking place in part successively and in part simultaneously, bring the animal at last to a state of complete formation. The process of development may then be briefly recapitulated as follows: — 1. The blastodermic membrane, produced by the segmentation of the vitellus, consists of two cellular layers, viz., an external and an internal blastodermic layer. 2. The external layer of the blastodermic membrane incloses by its dorsal plates the cerebro-spinal canal, and by its abdominal plates the abdominal or visceral cavity. FORMATION OF ORGANS IN THE FROG. 497 3. The internal layer of the blastodermic membrane forms the intestinal canal, which becomes lengthened and convoluted, and communicates with the exterior by a mouth and anus of secondary formation. 4. Finally the cerebro-spinal axis and its nerves, the skeleton, the organs of special sense, the integument, and the muscles, are developed from the external blastodermic layer ; while the anterior and posterior extremities are formed from the same layer by a pro- cess of sprouting, or continuous growth. 32 498 THE UMBILICAL VESICLE. CHAPTER VIII. THE UMBILICAL VESICLE, Fig. 187. In" the frog, as we have seen, the abdominal plates, closing together in front and underneath the body of the animal, shut in directly the whole of the vitellus, and join each other upon the median line, at the umbilicus. The whole remains of the vitellus are then inclosed in the abdomen of the animal, and in the intestinal sac formed by the internal blastodermic layer. In many instances, however, as, for example, in several kinds of fish, and in all the birds and quadrupeds, the abdominal plates do not immediately embrace the whole of the vitelline mass, but tend to close together about its middle ; so that the vitellus is constricted, in this way, and divided into two portions: one internal, and one external. (Fig. 187.) As the process of development proceeds, the body of the foetus increases in size, out of pro- portion to the vitelline sac, and the con- striction just mentioned becomes at the same time more strongly marked ; so that the separation between the internal and external portions of the vitelline sac is nearly complete. (Fig. 188.) The internal layer of the blastodermic membrane is by the same means divided into two portions, one of which forms the intestinal canal, while the other, remaining outside, forms a sac-like appendage to the abdo- men, which is known by the name of the umbilical vesicle. The umbilical vesicle is accordingly lined by a portion of the internal blastodermic layer, continuous with the mucous membrane of the intestinal canal; while it is covered on the outside by a por- tion of the external blastodermic layer, continuous with the integu- ment of the abdomen. EfiQOF Fish; showing forma tioa of umbilical vesicle. THE UMBILICAL VESICLE. 499 After the young animal leaves the egg, the umbilical vesicle sometimes becomes withered and atrophied by the absorption of its contents; while in some instances, the abdominal walls gradually Fig. 188. Young Fish with umbilical vesicle. extend over it, and crowd it back into the abdomen ; the nutritious matter which it contained passing from the cavity of the vesicle into that of the intestine by the narrow passage or canal which remains open between them. In the human subject, however, as well as in the quadrupeds, the umbilical vesicle becomes more completely separated from the abdo men than in the cases just mentioned. There is at first a wide com- munication between the cavity of the umbilical vesicle and that of the intestine ; and this communication, as in other instances, becomes gradually narrowed by the increasing constriction of the abdominal walls. Here, however, the constriction proceeds so far that the opposite surfaces of the canal come in contact with each other, and adhere; so that the narrow passage previously existing between the cavity of the intestine and that of the umbilical vesicle is obliterated, and the vesicle is then connected with the abdomen only by an impervious cord. This cord afterward elongates, and becomes con- verted into a slender, thread-like pedicle (Fig. 189), passing out from the abdomen of the foetus, and connected by its further extremity with the umbilical vesicle, which is filled with a transparent, colorless fluid. The umbilical human embryo, with .,. T- 1 ••II- 11 umbilical vesicle ; about the vesicle is very distinctly visible m the human gfth week. foetus so late as the end of the third month. After that period it diminishes in size, and is gradually lost in the advancing development of the neighboring parts. In the formation of the umbilical vesicle, we have the first varia- 500 THE UMBILICAL VESICLE. tion from the simple plan of development described in the preceding chapter. Here, the whole of the vitellus is not directly converted into the body of the embryo ; but while a part of it is taken, as usual, into the abdominal cavity, and used immediately for the pur- poses of nutrition, a part is left outside the abdomen, in the umbilical vesicle, a kind of secondary organ or appendage of the foetus. The contents of the umbilical vesicle, however, are afterward absorbed, and so appropriated, finally, to the nourishment of the newly formed tissues AMNION AND ALLANTOIS. 501 CHAPTER IX. AMNION AND A LL A NTOIS. — D E VE LO P M E N T OF THE CHICK. We shall now proceed to the description of two other accessory organs, which are formed, during the development of the fecundated egg, in all the higher classes of animals. These are the amnion and the allantois ; two organs which are always found in company with each other, since the object of the first is to provide for the forma- tion of the second. The amnion is formed from the external layer of the blastodermic membrane, the allantois from the internal layer. In the frog and in fish, as we have seen, the egg is abundantly supplied with moisture, air, and nourishment, by the water with which it is surrounded. It can absorb directly all the gaseous and liquid substances, which it requires for the purposes of nutrition and growth. The absorption of oxygen, the exhalation of carbonic acid, and the imbibition of albuminous and other liquids, can all take place without diflSculty through the simple membranes of the egg; particularly as the time required for the formation of the embryo is very short, and as a great part of the process of develop- ment remains to be accomplished after the young animal leaves the egg. But in birds and quadrupeds, the time required for the develop- ment of the foetus is longer. The young animal also acquires a much more perfect organization during the time that it remains inclosed within the egg; and the processes of absorption and exhala- tion necessary for its growth, being increased in activity to a corre- sponding degree, require a special organ for their accomplishment. This special organ, destined to bring the blood of the fcetus into relation with the atmosphere and external sources of nutrition, is the allantois. In the frog and the fish, the internal blastodermic layer, forming the intestinal mucous membrane, is inclosed everywhere, as above described, by the external layer, forming the integument; and 502 AMNION AND ALLANTOIS. consequently can nowhere come in contact with the investing membrane of the egg. But in the higher animals, the internal blastodermic layer, which is the seat of the greatest vascularity, and which is destined to produce the allantois, is made to come in contact with the external membrane of the egg for purposes of exhalation and absorption ; and this can only be accomplished by opening a passage for it through the external germinative layer. This is done in the following manner, by the formation of the amnion. Soon after the body of the foetus has begun to be formed by the thickening of the external layer of the blastodermic membrane, a double fold of this external layer rises up on all sides about the edges of the newly formed enabryo ; so that the body of the foetus appears as if sunk in a kind of depression, and surrounded with a membranous ridge or embankment, as in Fig. 190. The embryo (c) is here seen in profile, with the double membranous folds, above men- tioned, rising up just in advance of the head, and behind the posterior extremity. It must be understood, of course, that the same thing takes place on the two sides of the foetus, by the forma- Diagram of fecpn- tiou of lateral folds simultaneously with the BATED Ego; showing appearance of those in front and behind. As it formation of amnion. — ^ i^ a. viteiius. 6. External is thcsc folds which are dcstincd to form the LTbra1e.''TBod7of amniou, they are called the "amniotic folds." embryo d,rf. Amniotic The amuiotic folds contiuue to grow, and ex- folds. e. Vitelline mem- . t .■, -, n jt_i j j ^ j. ^^ ^jj^^g tend themselves lorward, backward and laterally, until they approach each other at a point over the back of the foetus (Fig. 191), which is termed the "amniotic umbilicus." Their opposite edges afterward actually come in con- tact with each other at this point, and adhere together, so as to shut in a space or cavity (Fig. 191, b) between their inner surface and the body of the foetus. This space, which is filled with a clear fluid, is called the amniotic cavity. At the same time, the intestinal canal has begun to be formed, and the umbilical vesicle has been partially separated from it, by the constriction of the abdominal walls on the under surface of the body. There* now appears a prolongation or diverticulum (Fig. 191, c) o-rowincr out from the posterior portion of the intestinal canal, and following the course of the amniotic fold which has preceded it ; occupying, as it gradually enlarges and protrudes, the space left AMNION AND ALLANTOIS. 503 Fecundated Eon, farther advanced. — a. Umbilical vesicle. h. Amniotic cavity, c. Al- laatois. vacant by the rising up of the amniotic fold. This diverticulum is the commencement of the allantois. It is an elongated mem- branous sac, continuous with the posterior portion of the intestine, and containing bloodvessels derived from those of the intestinal circulation. The cavity of the allantois is also continuous with the cavity of the intestine. After the amniotic folds have approached and touched each other, as already described, over the back of the foetus, at the amniotic umbilicus, the adjacent surfaces, thus brought in contact, fuse together, so that the cavities of the two folds, coming respectively from front and rear, are separated only by a single membranous par- tition (Fig. 192, c) running from the inner to the outer lamina of the amniotic folds. This parti- tion itself soon after atrophies and disappears; and the inner and outer larainse become consequently separated from each other. The inner lamina (Fig. 192, a) which remains con- tinuous with the integument of the foetus, in- closing the body of the embryo in a distinct cavity, is called the amnion (Fig. 193, h\ and its cavity is known as the amniotic cavity. The outer lamina of the amniotic fold, on the other hand (Fig. 192, h\ recedes farther and farther from the inner, until it comes in con- tact with the original vitelline membrane, still covering the exterior of the egg ; and by con- tinued growth and expansion it at last fuses with the vitelline membrane and unites with its substance, so that the two membranes form but one. This membrane, formed by the fusion and consolidation of two others, constitutes then the external investing membrane of the Qgg. The allantois, during all this time, is increas- ing in size and vascularity. Following the course of the amniotic folds as before, it insinuates itself between them, and of course soon comes in contact with the external investing membrane just de- scribed. It then begins to expand laterally in every direction, enveloping more and more the body of the foetus, and bringing its vessels into contact with the external membrane of the egg. Fecundated Ego, with allantois nearly com- plete. — a. Inner lamina of amniotic fold. b. Outer la- mina of ditto. c. Point where the amniotic folds come in contact. The allan- tois is seen penetrating be- tween the inner and outer laminse of the amniotic folds. 604 AMNION AND ALLANTOIS. Fig. 193. Fecundated Ego, with allantois fully formed. — a. Um- bilical vesicle, b. Amnion, c. Allantois. By a continuation of the above process, the allantois at last grows to such an extent as to envelope completely the body of the embryo, together with the amnion: its two extremities coming in contact with each other and fusing together over the back of the foetus, just as the amniotic folds had previously done. (Fig. 193.) It lines, there- fore, the whole internal surface of the in- vesting membrane with a flattened, vascu- lar sac, the vessels of which come from the interior of the body of the foetus, and which still communicates with the cavity of the intestinal canal. It is evident, from the above description, that there is a close connection between the formation of the amnion and that of the allantois. For it is only in this manner that the allantois, which is an extension of the in- ternal layer of the blastodermic membrane, can come to be situated outside the foetus and the amnion, and be brought into relation with external surrounding media. The two laminae of the amni- otic folds, in fact, by separating from each other as above described, open a passage for the allantois, and allow it to come in contact with the external membrane of the egg. In order to explain more fully the physiological action of the allantois, we shall now proceed to describe the process of develop- ment, as it takes place in the egg of the fowl. In order that the embryo may be properly developed in any case, it is essential that it be freely supplied with air, warmth, moisture, and nourishment. The egg of the fowl contains already, when discharged from the generative passages, a sufficient quantity of moisture and albuminous material. The necessary warmth is supplied by the body of the parent during incubation ; while the atmospheric gases can pass and repass through the porous egg- shell, and by endosmosis through the fibrous membranes which line its cavity. When the egg is first laid, the vitellus, or yolk, is enveloped in a thick layer of semi-solid albumen. On the commencement of incubation, a liquefaction takes place in the albumen immediately above that part of the vitellus which is occupied by the cicatri- cula ; so that the vitellus rises or floats upward toward the surface by virtue of iis specific gravity, and the cicatricula comes to be DEVELOPMENT OF THE CHICK. 505 placed almost immediately underneath the lining membrane of the egg-shell. As the cicatricula is the spot from which the process of embryonic development commences, the body of the young foetus is by this arrangement placed in the most favorable position for the reception of warmth and other necessary external influences through the egg-shell. The liquefied albumen is also absorbed by the vitelline membrane, and the vitellus thus becomes larger, softer, and more diffluent than before the commencement of incubation. As soon as the circulatory apparatus of the embryo has been fairly formed, two minute arteries are seen to run out from its lateral edges and spread out into the neighboring parts of the blastodermic membrane, breaking up into inosculating branches, and covering the adjacent portions of the vitellus with a plexus of capillary bloodvessels. The space occupied in the blastodermic membrane, on the surface of the vitellus, by these vessels, is called the area vasculosa. (Fig. 194.) It is of a nearly circular shape, Fig. 194. Egg of Fowl during early periods of incubation ; showing the body of the embryo, and the area vasculosa partially covering the surface of the vitellus. and is limited, on its outer edge, by a terminal vein or sinus, called the "sinus terminalis." The blood is returned to the body of the foetus by two veins which penetrate beneath its edges, one near the head and one near the tail. The area vasculosa tends to increase in extent, as the develop- ment of the foetus proceeds and its circulation becomes more active. It soon covers the upper half, or hemisphere, of the vitellus, and the terminal sinus then runs like an equator round the middle of 506 AMNION AND ALLANTOIS. the vitelline sphere. As the growth of the vascular plexus con- tinues, it passes this point, and embraces more and more of the in- ferior, as well as of the superior hemisphere, the vessels converging toward its under surface, until at last nearly the whole of the vitellus is covered with a network of inosculating capillaries. The function of the vessels of the area vasculosa is to absorb nourishment from the cavity of the vitelline sac. As the albumen liquefies during the process of incubation, it passes by endosmosis, more and more abundantly, into the vitelline cavity; the whole vitellus growing constantly larger and more fluid in consistency. The blood of the foetus, then circulating in the vessels of the area vasculosa, absorbs freely the oleagino-albuminous matters of the vitellus, and, carrying them back to the foetus by the returning veins, supplies the newly-formed tissues and organs with abun- dance of nourishment. During this period the amnion and the allantois have been also in process of formation. At first the body of the foetus lies upon its abdomen, as in the cases previously described ; but as it increases in size it alters its position so as to lie more upon its side. The allantois then, emerging from the posterior portion of the abdominal cavity, turns directly upward over the body of the foetus, and comes immediately in contact with the shell membrane. (Fig. 195.) It Fig. 195. Ego of Fowl at a more advanced period of development. The body of the foetus is enveloped by the amnion, and has the umbilical vesicle hanging from its under surface ; while the vascular allantois is seen turning upward and spreading out over the internal surface of the shell-membrane. then spreads out rapidly, extending toward the extremities and down the sides of the egg, enveloping more and more completely DEVELOPMENT OF THE CHICK. 507 the foetus and the vitelline sac, and taking the place of the albumen which has been liquefied and absorbed. It will also be seen, by reference to the figure, that the umbilical vesicle is at the same time formed by the separation of part of the vitellus from the abdomen of the chick; and the vessels of the area vasculosa, which were at first distributed over the vitellus, now ramify, of course, upon the surface of the umbilical vesicle. At last the allantois, by its continued growth, envelopes nearly the whole of the remaining contents of the egg ; so that toward the later periods of incubation, at whatever point we break open the egg, we find the internal surface of the shell-membrane everywhere lined with a vascular membranous expansion, supplied by arteries which emerge from the abdomen of the fcBtus. It is easy to see, accordingly, with what readiness the absorption and exhalation of gases may take place by means of the allantois. The air penetrates from the exterior through the minute pores of the calcareous shell, and then acts upon the blood in the vessels of the allantois very much in the same manner that the air in the minute bronchial tubes and air- vesicles of the lungs acts upon the blood in the pulmonary capillaries. Examination of the egg, furthermore, at various periods of incubation, shows that changes take place in it which are entirely analogous to the process of respiration. The egg, in the first place, during its development, loses water by exhalation. This exhalation is not a simple efl'ect of evaporation, but is the result of the nutritive changes going on in the interior of the egg ; since it does not take place, except in a comparatively slight degree, in unimpregnated eggs, or in those which are not in- cubated, though they may be freely exposed to the air. The ex- halation of fluid is also essential to the processes of development, for it has often been found, in hatching eggs by artificial warmth, that if the air of the chamber in which they are inclosed become unduly charged with moisture, so as to retard or prevent further exhalation, the eggs readily become spoiled, and the development of the embryo is arrested. The loss of weight during natural incu- bation, principally due to the exhalation of water, has been found by Baudrimont and St. Ange' to be over 15 per cent, of the entire weight of the egg. Secondly, the egg absorbs oxygen and exhales carbonic acid. The two observers mentioned above, ascertained that during eigh- ' Du Developpement du Foetus. Paris, 1850, p. 143. 508 AMNION AND ALLANTOIS. teen days' incubation, the egg absorbed nearly 2 per cent, of its weight of oxygen, while the quantity of carbonic acid exhaled from the sixteenth to the nineteenth day of incubation amounted to no less than 3 grains in the twenty-four hours.' It is curious to observe, also, that in the egg during incubation, as well as in the adult animal, more oxygen is absorbed than is returned by exhalation under the form of carbonic acid. It is evident, therefore, that a true respiration takes place by means of the allantois, through the membranes of the shell. The allantois, however, is not simply an organ of respiration ; it takes part also in the absorption of nutritious matter. As the pro- cess of development advances, the skeleton of the young chick, at first entirely cartilaginous, begins to ossify. The calcareous mat- ter, necessary for this ossification, is, in all probability, derived from the shell. The shell is certainly lighter and more fragile toward the end of incubation than at first; and, at the same time, the cal- careous ingredients of the bones increase in quantity. The lime- salts, requisite for the process of ossification, are apparently ab- sorbed from the shell by the vessels of the allantois, and by them transferred to the skeleton of the growing chick ; so that, in the same proportion that the former becomes weaker, the latter grows stronger. This diminution in density of the shell is connected not only with the development of the skeleton, but also with the final escape of the chick from the egg. This deliverance is accomplished mostly by the movements of the chick itself, which become, at a certain period, sufficiently vigorous to break out an opening in the attenuated and weakened egg-shell. The first fracture is generally accomplished by a stroke from the end of the bill ; and it is pre- cisely at this point that the solidification of the skeleton is most advanced. The egg-shell itself, therefore, which at first only serves for the protection of the imperfectly-formed embryo, afterward lurnishes the materials which are used to accomplish its own demo- lition, and at the same time to effect the escape of the fully deve- loped foetus. Toward the latter periods of incubation, the allantois becomes more and more adherent to the internal surface of the shell-mem- brane. At last, when the chick, arrived at the full period of de- velopment, escapes from its confinement, the allantoic vessels are torn off at the umbilicus ; and the allantois itself, cast off as a use- ' Op. cit., pp. 138 and 149. DEVELOPMENT OF THE CHICK. 509 less and effete organ, is left behind in the cavity of the abandoned egg-shell. The allantois is, therefore, strictly speaking, a foetal organ. Developed as an accessory structure from a portion of the intestinal canal, it is exceedingly active and important during the middle and latter periods of incubation ; but when the chick is completely formed, and has become capable of carrying on an in- dependent existence, both the amnion and the allantois are detached and thrown off as obsolete structures, their place being afterward supplied by other organs belonging to the adult condition. 510 DEVELOPMENT OF THE EGG IN HUMAN SPECIES. CHAPTER X, DEVELOPMENT OF THE EGG IN THE HUMAN SPECIES.— FORMATION OF THE CHORION. We have already described, in a preceding chapter, the manner in which the outer lamina of the amniotic fold becomes adherent to the adjacent surface of the vitelline membrane, so as to form with it but a single layer ; and in which these two membranes, thus fused and united with each other, form at that time the single ex- ternal investing membrane of the egg. The allantois, in its turn, afterward comes in contact with the investing membrane, and lies immediately beneath it, as a double vascular membranous sac. In the egg of the human subject the development of the membranes, though carried on essentially upon the same plan with that which we have already described, undergoes, in addition, some further modifications, which we shall now proceed to explain. The first of these peculiarities is that the allantois, after spread- ing out upon the inner surface of Fig. 196. the external investing membrane, adheres to, and fuses with it, just as the outer lamina of the amni- otic fold has previously fused with the vitelline membrane. At the same time, the two layers be- longing to the allantois itself also come in contact and fuse toge- ther ; so that the cavity of the allantois is obliterated, and instead of forming a membranous sac con- taining fluid, this organ is convert- ed into a simple vascular membrane. (Fig. 196.) This membrane, moreover, being, after a time, thoroughly fused and united with the two which have preceded it, takes the place which was previously occupied by them. It is then HrMAN Ovum, about the end of the first month; showing formation of chorion. — 1. Umbilical vesicle. 2. Amnion. 3. Chorion. FOKMATION OF THE CHORION. 511 termed the cJwrion, and thus becomes the sole external investing Tnembrane of the egg. We find, therefore, that the chorion, that is, the external coat or investment of the egg, is formed successively by three distinct mem- branes, as follows: first, the original vitelline membrane; secondly, the outer lamina of the amniotic fold; and, thirdly, the allantois; the last predominating over the two former by the rapidity of its growth, and absorbing them into its substance, so that they become finally completely incorporated with its texture. It is easy to see, also, how, in consequence of the above process, the body of the foetus, in the human egg, becomes inclosed in two distinct membranes, viz., the amnion, which is internal and conti- nuous with the foetal integument, and the chorion, which is external and supplied with vessels from the cavity of the abdomen. The umbilical vesicle is, of course, situated between the two ; and the rest of the space between the chorion and the amnion is occupied by a semi-fluid gelatinous material, somewhat similar in appearance to that of the vitreous body of the eye. The obliteration of the cavity of the allantois takes place very early in the human subject, and, in fact, keeps pace almost entirely with the progress of its growth ; so that this organ never presents, in the human egg, the appearance of a hollow sac, filled with fluid, but rather that of a flattened vascular membrane, enveloping the body of the foetus, and forming the external^membrane of the egg. Notwithstanding this difference, however, the chorion of the human subject is, in respect to its mode of formation, the same thing with the allantois of the lower animals; its chief peculiarity consisting in the fact tbat its opposite surfaces are adherent to each other, instead of remaining separate and inclosing a cavity filled with fluid. The next peculiarity of the human chorion is, that it becomes shaggy. Even while the egg is still very small, and has but recently found its way into the uterine cavity, its exterior is already seen to be covered with little transparent prominences, like so many villi (Fig. 196), which increase the extent of its surface, and assist in the absorption of fluids from without. The villi are at this time quite simple in form, and altogether homogeneous in structure. As the egg increases in size, the villi rapidly elongate, and be- come divided and ramified by the repeated budding and sprouting of lateral offshoots from every part. After this process of growth has gone on for some time, the external surface of the chorion presents 512 DEVELOPMENT OF THE EGG IN HUMAN SPECIES. a uniformly velvety or shaggy appearance, owing to its being co- vered everywhere with these tufted and compound villosities. The villosities themselves, when examined by the microscope, have an exceedingly well marked and characteristic appearance. (Fig. 197.) They originate from the surface of the chorion by a somewhat narrow stem, and divide into a multitude of secondary and tertiary branches, of varying size and figure; some of them slender and filamentous, others club-shaped, many of them irregularly swollen at various points. All of them termi nate by rounded extremities, giving to the whole tuft a certain resem- blance to some varieties of sea- weed. The larger trunks and branches of ,r-"''^\>\T^'"°'''/'S7/<'' '^-^^^^'"■'^^a?^ ^^® villosity are seen to contain nu- ilCr^^N^^^X/^^^^ merous minute nuclei, imbedded in a nearly homogeneous, or finely gra- nular substratum. The smaller ones appear, under a low magnifying power, simply granular in texture. These villi are altogether peculiar in appearance, and quite unlike any other structure which may be met with in the body. Wherever we find, in the uterus, any portion of a membrane having villosities like these, we may be sure that pregnancy has existed ; for such villosities can only belong to the chorion, and the chorion itself is a part of the foetus. It is developed, as we have seen, as an out- growth from the intestinal canal, and can only exist, accordingly, as a portion of the fecundated egg. The presence of portions of a shaggy chorion is therefore as satisfactory proof of the existence of pregnancy, as if we had found the body of the foetus itself. While the villosities which we have just described are in pro- cess of formation, the allantois itself has completed its growth, and has become converted into a permanent chorion. The bloodvessels coming from the allantoic arteries accordingly ramify over the chorion, and supply it with a tolerably abundant vascular network. The growth of the foetus, moreover, at this time, has reached such a state of activity, that it requires to be supplied with nourishment by vascular absorption, instead of the slow process of imbibition, Compound villosity of Human Cho- EION, ramified extremity. From a three mouths' foetus. Magnified .'50 diameters. FORMATION OF THE CHORION. 513 Fig. 198. Extrernily of vrLLOsiTT of Chorion, more highly magni- fied ; showing the ai-rangement of bloodvessels in its interior. which has heretofore taken place through the comparatively incom- })lete and structureless villi of the cho- rion. The capillary vessels, accordingly, with which the chorion is supplied, begin to penetrate into the substance of its vil- losities. They enter the base or stem of each villosity, and, following every divi- sion of its compound ramifications, finally reach its rounded extremities. Here they turn upon themselves in loops (Fig. 198), like the vessels in the papillae of the skin, and retrace their course, to unite finally with the venous trunks of the chorion. The villi of the chorion are therefore very analogous in structure to those of the intestine; and their power of absorp- tion, as in other similar instances, corresponds with the abundance of their ramifications, and the extent of their vascularity. It must be remembered, also, that these vessels all come from the abdomen of the foetus ; and that whatever substances are taken up by them are transported directly to the interior of the embryo, and used for the nourishment of its tissues. The chorion, therefore, as soon as its villi and bloodvessels are completely developed, becomes an exceedingly active organ in the nutrition of the foetus; and con- stitutes, in fact, the only means by which, new material can be in- troduced from without. The existence of this general vascularity of the chorion affords also, as Coste was the first to point out, a striking indication that this membrane is in reality identical with the allantois of the lower animals. If the reader will turn back to the illustrations of the formation of the amnion and allantois (Chap. IX.), he will see that the first chorion or investing membrane is formed exclusively by the vitelline membrane, which is never vascular and cannot be- come so by itself, since it has no direct connection with the foetus. The second chorion is formed by the union of the vitelline mem- brane with the outer lamina of the amniotic fold. Both laminas of the amniotic fold are at first vascular, since they are portions of the external blastodermic layer, and derive their vessels from the integument of the foetus. But after the outer lamina has become completely separated from the inner, by the disappearance of the 33 514 DEVELOPMENT OF THE EGG IN HUMAN SPECIES. partition which for a time connected the two with each other (Fig. 192, c), this source of vascular supply is cut off; and the second cho- rion cannot, therefore, remain vascular after that period. But the third or permanent chorion, that is, the allantois, derives its ves- sels directly from those of the foetus, and retains its connection with them during the whole period of gestation. A chorion, therefore, which is universally and permanently vascular, can be no other tlian the allantois, converted into an external investing membrane of the egg. Thirdly, the chorion, which is at one time, as we have seen, every- where villous and shaggy, becomes afterward partially bald. This change begins to take place about the end of the second month. It commences at a point opposite the situation of the foetus and the insertion of the foetal vessels. The villosities in this region cease growing; and as the entire egg continues to enlarge, the villosities at the point indicated fail to keep pace with its growth, and with the progressive expansion of the chorion. They accordingly be- come at this part thinner and more scattered, leaving the surface of the chorion comparatively smooth and bald. This baldness in- creases in extent and becomes more and more complete, spreading and advancing over the adjacent portions of the chorion, until at least two-thirds of its surface have become nearly or quite destitute of villosities. .' At the opposite point of the surface of the egg, however, that portion, namely, which corre- sponds with the insertion of the foetal vessels, the villosities, instead of becoming atrophied, continue to grow ; and this portion of the chorion becomes even more shaggy and thickly set than before. The conse- quence is that the chorion afterward presents a very dif- ferent appearance at different portions of its surface. (Fig. 199.) The greater part of it is HnMAN OvcM at end of third month; showing g^^yQ^Ji . ^ut a Certain portion, placental portion of the chorion fully formed. i • 1 £• constituting about one-third or the whole, is covered with a soft and spongy mass of long, thickly- set, compound villosities. It is this thickened and shaggy portion, Fig. 199. FORMATION OF THE CHORION. 515 which is afterward concerned in the formation of the placenta : while the remaining smooth portion continues to be known under the name of the chorion. The placental portion of the chorion becomes distinctly limited and separated from the remainder by about the end of the third month. The vascularity of the chorion keeps pace, in its different parts respectively, with the atrophy and development of its villosities. As the villosities shrivel and disappear over a part of its extent, the looped capillary vessels, which they at first contained, disappear also ; so that the smooth portion of the chorion shows afterward only a few straggling vessels running over its surface, and does not contain any abundant capillary plexus. In the thickened portion, on the other hand, the vessels lengthen and ramify to an extent cor- responding with that of the villosities in which they are situated. The allantoic arteries, coming from the abdomen of the foetus, enter the villi, and penetrate through their whole extent ; forming, at the placental portion of the chorion, a mass of tufted and ramified vas- cular loops, while over the rest of the membrane they are merely distributed as a few single and scattered vessels. The chorion, accordingly, is the external investing membrane of the Qgg^ produced by the consolidation and transformation of the allantois. The placenta, furthermore, so far as it has now been described, is evidently a part of the chorion ; that part, namely, which is thickened, shaggy and vascular, while the remainder is comparatively thin, smooth, and membranous. 516 DEVELOPMENT OF UTERINE MUCOUS MEMBRANE. CHAPTER XI. DEVELOPMENT OF UTERINE MUCOUS MEMBRANE.— FORMATION OF TUE DECIDUA. In fish, reptiles, and birds, tlae egg is either provided with a sup- ply of nutritious material contained within its membranes, or it is so placed, after its discharge from the body of the parent, that it can absorb these materials from without. Thus, in the egg of the bird, the young embryo is supported upon the albuminous matter deposited around the vitellus ; while in the frog and fish, moisture, oxygen, saline substances, &c., are freely imbibed from the water in which the egg is placed. But in the quadrupeds, as well as in the human species, the egg is of minute size, and the quantity of nutritious matter which it contains is sufficient to last only for a very short time. Moreover, the development of the foetus takes place altogether within the body of the female, and no supply, therefore, can be obtained directly from the external media. In these instances, accordingly, the mu- cous membrane of the uterus, which is found to be unusually developed and increased in functional activity during the period of gestation, becomes a source of nutrition for the fecundated egg. The uterine mucous membrane, thus developed and hypertrophied, is known by the name of the Decidua. It has received this name because, as we shall hereafter see, it becomes exfoliated and thrown off, at the same time that the egg itself is finally discharged. The mucous membrane of the body of the uterus, in the unimpreg- nated condition, is quite thin and delicate, and presents a smooth and slightly vascular internal surface. There is, moreover, no layer of submucous cellular tissue between it and the muscular substance of the uterus ; so that the mucous membrane cannot here, as in most other organs, be easily dissected up and separated from the subjacent parts. The structure of the mucous membrane itself, however, is sufficiently well marked and readily distinguishable FORMATION OF THE DECIDUA. 517 Fig. 200. Uterine MrroFs Membrakk, as seeu in vertical section. — a. Free surface. 6. Attached surface. from that of other parts. It consists, throughout, of minute tubular follicles, ranged side by side, and running perpendicularly to the free surface of the mucous membrane. (Fig. 200.) Near this free surface, they are nearly straight ; but toward the deeper surface of the mu- cous membrane, where they terminate in blind extremities, or cul-de-sacs, they become more or less wavy or spiral in their cour.se. The tubules are about yig of an inch in diameter, and are lined throughout with co- lumnar epithelium. (Fig. 201.) They occupy the entire thickness of the ute- rine mucous membrane, their closed extremities resting upon the subjacent muscular tissue, while their mouths open into the cavity of the ute- rus. A few fine bloodvessels penetrate the mucous membrane from below, and, running upward between the tubules, encircle Fig- 201. their superficial extremities with a capillary network. There is no areolar tissue in the uterine mucous mem- brane, but only a small quan- tity of spindle-shaped fibro- plastic fibres, scattered be- tween the tubules. As the fecundated egg is about to descend into the cavity of the uterus, the mu- cous membrane, above de- scribed, takes on an increased . . Uterixe Tubules, from mucous membrane of activity of growth and an unimpregnated human uterus. unusual development. It be- comes tumefied and vascular; and as it increases in thickness, it projects, in rounded eminences or convolutions, into the uterine cavity. (Fig. 202.) In this process, the tubules of the uterus in- crease in length, and also become wider; so that their open mouths may be readily seen by the naked e3^e upon the uterine surface, as numerous minute perforations. The bloodvessels of the mucous membrane also enlarge and multiply, and inosculate freely with 518 DEVELOPMENT OF UTERINE MUCOUS MEMBRANE. each other; so that the vascular network encircling the tubules be- comes more extensive and abundant. The internal surface of the uterus, accordingly, after this process has been for some time going on, presents a thick, rich, soft, vas- cular, and velvety lining, quite different from that which is to be found in the unimpregnated condition. In consequence of this difference, the lining membrane of the uterus, in the impregnated condition, was formerly supposed to be an entirely new product, thrown out by exudation from the uterine surface, and analogous, in this respect, to the inflammatory exudations of croup and pleu- risy. It is now known, however, to be no other than the mucous membrane of the uterus itself, thickened and hypertrophied to an extraordinary degree, but still retaining all its natural connections and its original anatomical structure. The hypertrophied mucous membrane, above described, consti- tutes the Decidua vera. Its formation is confined altogether to the body of the uterus, the mucous membrane of the cervix taking no part in the process, but retaining its original appearance. The decidua vera, therefore, commences above, at the orifices of the Fallopian tubes, and ceases below, at the situation of the os inter- num. The cavity of the cervix, at this time, begins to be filled with an abundant secretion of its peculiarly viscid mucus, which blocks up, more or less completely, its passage, and protects the internal cavity. But there is no membranous partition at this time covering the os internum, and the mucous membranes of the cervix and of the body of the uterus, though very different in appearance, are still perfectly continuous with each other. When we cut open the cavity of the uterus, therefore, in this condition, we find its internal surface lined with the decidua vera, with the opening of the OS internum below and the orifices of the Fallopian tubes above, perfectly distinct, and in their natural positions. (Fig. 202.) As the fecundated egg^ in its journey from above downward, passes the lower orifice of the Fallopian tube, it insinuates itself between the opposite surfaces of the uterine mucous membrane, and becomes soon afterward lodged in one of the furrows or de- pressions between the projecting convolutions of the decidua. (Fig. 202.) It is at this situation that an adhesion is subsequently to take place between the external membranes of the egg, on the one hand, and the uterine decidua on the other. Now at the point where the egg becomes fixed and entangled, as above stated, a still more rapid development than before takes place in the uterine FORMATION OF THE DECIDUA. 51^ mucous membrane. Its projecting folds begin to grow up around the egg in such a manner as to partially inclose it in a kind of circumvallation of the decidua, and to shut it off", more or less com- Fiff. 202. Fig. 203. Impreuxatep Uterus; showing formation of decidua. The decidua is represented in black ; and the egg is !*een, at the fundus of the uterus, engaged between two of its projecting convolu- tions. Impregnated Uterus, with pro- jecting folds of decidua growing up around the egg. The narrow opening, where the edges of the folds approach each other, is seen over the most promi- nent portion of the egg. Fi?. 204. pletely, from the general cavity of the uterus. (Fig. 203.) The egg is thus soon contained in a special cavity of its own, which still communicates for a time with the general cavity of the uterus by a small opening, situated over its most prominent portion, which is known as the "decidual umbilicus." As the above process of growth goes on, this opening becomes narrower and narrower, while the projecting folds of decidua approach each other over the sur- face of the egg. At last these folds actually touch each other and unite, forming a kind of cicatrix which remains for a certain time, to mark the situation of the original opening. When the development of the uterus and its contents has reached this point (Fig. 204), it will be seen that the egg is com- pletely inclosed in a distinct cavity of its own ; being everywhere covered with a decidual layer of new formation, which has thus gradually enveloped it, and by which it is concealed from view when the uterine cavity is laid open. This newly formed layer of decidua, enveloping, as , 1-111 • • • n showing egg completely inclosed above aescribea, the projecting portion oi by decidua reflexa. Impregnated Uterus; — 520 DEVELOPMENT OF UTERINE MUCOUS MEMBRANE. the egg, is called the Decidua refiexa ; because it is reflected over the egg, by a continuous growth from the general surface of the uterine mucous membrane. The orifices of the uterine tubules, ac- cordingly, in consequence of the manner in which the decidua reflexa is formed, will be seen not only on its external surface, or that which looks toward the cavity of the uterus, but .also on its internal surface, or that which looks toward the egg. The decidua vera, therefore, is the original mucous membrane lining the surface of the uterus; while the decidua reflexa is a new formation, which has grown up round the egg and inclosed it in a distinct cavity. If abortion occur at this time, the mucous membrane of the uterus, that is, the decidua vera, is thrown off", and of course brings away with it the egg and decidua reflexa. On examining the mass discharged in such an abortion, the egg will accordingly be found imbedded in the substance of the decidual membrane. One side of this membrane, where it has been torn away from its attachment to the uterine walls, is ragged and shaggy; the other side, corres- ponding to the cavity of the uterus, is smooth or gently convoluted, and presents very distinctly the orifices of the uterine tubules; while the egg itself can only be extracted by cutting through the decidual membrane, either from one side or the other, and opening in this way the special cavity in which it has been inclosed. During the formation of the decidua reflexa, the entire egg, as well as the body of the uterus which contains it, has considerably enlarged. That portion of the uterine mucous membrane situated immediately underneath the egg, and to which the egg first became attached, has also continued to increase in thickness and vascu- larity. The remainder of the decidua vera, however, ceases to grow as rapidly as before, and no longer keeps pace with the in- creasing size of the egg and of the uterus. It is still very thick and vascular at the end of the third month ; but after that period it becomes comparatively thinner and less glandular in appearance, while the unusual activity of growth and development is concen- trated in the egg, and in that portion of the uterine mucous mem- brane which is in immediate contact with it. Let us now see in what manner the egg becomes attached to the decidual membrane, so as to derive from it the requisite supply of nutritious material. It must be recollected that, while the above changes have been taking place in the walls of the uterus, the for- mation of the embryo in the egg, and the development of the FORMATION OF THE DECIDUA. 521 Fiff. 205. amnion and chorion have been going on sinnultaneously. Soon after the entrance of the egg into the uterine cavity, its external investing membrane becomes covered with projecting filaments, or villosities, as previously described, (Chap. X.) These villosities, which are at first, as we have seen, solid and non-vascular, insinuate themselves, as they grow, into the uterine tubules, or between the folds of the decidual surface with which the egg is in contact, pene- trating in this way into little cavities or follicles of the uterine mucous membrane, formed either from the cavities of the tubules themselves, or by the adjacent surfaces (>f minute projecting folds. When the formation of the decidua reflexa is accomplished, the chorion has already become uniformly shaggy; and its villosities, spreading in all directions from its external surface, pene- trate everywhere into the follicles above de- scribed, both of the decidua vera underneath it, and the contiguous surface of the decidua reflexa with which it is covered. (Fig. 206.) In this way the egg becomes entangled with the decidua, and cannot then be rea- dily separated from it, without rupturing some of the filaments which have grown from its surface, and have been received into the cavity of the follicles. The nu- tritious fluids, exuded from the soft and glandular textures of the decidua, are now readily imbibed by the villosities of the chorion; and a more rapid supply of nourishment is thus provided, corresponding in abun- dance with the increased and increasing size of the egg. Very soon, however, a still greater activity of absorption be- comes necessary ; and, as we have seen in a preceding chapter, the external membrane of the egg becomes vascular by the formation of the allantoic bloodvessels, which emerge from the body of the foetus, to ramify in the chorion, and penetrate everywhere into the villosities with which it is covered. Each villosity, then, as it lies imbedded in its uterine follicle, contains a vascular loop through which the foetal blood circulates, increasing the rapidity with which absorption and exhalation take place. Subsequently, furthermore, these vascular tufts, which are at first uniformly abundant throughout the whole extent of the chorion, disappear over a portion of its surface, while they at the same time Imprrgnatf, I) Uterus; showiug coiiuection between vil- losities of chorion and decidual membranes. 522 DEVELOPMENT OF UTERINE MUCOUS MEMBRANE. Fig. 206. become concentrated and still further developed at a particular spot, the situation of the future placenta. (Fig. 206.) This is the spot at which the egg is in contact with the decidua vera. Here, therefore, both the decidual membrane and the tufts of the chorion continue to increase in thickness and vascularity; while else- where, over the prominent portion of the egg, the chorion not only becomes bare of villosities, and comparatively destitute of vessels, but the decidua re- flexa, which is in contact with it, also loses its activity of growth, and be- comes expanded into a thin layer nearly destitute of vessels, and without any remaining trace of tubules or follicles. The uterine mucous membrane is therefore developed, during the process of gestation, in such a way as to provide for the nourishment of the foetus in the different stages of its growth. At first, the whole of it is uniformly increased in thickness (decidua vera). Next, a portion of it grows upward around the egg, and " covers its projecting surface (decidua reflexa). Afterward, both the decidua reflexa and the greater part of the decidua vera diminish in the activity of their growth, and lose their importance as a means , of nourishment for the egg; while that part which is in contact with the vascular tufts of the chorion continues to grow, becoming ex- ceedingly developed, and taking an active part in the formation of the placenta. In the following chapter, we shall examine more particularly the structure and development of the placenta itself, and of those parts which are immediately connected with it. Pre.jnaxt Utercs; showing formation of placenta, by the united development of a portion of the de- cidna and the villosities of the cho- THE PLACENTA. 523 CHAPTER XII. THE PLACENTA. We have shown in the preceding chapters that the foetus, during its development, depends for its supply of nutriment upon the lining membrane of the maternal uterus; and that the nutriment, so sup- plied, is absorbed by the bloodvessels of the chorion, and transported in this way into the circulation of the foetus. In all instances, ac- cordingly, in which the development of the foetus takes place within the body of the parent, it is provided for by the relation thus esta- blished between two sets of membranes; namely, the maternal membranes which supply nourishment, and the foetal membranes which absorb it. In some species of animals, the connection between the maternal and foetal membranes is exceedingly simple. In the pig, for ex- ample, the uterine mucous membrane is everywhere uniformly vascular; its only peculiarity consisting in the presence of nume- rous transverse folds, which project from its surface, analogous to the valvulse conniventes of the small intestine. The external in- vesting membrane of the egg, which is the allantois, is also smooth and uniformly vascular like the other. No special development of tissue or of vessels occurs at any part of these membranes, and no direct adhesion takes place between them ; but the vascular allantois or chorion of the foetus is everywhere closely applied to the vascular mucous membrane of the maternal uterus, each of the two contiguous surfaces following the undulations presented by the other. (Fig. 207.) By this arrangement, transudation and absorp- tion take place from the bloodvessels of the mother to those of the foetus, in sufficient quantity to provide for the nutrition of the latter. When parturition takes place, accordingly, in these animals, a very moderate contraction of the uterus is sufficient to expel its contents. The egg, displaced from its original position, slides easily forward over the surface of the uterine mucous membrane, and is at last discharged without any hemorrhage or laceration of connecting 524 THE PLACENTA. parts. In other instances, however, the development of the foetus requires a more elaborate arrangement of the vascular membranes. Fi!?. 207. FcETAL I'ifi, with its membranes, contaiiir-d in cavity of uterus.- c, c. Cavity uf uterus, d. Amnion, e, e. Allantoi.s. I, a, 6, 6. Walls of uterus. In the cow, for example, the external membrane of the Q^g^ beside being everywhere supplied with branching vessels, presents upon various points of its surface no less than from seventy to eighty oval spots, at each of which the vessels of the chorion are developed into abundant tufted prominences, hanging from its exterior as a thick, velvety, vascular mass. At each point of the uterine mucous mem- brane, corresponding with one of these tufted masses, the maternal bloodvessels are developed in a similar manner, projecting into the uterine cavity as a flattened rounded mass or cake; which, with that part of the foetal chorion which is adherent to it, is known by the Cotyledon op Cow's Uterus.— a, a. Surface of fretai chorion 6, 6. Bloodve'ssels of fcetal chorion, d, d. Bloodvessels of uterine mucous membrane, c, c. Surface of uterine mucous mem- braue. name of the Cotyledon. Each cotyledon forms, therefore, a little placenta. (Fig. 208.) In its substance the tufted vascular loops THE PLACENTA. 525 coming from the uterine mucous membrane {d, d) are entangled with those coming from the membranes of the foetus {b, b). There is no absolute adhesion between the two sets of vessels, but only an interlacement of their ramified extremities; and with a little care in manipulation the foetal portion of the cotyledon may be extricated from the maternal portion, without lacerating either. In consequence, however, of this intricate interlacement of the vessels, transudation of fluids will evidently take place with great readiness, from one system to the other. The form of placenta, therefore, met with in these animals, is one in which the bloodvessels of the foetal chorion are simply entangled with those of the uterine mucous membrane. In the human sub- ject, the structure of the placenta is a little more complicated, though the main principles of its formation are the same as in the above instances. From what has already been said in the foregoing chapters, it appears that in the human subject, as well as in the lower animals, the placenta is formed partly by the vascular tufts of the chorion, and partly by the thickened mucous membrane of the uterus in which they are entangled. During the third month, those portions of the chorion and decidua which are destined to undergo this transformation become more or less distinctly limited in their form and dimensions ; and a thickened vascular mass, partly maternal and partly foetal in its origin, shows itself at the spot where the placenta is afterward to be developed. This mass is constituted in the following manner. It will be recollected that the villi of the chorion, when first formed, penetrate into follicles situated in the substance of the uterine mucous membrane; and that after they have become vas- cular, they elongate rapidly and are developed into tufted ramifi- cations of bloodvessels, each one of which turns upon itself in a loop at the end of the villus. At the same time the uterine follicle, into which the villus has penetrated, enlarges to a similar extent ; sending out branching diverticula, corresponding with the mutiplied ramifications of the villus. In fact, the growth of the follicle and that of the villus go on simultaneously and keep pace with each other ; the latter constantly advancing as the cavity of the former enlarges. But it is not only the uterine follicles which increase in size and in complication of structure at this period. The capillary blood- vessels, which lie between them and ramify over their exterior, 52Q THE PLACENTA. also become unusually developed. They enlarge and inosculate freely with each other; so that every uterine follicle is soon covered with an abundant network of dilated capillaries, derived from the bloodvessels of the original decidua. At this time, therefore, each vascular loop of the foetal chorion is covered, first, with a layer forming the wall of the villus. This is in contact with the lining membrane of a uterine follicle, and outside of this again are the capillary vessels of the uterine mucous membrane ; so that two distinct membranes intervene between the walls of the foetal capil- laries on the one hand and those of the maternal capillaries on the other, and all transudation must take place through the substance of these two membranes. As the formation of the placenta goes on, the anatomical arrange- ment of the foetal vessels remains the same. They continue to form vascular loops, penetrating deeply into the decidual mem- brane; only they become constantly more elongated, and their ramifications more abundant and tortuous. The maternal capilla- ries, however, situated on the outside of the uterine follicles, become considerably altered in their anatomical relations. They enlarge excessively; and, by encroaching constantly upon the little islets or spaces between them, fuse successively with each other; and, losing gradually in this way the characters of a capillary network, become dilated into wide sinuses, which communicate freely with the enlarged vessels in the muscular walls of the uterus. As the original capillary plexus occupied the entire thickness of the hyper- trophied decidua, the vascular sinuses, into which it is thus con- verted, are equally extensive. They commence at the inferior surface of the placenta, where it is in contact with the muscular walls of the uterus, and extend through its whole thickness, quite up to the surface of the foetal chorion. As the maternal sinuses grow upward, the vascular tufts of the chorion grow downward, and extend also through the entire thick- ness of the placenta. At this period, the development of the bloodvessels, both in the foetal and maternal portions of the placenta, is so excessive that all the other tissues, which originally co-ex- isted with them, become retrograde and disappear almost altogether. If a villus from the foetal portion of the placenta be examined at this time by transparency, in the fresh condition, it will be seen that its bloodvessels are covered only with a layer of homogeneous, or finely granular material, -ggV^ of an inch in thickness, in which are im- bedded small oval-shaped nuclei, similar to those seen at an earlier THE PLACENTA. 527 Fig. 209. period in the villosities of the chorion. The villosities of the cho- rion are now, therefore, hardly anything more than ramified and tor- tuous vascular loops; the remaining sub- stance of the villi having been atrophied and absorbed in the excessive growth of the bloodvessels. (Fig. 209.) The uterine follicles have at the same time lost all trace of their original structure, and have be- come mere vascular sinuses, into which the tufted foetal bloodvessels are received, as the villosities of the chorion were at first received into the uterine follicles. Finally, the walls of the foetal blood- vessels having come into close contact with the walls of the maternal sinuses, the two become adherent and fuse toge- ther ; so that a time at last arrives, when we can no longer separate the foetal ves- sels, in the substance of the placenta, from the maternal sinuses without lacerating either the one or the other, owing to the second- ary adhesion which has taken place between them. The placenta, therefore, when perfectly formed, has the structure which is shown in the accompanying diagram (Fig. 2i0), repre- Extreraity of Foetal Tuft of humaa placenta; from an in- jected specimen. Magnified 40 diameters. Fiff. 210. Vertical section of Placenta, showing arrangement of maternal and fcetal vess rion. 6, 6. Decidua. c, c, c, c. Orifices of uterine sinuses. 528 THE PLACENTA. senting a vertical section of the organ through its entire thickness. At a, a, is seen the chorion, receiving the umbilical vessels from the body of the foetus through the umbilical cord, and sending out its compound and ramified vascular tufts into the substance of the pla- centa. At b b, is the attached surface of the decidua, or uterine mucous membrane ; and at c, c, c, c, are the orifices of uterine ves- sels which penetrate it from below. These vessels enter the placenta in an extremely oblique direction, though they are represented in the diagram, for the sake of distinctness, as nearly perpendicular. When they have once penetrated, however, the lower portion of the decidua, they immediately dilate into the placental sinuses (represented, in the diagram, in black), which extend through the whole thickness of the organ, closely embracing all the ramifica- tions of the foetal tufts. It will be seen, therefore, that the placenta, arrived at this stage of completion, is composed essentially of nothing but bloodvessels. No other tissues enter into its structure, for all those which it originally contained have disappeared, ex- cepting the bloodvessels of the foetus, entangled with and adherent to the bloodvessels of the mother. There is, however, no direct communication between the foetal and maternal vessels. The blood of the foetus is always separated from the blood of the mother by a membrane which has resulted from the successive union and fusion of four different membranes, viz,, first, the membrane of the foetal villus ; secondly, that of the uterine follicle; thirdly, the wall of the foetal bloodvessel; and, fourthly, the wall of the uterine sinus. The single membrane, how- ever, into which these four finally coalesce, is extremely thin, as we have seen, and of enormous extent, owing to the extremely abundant branching and subdivision of the foetal tufts. These tufts, accordingly, in which the blood of the foetus circulates, are bathed everywhere, in the placental sinuses, with the blood of the mo- ther ; and the processes of endosmosis and exosmosis, of exhala- tion and absorption, go on between the two with the greatest pos- sible activity. It is very easy to demonstrate the arrangement of the foetal tufts in the human placenta. They can be readily seen by the naked eye, and may be easily traced from their attachment at the under surface of the chorion to their termination near the uterine surface of the placenta. The anatomical disposition of the pla- cental sinuses, however, is much more difficult of examination. During life, and while the placenta is still attached to the uterus, THE PLACENTA. 529 they are filled, of course, with the blood of the mother and occupy fully one-half the entire mass of the placenta. But when the pla- centa is detached, the maternal vessels belonging to it are torn off' at their necks (Fig. 210, c, c, c, c), and the sinuses, being then emptied of blood by the compression to which the placenta is sub- jected, are apparently obliterated ; and the foetal tufts, falling to- gether and lying in contact with each other, appear to constitute the whole of the placental mass. The existence of the placental sinuses, however, and their true extent, may be satisfactorily de- monstrated in the following manner. If we take the uterus of a woman who has died undelivered at the full term or thereabout, and open it in such a way as to avoid wounding the placenta, this organ will be seen remaining attached to the uterine surface, with all its vascular connections complete. Let the foetus now be removed by dividing the umbilical cord, and the uterus, with the placenta attached, placed under water, with its internal surface uppermost. If the end of a blowpipe be now introduced into one of the divided vessels of the uterine walls, and air forced in by gentle insufflation, we can easily inflate, first, the venous sinuses of the uterus itself, and next, the deeper por- tions of the placenta; and lastly, the bubbles of air insinuate them- selves everywhere between the foetal tufts, and appear in the most superficial portions of the placenta, immediately underneath the transparent chorion (a a, Fig. 210); thus showing that the placental sinuses, which freely communicate with the uterine vessels, really occupy the entire thickness of the placenta, and are equally ex- tensive with the tufts of the chorion. We have verified this fact in the above manner, on four different occasions, and in the pre- sence of Prof. C. E. Gilman, Dr. Geo. T. Elliot, Dr. Henry B. Sands, Dr. T. G. Thomas, Dr. T. C. Finnell, and various other medical gentlemen of New York. If the placenta be now detached and examined separately, it will be found to present upon its uterine surface a number of openings which are extremely oblique in their position, and which are accordingly bounded on one side by a very thin, projecting, cres- centic edge. These are the orifices of the uterine vessels, passing into the placenta and torn off" at their necks, as above described ; and by carefully following them with the probe and scissors, they are found to lead at once into extensive empty cavities (the pla cental sinuses), situated between the foetal tufts. We have already shown that these cavities are filled during life with the maternal 34 530 THE PLACENTA. blood ; and there is every reason to believe that before delivery, and while the circulation is going on, the placenta is at least twice as large as after it has been detached and expelled from the uterus. The placenta, accordingly, is a double organ, formed partly by the chorion and partly by the decidua ; and consisting of maternal and foetal bloodvessels, inextricably entangled and united with each other. The part which this organ takes in the development of the foetus is an exceedingly important one. From the date of its formation, at about the beginning of the fourth month, it constitutes the only channel through which nourishment is conveyed from the mother to the foetus. The nutritious materials, which circulate in abun- dance in the blood of the maternal sinuses, pass through the inter- vening membrane by endosmosis, and enter the blood of the foetus. The healthy or injurious regimen, to which the mother is subjected, will accordingly exert an almost immediate influence upon the child. Even medicinal substances, taken by the mother and ab- sorbed into her circulation, may readily transude through the pla- cental vessels ; and they have been known in this way to exert a Sj^ecific effect upon the foetal organization. The placenta is, furthermore, an organ of exhalation as well as of absorption. The excrementitious substances, produced in the circulation of the foetus, are undoubtedly in great measure disposed of by transudation through the walls of the placental vessels, to be afterward discharged by the excretory organs of the mother. The system of the mother may therefore be affected in this manner by influences derived from the foetus. It has been remarked more than once, in the lower animals, that when the female has two suc- cessive litters of young by different males, the young of the second litter will sometimes bear marks resembling those of the first male. In these instances, the peculiar influence which produces the ex- ternal mark must have been transmitted by the first male directly to the foetus, from the foetus to the mother, and from the mother to the foetus of the second litter. It is also through the placental circulation that those disturbing effects are produced upon the nutrition of the foetus, which result from sudden shocks or injuries inflicted upon the mother. There is now little room for doubt that various deformities and deficiencies of the foetus, conformably to the popular belief, do really originate, in certain cases, from nervous impressions, such a^ disgust, fear or anger, experienced by the mother. The mode in which these effects may TFIE PLACENTA. 531 be produced is readily understood from what has been said above of the anatomy and functions of the placenta. We know very well how easily nervous impressions will disturb the circulation in the brain, the face, the lungs, &;c, ; and the uterine circulation is quite as readily influenced by similar causes, as physicians see everyday in cases of amenorrhoea, menorrhagia, &c. If a nervous shock may excite premature contraction in the muscular fibres of the pregnant uterus and produce abortion, as not unfrequently happens, it is cer- tainly capable of disturbing the course of the circulation through the same organ. But the foetal circulation is dependent, to a great extent, on the maternal. Since the two sets of vessels are so closely entwined in the placenta, and since the foetal blood has here much the same relation to the maternal, that the blood in the pulmonary capillaries has to the air in the air-vesicles, it will be liable to de rangement from similar causes. If the circulation of air through the pulmonary tubes and vesicles be suspended, that of the blood through the capillaries comes to an end also. In the same way, whatever disturbs or arrests the circulation through the vessels of the maternal uterus must necessarily be liable to interfere with that in the foetal capillaries forming part of the placenta. And lastly, as the nutrition of the foetus is provided for wholly by the placenta, it will of course suffer immediately from any such disturbance of the placental circulation. These effects may be manifested either in the general atrophy and death of the foetus ; or, if the disturbing cause be slight, in the atrophy or imperfect development of particular parts; just as, in the adult, a morbid cause operating through the entire system, may be first or even exclusively manifested in some particular organ, which is more sensitive to its influence than other parts. The placenta must accordingly be regarded as an organ which performs, during intra-uterine life, offices similar to those of the lungs and the intestine after birth. It absorbs nourishment, reno- vates the blood, and discharges by exhalation various excrementi- tious matters, which originate in the processes of foetal nutrition. 532 DISCHARGE OF THE OVUM. CHAPTER XIII. DISCHAKGE OF THE OVUM, AND RETROGRADE DEVELOPMENT (INVOLUTION) OF THE UTERUS. During the growth of the ovum and the formation of the pla- cental structures, the muscular substance of the uterus also in- creases in thickness, while the whole organ enlarges, in order to accommodate the growing foetus and its appendages. The relative positions of the amnion and chorion, furthermore, undergo a change during the latter periods of gestation, and the umbilical cord be- comes developed, at the same time, in the following manner. In the earlier periods of foetal life the umbilical cord consists simply of that portion of the allantois lying next the abdomen. It is then very short, and contains the umbilical vessels running in a nearly straight course, and parallel with each other, from the abdo- men of the foetus to the external portions of the chorion. At this time the amnion closely invests the body of the foetus, so that the size of its cavity is but little Fis- 211. larger than that of the foetus. (Fig. 211.) The space between the amnion and the chorion is then occupied by an amorphous gela- tinous material, in which lies im- bedded the umbilical vesicle. Afterward, however, the am- nion enlarges faster than the cho- rion, and encroaches upon the layer of gelatinous matter situated between the two (Fig. 212), at the same time that an albuminous fluid, the "amniotic fluid," is ex- uded into its cavity, in constantly increasing quantity. Subsequently, the gelatinous layer, above de- scribed, altogether disappears, and the amnion, at about the begin- HcMAX Ovum about the end of the first month. — 1. Umbilical vesicle. 2. Amnion. 3. Choiiuu. ENLARGEMENT OF THE AMNION, 533 Human Ovum at end of third month; showing enlargement of amnion. ning of the fifth month, comes in contact with the internal surface of the chorion. Finally, toward the end of gestation, the contact becomes so close between these two membranes that they are ^'?- ■^^^• partially adherent to each other, and it requires a little care to separate them without laceration. The quantity of the amniotic fluid continues to increase dur- ing the latter periods of gesta- tion in order to accommodate the movements of the foetus. These movements begin to be perceptible about the fifth month, at which time the muscular system has already attained a considerable degree of development, but become after- ward more frequent and more strongly pronounced. The space and freedom requisite for these movements are provided for by the fluid accumulated in the cavity of the amnion. The umbilical cord elongates, at the same time, in proportion to the increasing size of the amniotic cavity. During its growth, it becomes spirally twisted from right to left, the two umbilical arte- ries winding round the vein in the same direction. The gelatinous matter, already described as existing between the amnion and chorion, while it disappears elsewhere, accumulates in the cord in considerable quantity, covering the vessels with a thick, elastic en- velope, which protects them from injury and prevents their being accidentally compressed or obliterated. The whole is covered by a portion of the amnion, which is connected at one extremity with the integument of the abdomen, and invests the whole of the cord with a continuous sheath, like the finger of a glove. (Fig. 213,) The cord also contains, for a certain period, the pedicle or stem of the umbilical vesicle. The situation of this vesicle, it will be recollected, is always between the chorion and the amnion. Its pedicle gradually elongates with the growth of the umbilical cord; and the vesicle itself, which generally disappears soon after the third month, sometimes remains as late as the fifth, sixth, or seventh. According to Prof. Mayer, of Bonn, it may even be found, by care- ful search, at the termination of pregnancy. When discovered in 584 DISCHARGE OF THE OVUM. the middle and latter periods of gestation, it presents itself as a small, flattened, and shrivelled vesicle, situated underneath the amnion, at a variable distance from the insertion of the umbilical cord. A minute bloodvessel is often seen running to it from the cord, and ramifying upon its surface. Fig. 213. Gravid Human Utekus and Contents, showing the relations of the cord, placenta, mem- branes, &c., about the end of the seventh month. — 1. Decidua vera. 2. Decidua reflexa. .S. Chorion. 4. Amnion. The decidua reflexa, during the liitter months of pregnancy, is constantly distended and pushed back by the increasing size of the egg; so that it is finally pressed closely against the opposite surface of the decidua vera, which still lines the greater part of the uterine cavity. By the end of the seventh month, the opposite surfaces of the decidua vera and reflexa are in complete contact with each other, though still distinct and capable of being separated without difficulty. After that time, they fuse together and become con- founded with each other ; the two at last forming only a single, thin, friable, semi-opaque layer, in which no trace of their original oflandular structure can be discovered. This is the condition of things at the termination of pregnancy. Then, the time having arrived for parturition to take place, the hypertrophied muscular walls of the uterus contract forcibly upon its contents, and the egg is discharged, together with the whole of the decidual uterine mucous membrane. In the human subject, as well as in most quadrupeds, the mem- SEPARATION OF THE PLACENTA. 635 branes of the egg are usually ruptured during the process of par- turition; and the foetus escapes first, the placenta and the rest of the appendages following a few moments afterward. Occasionally, however, even in the human subject, the egg is discharged entire, and the foetus liberated afterward by the laceration of the mem- branes. In each case, however, the mode of separation and expul- sion is, in all important particulars, the same. The process of parturition, therefore, consists essentially in a separation of the decidual membrane, which, on being discharged, brings away the ovum with it. The greater part of the decidua vera, having fallen into a state of atrophy during the latter months of pregnancy, is by this time nearly destitute of vessels, and sepa- rates, accordingly, without any perceptible hemorrhage. That por- tion, however, which enters into the formation of the placenta, is, on the contrary, excessively vascular; and when the placenta is separated, and its maternal vessels torn off at their necks, as before mentioned, a gush of blood takes place, which accompanies or immediately follows the birth of the foetus. This hemorrhage, which occurs as a natural phenomenon at the time of parturition, does not come from the uterine vessels proper. It consists of the blood which was contained in the placental sinuses, and which is expelled from them owing to the compression of the placenta by the walls of the uterus. Since the whole amount of blood thus lost was previously employed in the placental circulation, and since the placenta itself is thrown off at the same time, no unpleasant effect is produced upon the mother by such a hemorrhage, because the natural proportion of blood in the rest of the maternal system remains the same. Uterine hemorrhage at the time of parturition, therefore, becomes injurious only when it continues after complete separation of the placenta ; in which case it is supplied by the mouths of the uterine vessels themselves, left open by failure of the uterine contractions. These vessels are usually instantly closed, after separation of the placenta, by the contraction of the muscular fibres of the uterus. They pass, as we have already mentioned, in a,n exceedingly oblique direction, from the uterine surface to the placenta ; and the muscular fibres, which cross them transversely above and below, necessarily constrict them, and effectually close their orifices, immediately on being thrown into a state of contraction. Another very remarkable phenomenon, connected with preg- nancy and parturition, is the appearance in the uterus of a new mucous membrane, growing underneath the old, and ready to take the place of the latter after its discharge. 536 DISCHARGE OF THE OVUM. If the internal surface of the body of the uterus be examined immediately after parturition, it will be seen that at the spot where the placenta was attached every trace of mucous membrane has disappeared. The muscular fibres of the uterus are here perfectly exposed and bare; while the mouths of the ruptured uterine sinuses are also visible, with their thin, ragged edges hanging into the cavity of the uterus, and their orifices plugged with more or less abundant bloody coagula. Over the rest of the uterine surface the decidua vera has also disappeared. Here, however, notwithstanding the loss of the ori- ginal mucous membrane, the muscular fibres are not perfectly bare, but are covered with a thin, semi-transparent film, of a whitish color and soft consistency. This film is an imperfect mucous membrane of new formation, which begins to be produced, underneath the old decidua vera, as early as the beginning of the eighth month. We have seen this new mucous membrane very distinctly in the uterus of a woman who died undelivered at the above period. The old mucous membrane, or decidua vera, is at this time some- what opaque, and of a slightly yellowish color, owing to a partial fatty degeneration which it undergoes in the latter months of preg- nancy. It is easily raised and separated from the subjacent parts, owing to the atrophy of its vascular connections; and the new mucous membrane, situated beneath it, is readily distinguished by its fresh color, and healthy, transparent aspect. The mucous membrane of the cervix uteri, which takes no part in the formation of the decidua, is not thrown off in parturition, but remains in its natural position ; and after delivery it may be seen to terminate at the os internum by an uneven, lacerated edge, where it was formerly continuous with the decidua. Subsequently, a regeneration of the mucous membrane takes place over the whole extent of the body of the uterus. The mucous membrane of new formation, which is already in existence at the time of delivery, becomes thickened and vascular; and glandular tubules are gradually developed in its substance. At the end of two months after delivery, according to Heschl' and Longet,^ it has entirely regained the natural structure of the uterine mucous mem- brane. It unites at the os internum, by a linear cicatrix, with the mucous membrane of the cervix, and the traces of its laceration at ' Zeitschrift der K. K. Gesellschaft der Aerzte, in Wien, 1852. * Traite de Physiol ogie. De la Generation, p. 173. RETROGRADE DEVELOPMENT OF THE UTERUS. 537 this spot afterward cease to be visible. At the point, however, where the placenta was attached, the regeneration of the mucoas membrane is less rapid; and a cicatrix-like spot is often visible at this situation for several months after delivery. The only further change, which remains to be described in this connection, is the fatty degeneration and reconstruction of the muscular substance of the uterus. This process, which is some- times known as the " involu- tion" of the uterus, takes place in the following man- ner. The muscular fibres of the unimpregnated uterus are pale, flattened, spindle- shaped bodies (Fig. 214) near- ly homogeneous in structure or very faintly granular, and measuring from -^^ to gi^ of an inch in length, by T(T5oo to gtj'oo of an inch in width. During gestation these fibres increase very considerably in size. Their texture becomes much more distinctly granular, and their outlines more strongly mark- ed. An oval nucleus also shows itself in the central part of each fibre. The en- tire walls of the uterus, at the time of delivery, are com- posed of such muscular fibres as these, arranged in circu- lar, oblique, and longitudinal bundles. About the end of the first week after delivery, these fibres begin to undergo a fatty degeneration. (Fig. 215.) Their granules be- come larger arid more pro- minent, and very soon as- sume the appearance of Muscular Fibres op Unimpregnated Uterus; from a woman aged 40, dead of pbihisis pulmonalis. Fig. 215. Muscular Fibres of Human Uterus, ten days after parturition ; from a woman dead of puer- peral fever. 538 BISCHAEGE OF THE OVUM. Fig. 216. molecules of fat, deposited in the substance of the fibre. The fatty- deposit, thus commenced, increases in abundance, and the mole- cules continue to enlarge until they become converted into fully formed oil-globules, which fill the interior of the fibre more or less completely, and mask, to a certain extent, its anatomical characters. (Fig. 216.) The universal fatty degeneration, thus induced, gives to the uterus a softer consistency, and a pale yellowish color which is characteristic of it at this period. The muscu- lar fibres which have become altered by the fatty deposit are afterward gradually ab- sorbed and disappear; their place being subsequently taken by other fibres of new M0SCCLAR Fibres of Human Uterus, three formation, which already bc- weeks after parturition ; from a woman dead of peri- . i i • tonitis. gin to make their appearance before the old ones have been completely destroyed. As this process goes on, it results finally in a complete renovation of the muscular substance of the uterus. The organ becomes again reduced in size, compact in tissue, and of a pale ruddy hue, as in the ordinary unimpregnated condition. This entire renewal or reconstruction of the uterus is completed, according to Heschl,' about the end of the second month after delivery. ' Op. cit. DEVELOPMENT OF THE EMBRYO. 539 CHAPTER XIV. DEVELOPMENT OF THE E M B R YO— N E R V U S SYSTEM, ORGANS OF SENSE, SKELETON, AND LIMBS. Fig. 217. The first trace of a spinal cord in the embryo consists of the double longitudinal fold or ridge of the blastodermic membrane, which shows itself at an early period, as above described, on each side the median furrow. The two laminas of which this is com- posed, on the right and left sides (Fig. 217, a, b), unite with each other in front, forming a rounded dilatation (c), the cephalic extremity, and behind at c?, forming a pointed or caudal extremity. Near the poste- rior extremity, there is a smaller dilatation, which marks the future situation of the lumbar enlargement of the spinal cord. As the laminae above described grow upward and backward, they unite with each other upon the median line, so that the whole is converted into a hollow cylindrical cord, terminating ante- riorly by a bulbous enlargement, and posteriorly by a pointed enlargement; the central cavity which it contains running continuously through it, from front to rear. The next change which shows itself is a divi- sion of the anterior bulbous enlargement into caudai extremity three secondary compartments or vesicles (Fig. 218), which are partially separated from each other by transverse constrictions. These vesicles are known as the three cerebral vesi- cles, from which all the different parts of the encephalon are after- ward to be developed. The first, or most anterior cerebral vesicle is destined to form the hemispheres; the second, or middle, the tubercula quadrigemina ; and the third, or posterior, the medulla oblongata. All three vesicles are at this time hollow, and their Formation of Cere- bro-Spinal Axis — a, b. Spinal cord. c. Ce- phalic extremity. d. 540 DEVELOPMENT OF THE EMBRYO. Fig. 218. cavities communicate freely with each other, through the inter- vening constrictions. Yerj soon the anterior and the posterior cerebral vesicles suffer a further division ; the middle one remain- ing undivided. The anterior vesicle thus separates into two portions, of which the first, or larger, constitutes the hemispheres, while the second, or smaller, becomes the optic thalami. The third vesicle also sepa- rates into two portions, of which the ante- rior becomes the cerebellum, and the pos- terior the medulla oblongata. There are, therefore, at this time, five cerebral vesicles, all of whose cavities com- municate with each other and with the central cavity of the spinal cord. The entire cerebro-spinal axis, at the same time, becomes very strongly curved in an ante- rior direction, corresponding with the ante- rior curvature of the body of the embryo (Fig. 219); so that the middle vesicle, or that of the tubercula quadrigemina, occu- pies a prominent angle at the upper part of the encephalon, while the hemispheres and the medulla oblongata are situated below it, anteriorly and posteriorly. At first, it will be observed, the relative size of the various parts of the encephalon is very different from that which they afterward attain in the adult condition. The hemispheres, for example, are hardly larger than the tubercula quadrigemina; and the cerebellum is very much inferior in size to the medulla oblon- gata. Soon afterward, the relative position and size FffiTAL Pig, fire- of the parts bcgiu to alter. The hemispheres and long! Iho°win''g° bra^a tubcrcuk quadrigcmina grow faster than the poste- rior portions of the encephalon ; and the cerebellum becomes doubled backward over the medulla oblon- gata. (Fig. 220.) Subsequently, the hemispheres rapidly enlarge, growing upward and backward, so as to cover in and conceal both the optic thalami and the tuber- cula quadrigemina (Fig. 221); the cerebellum tending in the same way to grow backward, and projecting foriher and farther over the Formation of the Cerebro- spinal Axis. — 1. Vesicle of the hemispheres. 2. Vesicle of the tubercula quadrigemina. 3. Vesicle of the medulla oblongata. Fig. 2 219. 3 % and spinal cord. — 1. Hemispheres. 2. Tu- bercula quadrigemi- na. 3. Cerebellum. 4. Medulla oblongata. NERVOUS SYSTEM. 541 medulla oblongata. The subsequent history of the development of the encephalon is little more than a continuation of the same Fig. 221. F(KTAL Pi«, one and a quai-tor inch long. — 1. Hemispheres. 2. Tubercula. qnadrigemina. 3. Cerebellum. 4. Me- dulla oblongata. HnAn OF FcF.TAL Pio. tbrep and a half inches long. — 1. Hemispheres. 3. Cerebellum. 4. Medulla oblongata. process ; the relative dimensions of the parts constantly changing, so that the hemispheres become, in the adult condition (Fig. 222), Fig. 222. Bratx of .\DrLT Pig. — 1. Hemispheres. 3. Cerebellum. 4. Medulla oblongata. the largest of all the divisions of the encephalon, while the cere- bellum is next in size, and covers entirely the upper portion of the medulla oblongata. The surfaces, also, of the hemispheres and cerebellum, which were at first smooth, become afterward convoluted ; increasing, in this way, still farther the extent of their nervous matter. In the human foetus, these convolutions begin to appear about the beginning of the fifth month (Longet), and grow con- stantly deeper and more abundant during the remainder of foetal life. The lateral portions of the brain growing at the same time more rapidly than that which is situated on the median line, they soon project on each side outward and upward; and, by folding over against each other in the median line, form the right and left hemi- spheres, separated from each other by the longitudinal fissure. 542 DEVELOPMENT OF THE EMBRYO. A similar process of growth taking place in tlie spinal cord results in the formation of the two lateral columns and the anterior and pos- terior median fissures of the cord. Elsewhere the median fissure is less complete, as, for example, between the two lateral halves of the cerebellum, the two optic thalami and corpora striata, and the two tubercula quadrigemina; but it exists everywhere, and marks more or less distinctly the division between the two sides of the nervous centres, produced by the excessive growth of their lateral portions. In this way the whole cerebro-spinal axis is converted into a double organ, equally developed upon the right and left sides, and partially divided by a longitudinal median fissure. Organs of Special Sense. — The eyes are formed by a diverticulum which grows out on each side from the first cerebral vesicle. This diverticulum is at first hollow, its cavity communicating with that of the hemisphere. Afterward, the passage between the two is filled up with a deposit of nervous matter, and becomes the optic nerve. The globular portion of the diverticulum, which is converted into the globe of the eye, has a very thin layer of nervous matter depo- sited upon its internal surface, which becomes the retina; the rest of its cavity being occupied by a gelatinous semi-fluid substance, the vitreous body. The crystalline lens is formed in a distinct fol- licle, which is an offshoot of the integument, and becomes partially imbedded in the anterior portion of the globe of the eye. The cornea also is originally a part of the integument, and remains partially opaque until a very late period of development. Its tissue clears up, however, and becomes perfectly transparent, shortly be- fore birth. The iris is a muscular septum which is formed in front of the crystalline lens, separating the anterior and posterior chambers of the aqueous humor. Its central opening, which afterward becomes the pupil, is at first closed by a vascular membrane, the pupillary membrane^ passing directly across the axis of the eye. The vessels of this membrane, which are derived from those of the iris, subse- quently become atrophied. They disappear first from its centre, and afterward recede gradually toward its circumference; returning always upon themselves in loops, the convexities of which are directed toward the centre of the membrane. The pupillary membrane itself finally becomes atrophied and destroyed, following in this retro- grade process the direction of its receding bloodvessels, viz., from the centre.toward the circumference. It has completely disappeared by the end of the seventh month. (Cruveilhier.) SKELETON AND LIMBS. 543 The eyelids are formed by folds of the integument, which gradually project from above and below the situation of the eye- ball. They grow so rapidly during the second and third months that their free margins come in contact and adhere together, so that they cannot be separated at that time without some degree of vio- lence. They remain adherent from this period until the seventh month (Guy), when their margins separate and they become per- fectly free and movable. In the carnivorous animals, however (dogs and cats), the eyelids do not separate from each other until eight or ten days after birth. The internal ear is formed in a somewhat similar manner with the eyeball, by an oft'shoot from the third cerebral vesicle; the passage between them filling up by a deposit of white substance, which becomes the auditory nerve. The tympanum and auditory meatus are both offshoots from the external integument. Skeleton. — At a very early period of development there appears, as we have already described (Chap. YIL), immediately beneath the cerebro-spinal axis, a cylindrical cord, of a soft, cartilaginous con- sistency, termed the chorda dorsalis. It consists of a fibrous sheath containing a mass of simple cells, closely packed together and united by adhesive material. This cord is not intended to be a permanent part of the skeleton, but is merely a temporary organ destined to disappear as development proceeds. Immediately around the chorda dorsalis there are deposited soon afterward a number of cartilaginous plates, which encircle it in a series of rings, corresponding in number with the bodies of the future vertebrae. These rings increase in thickness from without inward, encroaching upon the substance of the chorda dorsalis, and finally taking its place altogether. The thickened rings, which have been filled up in this way and solidified by cartilaginous deposit, become the bodies of the vertebrae ; while their transverse and articulating processes, with the laminae and spinous processes, are formed by subsequent outgrowths from the bodies hi various directions. When the union of the dorsal plates upon the median line fails to take place, the spinal canal remains open at that situation, and presents the malformation known as spina bifida. This malforma- tion may consist simply in a fissure of the spinal canal, more or less extensive, in which case it may often be cured, or even close spontaneously; or it may be complicated with an imperfect deve- lopment or complete absence of the spinal cord at the same spot. 544 DEVELOPMENT OF THE EMBRYO. when it is accompanied of course by paralysis of the lower ex- tremities, and almost necessarily results in early death. The entire skeleton is at first cartilaginous. The first points of ossification show themselves about the beginning of the second month, almost simultaneously in the clavicle and the upper and lower jaw. Then come, in the following order, the long bones of the extremities, the bodies and processes of the vertebrae, the bones of the head, the ribs, pelvis, scapula, metacarpus and metatarsus, and the phalanges of the fingers and toes. The bones of the carpus, however, are all cartilaginous at birth, and do not begin to ossify until a year afterward. The calcaneum and astragalus begin to ossify, according to Cruveilhier, during the latter periods of foetal life, but the remainder of the tarsus is cartilaginous at birth. The lower extremity of the femur begins to ossify, according to the same author, during the last half of the ninth month. The pisiform bone of the carpus is said to commence its ossification later than any other bone in the skeleton, viz., at from twelve to fifteen years after birth. Nearly all the bones ossify from several distinct points ; the ossification spreading as the cartilage itself increases in size, and the various bony pieces, thus produced, uniting with each other at a later period, usually some time after birth. The limbs appear, by a kind of budding process, as offshoots of the external layer of the blastodermic membrane. They are at first mere rounded elevations, without any separation between the fingers and toes, or any distinction between the different articula- tions. Subsequently the free extremity of each limb becomes di- vided into the phalanges of the fingers or toes ; and afterward the articulations of the wrist and ankle, knee and elbow, shoulder and hip, appear successively from below upward. The posterior extremities, in the human subject, are less rapid in their development than the anterior. Throughout the term of foetal life, indeed, the anterior parts of the body are generally more voluminous than the posterior. The younger the embryo, the larger are the head and upper extremities in proportion to the rest of the body. The lower limbs, and the pelvis more particularly, are very slightly developed in the early periods of growth, as compared with the spinal column, to which they are attached. The inferior ex- tremity of the spinal column, formed by the sacrum and coccyx, pro- jects at this time considerably beyond the pelvis, forming a tail, like that of the lower animals, which is curled forward toward the ab- domen, and terminates in a pointed extremity. Subsequently the SKELETON AND LIMBS. 545 pelvis and the muscular parts seated upon it grow so much faster than the sacrum and coccyx, that the latter become concealed under the adjoining soft parts, and the rudimentary tail accordingly disappears. The integument o^ the embryo is at first thin, vascular, and ex- ceedingly transparent. It afterward becomes thicker, more opaque, and whitish in color; though even at birth it is more vascular than in the adult condition, and the ruddy color of its abundant capil- lary vessels is then very strongly marked. The hairs begin to appear about the middle of intra-uterine life; showing themselves first upon the eyebrows, and afterward upon the scalp, trunk and extremities. The nails are in process of formation from the third to the fifth month ; and, according to Kolliker, are still covered with a layer of epidermis until after the latter period. The seba- ceous matter of the cutaneous glandules accumulates upon the skin after the sixth month, and forms a whitish, semisolid, oleaginous layer, termed the vernix caseosa, which is most abundant in the flexures of the joints, between the folds of the integument, behind the ears and upon the scalp. The cells of the epidermis are repeatedly exfoliated after the first five months of foetal life (Kolliker), and replaced by others, of new formation and of larger size. These exfoliated epidermic cells are found mingled with the sebaceous matter of the vernix caseosa in great abundance. This semi-oleaginous layer, with which the in- tegument is covered, becomes exceedingly useful in the process of parturition, by lubricating the surface of the body, and allowing it to pass easily through the generative passages. 35 546 DEVELOPMENT OF THE ALIMENTARY CANAL CHAPTER XV. DEVELOPMENT OF THE ALIMENTARY CANAL AND ITS APPENDAGES. We have already seen, in a preceding chapter, that the intestinal canal is formed by the internal layer of the blastodermic membrane, which curves forward on each side, and is thus converted into a nearly straight cylindrical tube, terminating at each extremity in a rounded cul-de-sac, and inclosed by the external layer of the blastodermic membrane. The abdominal walls, however, do not unite with each other upon the median line until long after the forjnation of the intestinal canal ; so that, during a certain period, the abdomen of the embryo is widely open in front, presenting a long oval excavation, in which the nearly straight intestinal tube is to be seen, running from its anterior to its posterior extremity. The formation of the stomach takes place in the following man- ner : The alimentary canal, originally straight, soon presents two lateral curvatures at the upper part of the abdomen ; the first to the left, the second to the right. The first of these curvatures be- comes expanded into a wide sac, projecting laterally from the median line into the left hypochondrium, forming the great pouch of the stomach. The second curvature, directed to the right, marks the boundary between the stomach and the duodenum; and the tube at that point becoming constricted and furnished with a circular layer of muscular fibres, is converted into the pylorus. Immedi- ately below the pylorus, the duodenum again turns to the left ; and these curvatures, increasing in number and complexity, form the convolutions of the small intestine. The large intestine forms a spiral curvature ; ascending on the right side, then crossing over to the left as the transverse colon, and again descending on the left side, to terminate by the sigmoid flexure in the rectum. The curvatures of the intestinal canal take place, however, in an antero-posterior, as well as in a lateral direction, and may be best studied in a profile view, as in Fig. 223. The abdominal walls are AND ITS APPENDAGES, 547 here still imperfectly closed, leaving a wide opening at a h, where the integument of the foetus becomes continuous with the com- mencement of the amniotic membrane. The intestine makes at Fiff. 223. Formation of Alimentary Canal. — n,b. Commeneement of amnion, c, c. Intestine, d. Pharynx, e. Uiinai y bladder. /. Allautois. g. Umbilical vesicle, x. Dotted line, showing the place of formation of the oesophagus. first a single angular turn forward, and opposite the most promi- nent portion of this angle is to be seen the obliterated duct, which forms the stem of the umbilical vesicle. A short distance below this point the intestine subsequently enlarges in its calibre, and the situation of this enlargement marks the commencement of the colon. The two portions of the intestine, after this period, becomie widely different from each other. The upper portion, which is the small intestine, grows mostly in the direction of its- length, and be- comes a very long, convoluted, and narrow tube; while the lower portion, which is the large intestine, increases rapidly in diameter, but elongates less than the former. At the point of junction of the small and large intestines, a late- ral bulging or diverticulum of the latter shows itself, and increases in extent, until the ileum seems at last to be inserted obliquely into the side of the colon. This diverticulum of the colon is at first uni- formly tapering or conical in shape ; but afterward that portion which forms its free extremity, becomes narrow and elongated, and is slightly twisted upon itself in a spiral direction, forming the ap- pendix vermiformis; while the remaining portion, which is con- tinuous with the intestine, becomes exceedingly enlarged, and forms the caput coli. The ciiput coli and the appendix are at first situated near the um- 548 DEVELOPMENT OF THE ALIMENTARY CANAL bilicus; but between the fourth and fifth months (Cruveilhier) their position is altered, and they then become fixed in the right iliac region. During the first six months, the internal surface of the small intestine is smooth. At the seventh month, according to Cruveilhier, the valvulse conniventes begin to appear, after which they increase in size till birth. The division of the colon into sac- culi by longitudinal and transverse bands, is also an appearance which presents itself only during the last half of foetal life. Pre- vious to that time, the colon is smooth and cylindrical in figure, like the small intestine. After the small intestine is once formed, it increases very rapidly in length. It grows, indeed, at this time, faster than the walls of the abdomen; so that it can no longer be contained in the abdominal cavity, but protrudes under the form of an intestinal loop, or her- nia, from the umbilical opening. At a subsequent period, on the contrary, the walls of the abdomen grow more rapidly than the intestine. They accordingly gradually envelop the hernial protru- sion, and at last inclose it again in the cavity of the abdomen. Owing to an imperfect development of the abdominal walls, and an imperfect closure of the umbilicus, this intestinal protrusion, which is normal during the early stages of foetal life, sometimes remains at birth, and we then have a congenital umbilical hernia. As the parts at that time, however, have a natural tendency to cicatrize and unite with each other, simple pressure is generally effectual, in such cases, in retaining the hernia within the abdomen, and in producing at last a complete cure. Urinary Bladder^ Urethra, &c. — It will be recollected that very soon after the formation of the intestine, a vascular outgrowth takes place from its posterior portion, which gradually protrudes from the open walls of the abdomen in front, until it comes in contact with the external investing membrane of the egg, and forms, by its con- tinued growth and expansion, the allantois. (Fig. 228,/.) It is at first, as we have shown above, a hollow sac; but, as it spreads out over the surface of the investing membrane of the egg, its two opposite walls adhere to each other, so that its cavity is obliterated at this situation, and it is thus converted into a single vascular membrane, the chorion. This obliteration of the cavity of the allantois commences at its external portion, and gradually extends inward toward the point of its emergence from the abdomen. The hollow tube, or duct, which connects the cavity of the allantois with the posterior part of the intestine, is accordingly converted, as the AND ITS APPENDAGES. 549 process of obliteration proceeds, into a solid, rounded cord. This cord is termed the urachus. After the walls of the abdomen have come in contact, and united with each other at the umbilicus, that portion of the above duct which is left outside the abdominal cavity, forms a part of the um- bilical cord, and remains connected with the umbilical arteries and vein. That portion, on the contrary, which is included in the ab- domen, does not close completely, but remains as a pointed fusiform sac, terminating near the umbilicus in the solid cord of the urachus, and still communicating at its base with the lower extremity of the intestinal canal. This fusiform sac (Fig. 223, e), becomes the wn- nary bladder; and in the foetus at term, the bladder is still conical in form, its pointed extremity being attached, by means of the ura- chus, to the internal surface of the abdominal walls at the situation of the umbilicus. Afterward, the bladder loses this conical form, and its fundus in the adult becomes rounded and bulging. The urinary bladder, as it appears from the abov^e description, at first communicates freely with the intestinal cavity. The intestine, in fact, terminates, at this time, in a wide passage, or cloaca, at its lower extremity, which serves as a common outlet for the urinary and intestinal passages. Subsequently, however, a horizontal par- tition makes its appearance just above the point of junction between the bladder and rectum, and grows downward and forward in such a manner as to divide the above-mentioned cloaca into two parallel and unequal passages. The anterior or smaller of these passages becomes the urethra, the posterior or larger becomes the rectum; and the lower edge of the septum between them becomes finally united with the skin, forming, at its most superficial part, a tole- rably wide band of integument, the perineum, which intervenes be- tween the anus and the external portion of the urethra. The contents of the intestine, which accumulate during foetal life, vary in different parts of the alimentary canal. In the small intes- tine they are semifluid or gelatinous in consistency, of a light yellowish or grayish- white color in the duodenum, becoming yellow, reddish-brown and greenish brown below. In the large intestine they are of a dark greenish hue, and pasty in consistency ; and the contents of this portion of the alimentary canal have received the name of meconium, from their resemblance to inspissated poppy- juice. The meconium contains a large quantity of fat, as well as various insoluble substances, probably the residue of epithelial and mucous accumulations. It does not contain, however, any trace of 550 DEVELOPMEXT OF THE ALIMEXTAEY CAXAL the biliary substances (tauro-cholatesand glyko-cholates) when care- fallj examined by Pettenkofer's test ; and cannot therefore properly be regarded, as is sometimes incorrectly asserted, as resulting from the accumulation of bile. In the contents of the small intestine, on the contrary, traces of bile may be found, according to Lehmann,' so early as between the fifth and sixth months. We have also found distinct traces of bile in the small intestine at birth, but it is even then in extremely small quantity, and is sometimes altogether absent. The meconium, therefore, and the intestinal contents generally, are not composed principally, or even to any appreciable extent, of the secretions of the liver. They appear rather to be produced by the mucous membrane of the intestine itself. Even their yellowish and greenish color does not depend on the presence of bile, since the yellow color first shows itself, in very 3'Oung foetuses, tibout the middle of the small intestine, and not at its upper extremity. The material which accumulates afterward appears to extend from this point upward and downward, gradually filling the intestine, and becoming, in the ileum and large intestine, darker and more pasty as gestation advances. It is a singular fact, perhaps of some importance in this connec- tion, that the amniotic fluid, during the latter half of fcetal life, finds its way, in greater or less abundance, into the stomach, and through that into the intestinal canal. Small cheesy-looking masses may sometimes be found at birth in the fluid contained in the stomach, which are seen on microscopic examination to be no other than portions of the vernix caseosa exfoliated from the skin into the amniotic cavity, and afterward swallowed into the stomach. According to Kcilliker,^ the soft downy hairs of the foetus, exfoliated from the skin, are often swallowed in the same way, and may be found in the meconium. The gastric juice is not secreted before birth ; the contents of the stomach being generally in small quantity, clear, nearly colorless, and neutral or alkaline in reaction. The liver is developed at a very early period. Its size in pro portion to that of the entire body is, in fact, very much greater in the early months than at birth or in the adult condition. In the fcetal pig we have found the relative size of the liver greatest within the first month, when it amounts to very nearly 12 per cent. ' Physiological Chemistry, Philadelphia edition, vol. i. p. 532. * Gewebelehie. Leipzig, 1852, p. 139. AND ITS APPENDAGES. 551 of the entire weight of the body. Afterward, as it grows less rapidly than other parts, its relative weight diminishes successively to 10 per cent, and 6 per cent. ; and is reduced before birth to 3 or 4 per cent. In the human subject, also, the weight of the liver at birth is between 3 and 4 per cent, of that of the entire body. The secretion of hue takes place, as we have intimated above, during foetal life, in a very scanty manner. "We have found it, in minute quantity, in the gall-bladder as well as in the small intes- tine at birth ; but it does not probably take any active part in the nutritive or other functions of the foetus before that period. The glycogenic function of the liver commences during foetal life, and at birth the tissue of the organ is abundantly saccharine. It is remarkable, however, that in the early periods of gestation sugar is produced in the foetus from other sources than the liver. In very young foetuses of the pig, for example, both the allantoic and amniotic fluids are saccharine, a considerable time before any sugar makes its appearance in the tissue of the liver. Even the urine, in half grown foetal pigs, contains an appreciable quantity of sugar, and the young animal is therefore, at this period, in a diabetic con- dition. This sugar, however, disappears from the urine before birth, and also from the amniotic fluid, as has been ascertained by M. Ber- nard;' while the liver begins to produce a saccharine substance, and to exercise the glycogenic function, which it continues after birth. Development of the Pharynx, (Esophagus, &c. — We have already seen that the intestinal canal consists at first of a cylindrical tube, terminated, at each extremity of the abdominal cavity, by a rounded cul-de-sac (Fig. 223, c, c); and that the openings of the mouth and anus are subsequently formed by perforations which take place through the integument and the intervening tissues, and so estab- lish a communication with the intestinal tube. The formation of the anterior perforation and its appendages takes place in the fol- lowing manner: — After the early development of the intestinal tube in the mode above described, the head increases in size out of all proportion to the remainder of the foetus, projecting as a large rounded mass from the anterior extremity of the body, and containing the brain and the organs of special sense. This portion soon bends over toward the abdomen, in consequence of the increasing curvature of the whole body which takes place at this time. In the interior of this ' Lemons de Phjsiologie ExpSrimentale, Paris, 1855, p. 398. 552 DEVELOPMENT OF THE ALIMENTARY CANAL cephalic mass there is now formed a large cavity (Fig. 223, d\ by the melting down and liquefaction of a portion of its substance. This cavity is the pharynx. It corresponds by its anterior extre- mity to the future situation of the mouth; and by its posterior portion to the upper end of the intestinal canal, the future situation of the stomach. It is still, however, closed on all sides, and does not as yet communicate either with the exterior or with the cavity of the stomach. There is, accordingly, at this time, no thorax whatever; but the stomach lies at the upper extremity of the abdomen, immediately beneath the lower extremity of the pharynx, from which it is separated by a wall of intervening tissue. Subsequently, a perforation takes place between the adjacent extremities of the pharynx and stomach, by a short narrow tube, the situation of which is marked by the dotted lines cc, in Fig. 223. This tube afterward lengthens by the rapid growth of that portion of the body in which it is contained, and becomes the oesophagus. Neither the pharynx nor oesophagus, therefore, are, properly speak- ino-, parts of the intestinal canal, formed from the internal layer of the blastodermic membrane; but are, on the contrary, formations of the external layer, from which the entire cephalic mass is pro- duced. The lining membrane of the pharynx and oesophagus is to be regarded, also, for the same reason, as rather a continuation of the integument than of the intestinal mucous membrane; and even in the adult, the thick, whitish, and opaque pavement epithe- lium of the oesophagus may be seen to terminate abruptly, by a well-defined line of demarcation, at the cardiac orifice of the sto- mach ; beyond which, throughout the remainder of the alimentary canal, the epithelium is of the columnar variety, and easily dis- tinguishable by its soft, ruddy, and transparent appearance. As the oesophagus lengthens, the lungs are developed on each side of it by a protrusion from the pharynx, which extends and becomes repeatedly subdivided, forming the bronchial tubes and their ramifications. At first, the lungs project into the upper part of the abdominal cavity; for there is still no distinction be- tween the chest and abdomen. Afterward, a horizontal partition begins to form on each side, at the level of the base of the lungs, which gradually closes together at a central point, so as to form the diaphragm, and finally to shut off altogether the cavity of the chest from that of the abdomen. Before the closure of the diaphragm, thus formed, is complete, a circular opening exists on each side the median line, by which the peritoneal and pleural AND ITS APPENDAGES, 553 cavities communicate with each other. In some instances the de- velopment of the diaphragm is arrested at this point, either on one side or the other, and the opening accordingly remains permanent. The abdominal organs then partially protrude into the cavity of the chest on that side, forming congenital diaphragmatic hernia. The lung on the affected side also usually remains in a state of imperfect development. Diaphragmatic hernia of this character is more frequently found upon the left side than upon the right. It may sometimes continue until adult life without causing any serious iuconvenience. The heart is formed, at a very early period, directly in front of the situation of the oesophagus. Its size soon becomes very large in proportion to the rest of the body; so that it protrudes beyond the level of the thoracic parietes, covered only by the pericardium. Subsequently, the walls of the thorax, becoming more rapidly developed, grow over it and inclose it. In certain instances, how- ever, they fail to do so, and the heart then remains partially or completely uncovered, in front of the chest, presenting the condi- tion known as ectopia cordis. This malformation is necessarily fatal. Development of the Face. — While the lower extremity of the pharynx communicates with the cavity of the stomach, as above described, its upper extremity also becomes perforated in a similar manner, and establishes a communication with the exterior. This perforation is at first wide and gaping. It afterward becomes divided into the mouth and nasal passages; and the different parts of the face are formed round it in the following manner: — From the sides of the cephalic mass five buds or processes shoot out, and grow toward each other, so as to approach the centre of the oral orifice above men- tioned. (Fig. 224.) One of them grows directly downward from the frontal region (i), and is called the frontal or inter- maxillary process, because it afterward contains in its lower extremity the inter- maxillary bones, in which the incisor HeadofHumaxEmbryo, teeth of the upper jaw are inserted. The at about the twentieth day. After next process (2) originates from the side ^"°^''= ^'■°™ ^ specimen iu the ^ ^ ^ ^ ° collection of M.Coste.—l. Frontal of the opening, and, advancing toward the or intermaxillary process. 2. Pro- median line, forms, with its fellow of the -- of ^yi-io-- "-^;>'- s- i'^^- ' ' cess of inferior maxiUa. Fig. 224. 554 DEVELOPMENT OF THE ALIMENTAKY CANAL opposite side, the superior maxilla. The processes of the remain- ing pair (3) also grow from the side, and form, by their subsequent union upon the median line, the inferior maxilla. The inferior maxillary bone is finally consolidated, in man, into a single piece, but remains permanently divided, in the lower animals, by a suture upon the median line. As the frontal process grows from above downward, it becomes double at its lower extremity (Fig. 225), and at the same time two offshoots show themselves upon its sides (1), which curl round and inclose two circular orifices (5), the opening of the an- terior nares; the offshoots themselves be- coming the alas nasi. The processes of the superior maxilla continue their growth, but less rapidly than those of the inferior ; so that the two sides of the lower jaw are already consolidated with each other, while those of the upper jaw are still separate. As the processes of the superior maxilla continue to enlarge, they also tend to unite with each other on the median line, but are prevented from doing so by the intermax- illary processes which grow down between them. They then unite with the inter- maxillary processes, which have at the same time united with each other, and the upper jaw and lip are thus completed. (Fig. 226.) The external edge of the ala nasi also adheres to the superior maxillary pro- cess and unites with it, leaving only a curved crease or furrow, as a sort of cicatrix, to mark the line of union be- tween them. Sometimes the superior maxillary and the intermaxillary processes fail to unite with each other; and we then have the malformation known as hare- lip. The fissure of hare-lip, conse- TiEAD OF Human Embryo, about queutly, is ncvcr cxactly in the median tbo end of the second month —From a ,. i t „ i;<.*.1„ 4.^ ^v^^ c,\A^ ^P if ,->ti ., , line, but a little to one side oi it, on specimen in the author's possession. iiuc, uliu c* i.tvi,i.^ uv. vy v/ , Head of Human Embryo at the end of the first month. After Longet ; from a specimen in the collection of M. Co-^te — ]. AUnasi. 2. Superior maxilla. .3. Inferior maxilla. 4. Inter- maxillary process. 0. Nostril. 6. Eye. 226. AND ITS APPENDAGES. 555 the external edge of the intermaxillary process. Occasionally, the same deficiency exists on both sides, producing "double hare-lip;" in which case, if the fissures extend through the bony structures, the central piece of the superior maxilla, which is detached from the remainder, contains the four upper incisor teeth, and corres- ponds with the intermaxillary bone of the lower animals. The eyes at an early period are situated upon the sides of the head, so that they cannot be seen in an anterior view. (Fig. 224.) As development proceeds, they come to be situated farther forward (Fig. 225), their axes being divergent and directed obliquely for- ward and outward. At a later period still they are placed on the anterior plane of the face (Fig. 226) and have their axes nearly parallel and looking directly forward. This change in the situa- tion of the eyes is effected by the more rapid growth of the pos- terior and lateral parts of the head, which enlarge in such a manner as to alter the relative position of the parts seated in front of them. The palate is formed by a septum between the mouth and nares, which arises on each side as a horizontal plate or offshoot from the superior maxilla. These two plates afterward unite with each other upon the median line, forming a complete partition between the oral and nasal cavities. The right and left nasal passages are also separated from each other by a vertical plate (vomer), which grows from above downward and fuses with the palatal plates be- low. Fissure of the palate is caused by a deficiency, more or less complete, of one of the horizontal maxillary plates. It is accord- ingly situated a little to one side of the median line, and is fre- quently associated with hare-lip and fissure of the upper jaw. The fissures of the palate and the lip are very often continuous with each other. The anterior and posterior pillars of the fauces are incomplete vertical partitions, which grow from the sides of the oral cavity, and tend to separate, by a slight constriction, the cavity of the mouth from that of the pharynx. When all the above changes are accomplished, the pharynx, oesophagus, mouth, nares, and fauces, with their various projections and divisions, have been successively formed; and the development of the upper part of the alimentary, canal is then complete. 556 DEVELOPMENT OF THE KIDNEYS. CHAPTER XVI. DEVELOPMENT OF THE KIDNEYS, WOLFFIAN BODIES, AND INTERNAL ORGANS OF GENE- RATION. The first trace of a urinary apparatus in the embryo, consists of two long, fusiform bodies, which make their appearance in the ab- domen at a very early period, situated on each side the spinal column. I^ese are known by the name of the Wolffian bodies. They are fully formed, in the human subject, toward the end of the first month (Coste), at which time they are the largest organs in the cavity of the abdomen, extending from just below the heart, nearly to the posterior extremity of the body. In the foetal pig, when a little over half an inch in length (Fig. 227), the Wolffian bodies are rounded and kidney-shaped, and occupy a very large part of the abdominal cavity. Their importance may be estimated from the fact that their weight at this time is equal to a little over -3-^3 of that of the entire body — a proportion which is seven or eight times as large as that of the kidneys, in the adult condition. There are, indeed, at this period, only three organs perceptible in the abdo- men, viz., the liver, which has begun to be formed at the upper part of the abdominal cavity; the intestine, which is already some- what convoluted, and occupies its central portion ; and the Wolffian bodies, which pro- ject on each side the spinal column. The WolflBan bodies, in their intimate structure, resemble very closely the adult kidney. They consist of secreting tubules, lined with epithelium, which run from the outer toward the inner edge of the organ, terminating at their free extremities in small rounded dilatations, or culs-de-sac. Into each FfETALPiu, 5^ of an inch long; from a specimen in the autlior's possession. 1. Heart. 2. Anterior extremity. 3. Pos- terior extremity. 4. Wolfiian body The abdominal walls have been cut away, in order to show the position of the Wolffian bodies WOLFFIAN BODIES. 557 of these dilated extremities is received a globular coil of capillary bloodvessels, or glomerulus, similar to that of the adult kidney. The tubules of the Wolffian body all empty into a common excre- tory duct, which leaves the organ at its lower extremity, and com- municates afterward with the lower part of the intestinal canal, just at the point where the diverticulum of the allantois is given off", and where the urinary bladder is afterward to be situated. The prin- cipal, if not the only distinction, between the minute structure of the Wolffian bodies and that of the true kidneys, consists in the size of the tubules and of their glomeruli, these elements being considerably larger in the Wolffian body, than in the kidney. In the foetal pig, for example, about an inch and a half in length, the diameter of the tubules of the Wolffian body is 5^^ of an inch, while in the kidney of the same foetus, the diameter of the tubules is only ^i^ of an inch. The glomeruli in the Wolffian bodies measure ^'^ of an inch in diameter, wbile those of the kidney mea- sure only y| of an inch. The Wolffian bodies are therefore urinary organs, so far as regards their anatomical structure, and are some- times known, accordingly, by the name of the "false kidneys." There is little doubt that they perform, at this early period, a func- tion analogous to that of the kidneys, and separate from the blood of the embryo an excrementitious fluid which is discharged by the ducts of the organ into the cavity of the allantois. Subsequently, the Wolffian bodies increase for a time in size, though not so rapidly as the rest of the body; and consequently their relative magnitude diminishes. Still later, they begin to suffer an absolute diminution or atrophy, and become gradually less and less perceptible. In the human subject, they are hardlv to be detected after the end of the second month (Longet), and in the quadrupeds also they disappear completely long before birth. They are consequently foetal organs, destined to play an important part during a certain stage of development, but to become after- ward atrophied and absorbed, as the physiological condition of the foetus alters. During the period, however, of their retrogression and atrophy, other organs appear in their neighborhood, which become afterward permanently developed. These are, first, the kidneys, and secondly, the internal organs of generation. The kidneys are formed just behind the Wolffian bodies, and are at first entirely concealed by them in a front view, the kidneys being at this time not more than a fourth or a fifth part the size of 558 DEVELOPMENT OF THE KIDNEYS. FfETAL PiQ, one and a half inches long. From a specimen in the author's possession. — 1. Wolffian body. 2. Kidney. the Wolffian bodies. (Fig. 228.) As the kidneys, however, subse- quently enlarge, while the Wolffian bodies diminish, the propor- tions existing between the two organs are reversed; and the Wolffian bodies at last come to be mere small rounded or ovoid masses, situated on the anterior surface of the kidneys. (Figs. 229 and 230.) The kidneys, during this period, grow more rapidly in an upward than in a downward direction, so that the Wolffian bodies come to be situated near their inferior extremity, and seem to have performed a sliding movement from above down- ward, over their anterior surface. This apparent sliding movement, or descent of the AYolffian bodies, is owing entirely to the rapid growth of the kidneys in an upward direction, as we have already explained. The kidneys, during the succeeding periods of foetal life, become in their turn very largely developed in proportion to the rest of the organs; attaining a size, in the foetal pig, equal to ^^ (in weight) of that of the entire body. This proportion, however, diminishes again very considerably before birth, owing to the increased deve- lopment of other parts. In the human foetus at birth, the weight of the two kidneys taken together is y^g that of the entire body. . Internal Organs of Generation. — About the same time that the kid- neys are formed behind the Wolffian bo- dies, two oval shaped organs make their appearance in front, on the inner side of the Wolffian bodies and between them and the spinal column. These bodies are the internal organs of generation ; viz., the testicles in the male, and the ovaries in the female. At first they occupy pre- cisely the same situation and present precisely the same appearance, whether IXT.RXAT.OROAXS OF nK>-K- tijgfoetus Is aftcrwardto belong to the KATioN, &c. ; lu a foetal pig three ° inches long. From a specimen in the male Or the fcmalc SCX. (Fig. 229.) author's Possession. — 1,1. Kidneys. \ ^ i. j- i. i, j.i • j. ^ 2,2. Wolffian bodies. .3,3. Internal ^ ^hort distancc abovc thc mtcmal organsofgeneration; testicles or ova- orgaDS of generation there commenccs, ries. 4. Urinary bladder, turned over i • i , i i j iu front. 5. -Intestine. ou cach Side, a narrow tube or duct, MALE ORGANS OF GENERATION. 559 which runs from above downward along the anterior border of the Wolffian body, immediately in front of and parallel with the excre- tory duct of this organ. The two tubes, right and left, then approach each other below; and, joining upon the median line, empty, together with the ducts of the Wolffian bodies, into the base of the allantois, OT what will afterward be the base of the urinary bladder. These tubes serve as the excretory ducts of the internal organs of genera- tion ; and will afterward become the vasa deferentia in the male, and the Fallopian tubes in the female. According to Coste, the vasa de- ferentia at an early period are disconnected with the testicles ; and originate, like the Fallopian tubes, by free extremities, presenting each an open orifice. It is only afterward, according to the same author, that the vasa deferentia become adherent to the testicles, and a communication is established between them and the tubuli serni- niferi. In the female, the Fallopian tubes remain permanently disconnected with the ovaries, except by the edge of the fimbriated extremity ; which in many of the lower animals becomes closely adherent to the ovary, and envelopes it more or less completely. Male Organs of Generation ; Descent of the Testicles. — In the male foetus there now commences a movement of translation, or change of place, in the internal organs of generation, which is known as the " descent of the testicles." In consequence of this movement, the above organs, which are at first placed near the middle of the abdomen, and directly in front of the kidneys, come at last to be situated in the scrotum, altogether outside and below the abdominal cavity. They also become inclosed in a distinct serous sac of their own, the tunica vaginalis testis. This apparent movement of the testicles is accomplished in the same manner as that of the Wolf- fian bodies, above mentioned, viz., by a disproportionate growth of the middle and upper portions of the abdomen and of the organs situated above the testicles, so that the relative position of these or- gans becomes altered. The descent of the testicles is accompanied by certain other alterations, in the organs themselves and their ap- pendages, which take place in the following manner. By the upward enlargement of the kidneys, both the Wolffian bodies and the testicles are soon found to be situated near the lower extremity of these organs. (Fig. 230.) At the same time, a slender rounded cord (not represented in the figure) passes from the lower extremity of each testicle in an outward and downward direction, crossing the corresponding vas deferens a short distance above its union with its fellow of the opposite side. Below this 660 DEVELOPMENT OF THE KIDNEYS. Internal Organs of Generation, &c , in a foetal pig nearly four inches long. From a specimen in the author's possession. — 1, 1 Kidneys. 2, 2. Wolffian bodies. 3, 3. Testicles. 4. Urinary bladder. 5. Intestine. converted into the epididymis. point, the cord spoken of continues to run obliquely outward and downward ; and, passing through the abdominal walls at the situ- ation of the inguinal canal, is inserted into the subcutaneous tissues near the symphysis pubis. The lower part of this cord becomes the gubernaculum testis ; and mus- cular fibres are soon developed in its substance which may be easily detected, even in the human foetus, during the latter half of gestation. At the period of birth, however, or soon afterward, these muscular fibres disappear and can no longer be recognized. All that portion of the excre- tory tube of the testicle which is situated outside the crossing of the gubernaculum, is destined to be- come afterward convoluted, and That portion which is situated in- side the same point remains comparatively straight, but becomes considerably elongated, and is finally known as the vas deferens. As the testicles descend still farther in the abdomen, they con- tinue to grow, while the Wolffian bodies, on the contrary, diminish rapidly in size, until the latter become much smaller than the tes- ticles ; and at last, when the testicles have arrived at the internal inguinal ring, the Wolffian bodies have altogether disappeared, or at least have become so much altered that their characters are no longer recognizable. In the human foetus, the testicles arrive at the internal inguinal ring, about the termination of the sixth month (Wilson). During the succeeding month, a protrusion of the peritoneum takes place through the inguinal canal, in advance of the testicle ; while the last named organ still continues its descent. As it then passes downward into the scrotum, certain muscular fibres are given off from the lower border of the internal oblique muscle of the abdomen, growing downward with the testicle, in such a manner as to form a series of loops upon it, and upon the elongating spermatic cord. These loops constitute afterward the cremaster muscle. At last, the testicle descends fairly to the bottom of the scrotum, the gubernaculum constantly shortening, and the vas deferens MALE ORGANS OF GENERATION". 561 elongating as it proceeds. The convoluted portion of the efferent duct, viz., the epididymis, then remains closely attached to the body of the testicle ; while the vas deferens passes upward, in a reverse direction, enters the abdomen through the inguinal canal, again bends downward, and joins its fellow of the opposite side; after which they both open into the prostatic portion of the urethra, upon the median line, by a common orifice (sinus pocularis). At the same time, two diverticula arise from the median portion of the vasa deferentia, and, elongating in a backward direction, underneath the base of the bladder, become developed into two compound sacculated reservoirs — the vesiculoe seminales. The left testicle is a little later in its descent than the right, but it afterward passes farther into the scrotum, and, in the adult condi- tion, usually hangs a little lower than its fellow of the opposite side. After the testicle has fairly passed into the scrotum, the serous pouch, which preceded its descent, remains for a time in communi- cation with the peritoneal cavity. In many of the lower animals, as, for example, the rabbit, this condition is permanent ; and the testicle, even in the adult animal, may be alternately drawn down- ward into the scrotum, or retracted into the abdomen, by the action of the gubernaculum and the cremaster muscle. But in the human foetus, the two opposite surfaces of the peritoneal pouch, covering the testicle, approach each other at the inguinal canal, forming at that point a constriction or neck, which partly shuts off the testicle from the cavity of the abdomen. By a continuation of this process, the serous surfaces come actually in contact with each other, and, adhering together at this situation (Fig. 281, 4), form a kind of cicatrix, or umbilicus, by the complete closure and consolidation of which the cavity of the tunica vaginalis (2) is finally shut off altogether from the general cavity of the peritoneum (3). The tunica vagi- nalis testis is, therefore, originally a part of the peritoneum, from which it is sub- sequently separated by the process just described. The separation of the tunica vaginalis from the peritoneum is usually completed in the human subject before birth. But sometimes it fails to take 36 Fig. 231. Formation of Tunica Va- ginalis Testis.— 1. Testicle^ nearly at the bottom of the scro- tum. 2. Cavity of tunica Tdginali.K, 3. Cavity of peritoneum. 4. Obliter- ated neck of peritoneal sac. 562 DEVELOPMENT OF THE KIDNEYS, ETC. place at the proper time, and the intestine is then apt to protrude into the scrotum, in front of the spermatic cord, giving rise, in this way, to a congenital inguinal hernia. (Fig. 232.) The parts impli- cated, however, in this malformation, have ^^' still, as in the case of congenital umbili- cal hernia, a tendency to unite with each other and obliterate the unnatural open- ings ; and if the intestine be retained by pressure in the cavity of the abdomen, cicatrization usually takes place at the inguinal canal, and a cure is effected. The descent of the testicle, above de- scribed, is not accomplished by the forci- ble traction of the muscular fibres of the co.v-o^s s^s^m; show- ^ ' i- tJ mg the vertebral veins itself into the lower extremity of the heart. The emptying into the heart . ^ ^ -I r 1 • T J.^ ■ n ^7 two lateral trunks, two branches, by means o± which the veins of the "eauaisof cuvier." Fig 239. 574 DEVELOPMENT OF THE CIRCULATORY APPARATUS. Fig. 240. Venous System far- ther advanced, showing formation of iliac and sub- clavian veins. — a. Vein of new formation, which be- comes the inferior vena cava. ft. Transverse branch of new formation, which afterward becomes the left vena innominata. Fig. 241. Further development of the Venous System — The vertebral veins are much diminished in size, and the canal of Cuvier, on the left side, is gradually- disappearing, c. Trans- verse branch of new forma- tion, which is to become the vena azygos minor. the lower extremities thus unite, become after- ward, by enlargement, the common iliac veins; while the single trunk (a) resulting from their union becomes the vena cava inferior. Subse- quently, the vena cava inferior becomes very much larger than the vertebral veins ; and its two branches of bifurcation are afterward re- presented by the two iliacs. Above the level of the heart, the vertebral and intercostal veins retain their relative size until the development of the superior extremi- ties has commenced. Then two of the inter- costal veins increase in diameter (Fig. 240), and become converted into the right and left sub- clavians ; while those portions of the vertebral veins situated above the subclavians become the right and left jugulars. Just below the junction of the jugulars with the subclavians, a small branch of communication now appears between the two vertebrals (Fig. 240, 6), pass- ing over from left to right, and emptying into the right vertebral vein a little above the level of the heart ; so that a part of the blood coming from the left side of the head, and the left upper extremity, still passes down the left vertebral vein to the heart upon its own side, while a part crosses over by the communicating branch (6), and is finally conveyed to the heart by the right descending vertebral. Soon afterward, this branch of communication enlarges so rapidly that it preponderates altogether over the left superior vertebral vein, from which it ori- ginated (Fig. 241), and, serving then to convey all the blood coming from the left side of the head and left upper extremity over to the right side above the heart, it becomes the left vena innominata. On the left side, that portion of the superior vertebral vein, which is below the subclavian, remains as a small branch of the vena innomi nata, receiving the six or seven upper intercostal DEVELOPMENT OF THE VENOUS SYSTEM. 575 Fig. 242. veins; while on the right side it becomes excessively enlarged, receiving the blood of both jugulars and both subclavians, and is converted into the vena cava superior. The left canal of Cuvier, by which the left vertebral vein at first communicates with the heart, subsequently becomes atrophied and disappears; while on the right side it becomes excessively enlarged, and forms the lower extremity of the vena cava superior. The superior and inferior venae cavae, accordingly, do not cor- respond with each other so far as regards their mode of origin, and are not to be regarded as analogous veins. For the superior vena cava is one of the original vertebral veins ; while the inferior vena cava is a totally distinct vein, of new formation, resulting from the union of the two lateral trunks coming from the infe- rior extremities. The remainder of the vertebral veins finally assume the condition shown in Fig. 242, which is the complete or adult form of the venous circulation. At the lower part of the abdomen, the vertebral veins send inward small trans- verse branches, which communicate with the vena cava inferior, between the points at which they receive the intercostal veins. These branches of communication, by increasing in size, become the lumbar veins (7), which, in the adult condition, communicate with each other by arched branches, a short distance to the side of the vena cava. Above the level of the lumbar arches, the vertebral veins retain their .5. Vena cava inferior. 6, 6. original direction. .That upon the right side niac veins v. Lumbar veins, still receives all the right intercostal veins, and '; ^'''"^ ■''''^'' ""'"'Z' J' o ' Vena azygos minor. 10. su- becomes the vena azyjos major (s). It also perior intercostal vein. receives a small branch of communication from its fellow of the left side (Fig. 24.1, c), and this branch soon enlarges to such an extent as to bring over to the vena azygos major all the blood of the five or six lower intercostal veins of the left side, becoming, in this way, the vena azygos minor {9). The six or seven upper intercostal veins on the left side still empty, as before, into their own vertebral vein (10), which, joining the left vena innomi- nata above, is known as the superior intercostal vein. The left canal Adult condition of Ve- xors System. — 1. Riglit auricle of heart. 2. Vena cava superior. 3, 3. Jugular veins. 4,1:. Subclavian veins. 576 DEVELOPMENT OF THE CIECULATORY APPARATUS. of Cuvier lias by this time entirely disappeared; so that all the venous blood now enters the heart by the superior or the inferior vena cava. But the original vertebral veins are still continuous throughout, though very much diminished in size at certain points; since both the greater and lesser azygous veins inosculate below with the superior lumbar veins, and the superior intercostal vein also inosculates below with the lesser azygous, just before it passes over to the right side. There are still two parts of the circulatory apparatus, the deve- lopment of which presents peculiarities sufficiently important to be described by themselves. These are, first, the liver and the ductus venosus, and secondly, the heart, with the ductus arteriosus. Development of the Hepatic Circulation and the Ductvs Venosus. — The liver appears at a very early period in the upper part of the abdomen, as a mass of glandular and vascular tissue, which is deve- loped around the upper portion of the om- Fig. 243. phalo-rnesenteric vein, just below its termi- nation in the heart. (Fig. 243.) As soon as the organ has attained a considerable size, the omphalo-mesenteric vein (i) breaks up in its interior into a capillary plexus, the vessels of which unite again into venous trunks, and so convey the blood finally to the heart. The omphalo-mesenteric vein below the liver Early form of Hepatic then hecomes the portal veiu ; while above tiie ciKccLATioN. 1 ompha- ^. ^^^ between that organ and the heart, lo mesenteric vein. 2. Hepa- ' o ' tic vein. 3. Heart. Tiiedotted [{, rcccivcs the uamc of the hcpatic Vein (2). line shows the situation of „, ,. t i • . .1 • . • v i the fnture umbilical vein. The hvcr, accordiugly, is at this time supplied with blood entirely by the portal vein, com- ing from the umbilical vesicle and the intestine; and all the blood derived from this source must pass through the hepatic circulation before reaching the venous extremity of the heart. But soon afterward the allantois makes its appearance, and be- comes rapidly developed into the placenta ; and the umbilical vein comino- from it joins the omphalo-mesenteric vein in the substance of the liver, and takes part in the formation of the hepatic capillary plexus. As the umbilical vesicle, however, becomes atrophied, and the intestine also remains inactive, while the placenta increases in size and in functional importance, a time soon arrives when the liver receives more blood by the umbilical vein than by the portal vein. (Fig. 244.) The umbilical vein then passes into the liver at DEVELOPMENT OF THE HEPATIC CIRCULATION. 577 Hepatic CiRCULATioif further advanced. — 1. Portal vein. 2. Umbilical vein. 3. Hepatic vein. the longitudinal fissure, and supplies the left lobe entirely with its own branches. To the right it sends off a large branch of com- munication, which opens into the portal vein, and partially supplies the right lobe witb umbilical blood. The liver is thus supplied with blood from two different sources, the most abundant of which is the umbilical vein ; and all the blood entering the liver circulates, as before, through its capillary vessels. But we have already seen that the liver is much larger in proportion to the entire body at an early period of foetal life than in the later months. In the foetal pig, when very young, it amounts to nearly twelve per cent, of the weight of the whole body; but before birth it diminishes to seven, six, and even three or four per cent. For some time, therefore, previous to birth, there is mucb more blood returned from the placenta than is required for the capillary circulation of the liver. Accordingly, a vascular duct or canal is formed in its interior, by which a portion of the placental blood is carried directly through the organ, and conveyed to the heart Avithout having passed through the hepatic capillaries. This duct is called the Ductus venosus. The ductus venosus is formed by a gradual dilatation of one of the he- patic capillaries at (s) (Fig. 245), which, enlarging excessively, be- comes at last converted into a wide canal, or branch of communication, passing directly from the umbilical vein below to tlie hepatic vein above. The circulation through the liver, thus established, is as follows : A certain quantity of venous blood still enters through the portal vein (i), and circulates in a part of the capil- lary system of the right lobe. The umbilical vein (2), bringing a much larger quantity of blood, enters the liver also, a little to the 37 Hepatic CiRCDLATioN during lat- ter part of foetal life. — 1. Portal vein. 2. Umbilical vein. 3. Left branch of umbili- cal vein. 4. Right branch of umbilical vein. 5. Ductus venosus. 6. Hepatic vein. 578 DEVELOPMENT OF THE CIRCULATORY APPARATUS. left, and the blood which it contains divides into three principal streams. One of them passes through the left branch (3) into the capillaries of the left lobe ; another turns oft' through the right branch (4), and, joining the blood of the portal vein, circulates through the capillaries of the right lobe ; while the third passes directly onward through the venous duet (5), and reaches the he- patic vein without having passed through any part of the capillary plexus. This condition of the hepatic circulation continues until birth. At that time, two important changes tai^e place. First, the pla- cental circulation is altogether cut off; and secondly, a much larger quantity of blood than before begins to circulate through the lungs and the intestine. The superabundance of blood, previously coming from the placenta, is now diverted into the lungs ; while the intes- tinal canal, entering upon the active performance of its functions, becomes the sole source of supply for the hepatic venous blood. The following changes, therefore, take place at birth in the vessels of the liver. (Fig. 246.) First, the umbilical vein shrivels and becomes converted into a solid rounded cord (2). This cord may be seen, in the adult condition, running from the internal surface of the abdominal walls, at the umbilicus, to the longitudinal fissure of the liver. It is then known under the name of the round ligament. Secondly, the ductus venosus also becomes ob- literated, and converted into a fibrous cord. Thirdly, the blood entering the liver by the portal vein ( i ), passes off by its right branch, as before, to the right lobe. But in the branch (4), the course of the blood is reversed. This was formerly the right branch of the umbilical vein, its blood passing in a direction from left to right. It now becomes the left branch of the portal vein ; and its blood passes from right to left, to be distributed to the capil- laries of the left lobe. According to Dr. Guy, the umbili- cal vein is completely closed at the end of the fifth day after birth. Development of the Hearty and the Ductus Arteriosus. — When the Yxs.. 246. Adult form of Hepatic Ciuccla- TION. — 1. Portal veiu. 2. Obliterated umbilical vein, forming the round liga ment ; the continuation of the dotted lines through the liver shoves the situa- tion of the obliterated ductus venosus. 3. Hepatic vein. 4. Left branch of portal vein. DEVELOPMENT OF THE HEART. 579 embryonic circulation is first established, the heart is a simple tubu- lar sac (Fig. 247), receiving the veins at its lower extremity, and giving off' the arterial trunks at its upper extremity. By the pro- gress of its growth, it soon becomes twisted upon itself; so that the entrance of the veins, and the exit of the arteries, come to be placed more nearly upon the same horizontal level (Fig. 248) ; but the entrance of the veins (i) is behind and a little below, while the exit of the arteries (2) is in front and a little above. The heart is, at this time, a simple twisted tube; and the blood passes through it in a single continuous stream, turning upon itself at the point of curvature, and passing directly out by the arterial orifice. Fig. 247. 2 Earliest form of F(etal Heart. — 1. venous ex- tremity. 2. Arterial ex- tremity. Fis. 248. FffiTAL Heart, twi.sted upon itself. — 1. Venous ex- tremity. 2. Arterial ex- tremitv. FfKfAL Heart, divided into right and left cavities. — 1. Venous extremity. 2. Arterial extremity. 3, 3. Pulmonary branches. Soon afterward, this single cardiac tube is divided into two paral- lel tubes, right and left, by a longitudinal partition, which grows from the inner surface of its walls and follows the twisted course of the organ itself. (Fig. 249.) This partition, which is indicated in the figure by a dotted, line, extends a short distance into the commencement of the primitive arterial trunk, dividing it into two lateral halves, one of which is in communication with the right side of the heart, the other with the left. About the same time, the pulmonary branches (3,3) are given off" from each side of the arterial trunk near its origin ; and the longitudinal partition, above spoken of, is so placed that both these branches fall upon one side of it, and are both, consequently, given off' from that division of the artery which is connected with the right side of the heart. Very soon a superficial line of demarcation, or furrow, shows itself upon the external surface of the heart, corresponding in situa- tion with the internal septum ; while at the root of the arterial trunk, this furrow becomes much deeper, and finally the two lateral portions of the vessel are separated from each other altogether, in 580 DEVELOPMENT OF THE CIKCULATORY APPARATUS. F(ETAL Heart still far- ther developed. — 1. Aorta. 2. Pulmonary artery. 3, 3. Pulmonary branches. 4. Ductus arteriosus. Fig. 250. the immediate neighborhood of the heart, joining again, however, a short distance beyond the origin of the pulmonary branches. (Fig. 250.) It then becomes evident that the left lateral division of the arterial trunk is the commencement of the aorta (i); while its right lateral division is the trunk of the pulmonary artery (2), giving off the right and left pulmo- nary branches (3, 3), at a short distance from its origin. That portion of the pulmonary trunk (4) which is beyond the origin of the pulmonary branches, and which communicates freely with the aorta, is the Ductus arteriosus. The ductus arteriosus is at first as large as the pulmonary trunk itself; and nearly the whole of the blood, coming from the right ventricle, passes directly onward through the arterial duct, and enters the aorta without going to the lungs. But as the lungs gradually become developed, they require a larger quantity of blood for their nutrition, and the pulmonary branches increase in proportion to the pulmonary trunk and the ductus arteriosus. At the termination of foetal life, in the human subject, the ductus arteriosus is about as large as either one of the pulmonary branches; and a very considerable portion of the blood, therefore, eoming from the right ventricle still passes onward to the aorta without being distributed to the lungs. But at the period of birth, the lungs enter upon the active per- formance of the function of respiration, and immediately require a much larger supply of blood. The right and left pulmonary branches then enlarge, so as to become the two principal divis- ions of the pulmonary trunk. (Fig. 251.) The ductus arteriosus at the same time becomes contracted and shrivelled to such an extent that its cavity is obliterated ; and it is finally converted into an im- heabt of Infant, showing pcrvious, rouudcd cord, wliich remains disappearance of arterial duct after yxn\i\ adult life, rUUniug frOm the poiut birth.— 1. Aorta. 2. Pulmonary ar- p • r- i i tery. 3, 3. Pulmonary branches. 4. of bifurcatiou of the pulmouary artery Ductus arteriosus becoming oblite- ^^ ^^^ ^^^j^^ gj^^, ^f ^^iQ arch of the rated. Fig. 2^1. DEVELOPMENT OF THE HEART, 581 aorta. The obliteration of the arterial duct is complete, at latest, by the tenth week after birth. (Gruy.) The two auricles are separated from the two ventricles by hori- zontal septa which grow from the internal surface of the cardiac walls ; but these septa remaining incomplete, the auriculo-ventricu- lar orifices continue pervious, and allow the free passage of the blood from the auricles to the ventricles. The interventricular septum, or that which separates the two ventricles from each other, is completed at a very early date ; but the interauricular septum, or that which is situated between the two auricles, remains incomplete for a long time, being perforated by an oval-shaped opening, the foram£n ovale, allowing, at this situation, a free passage from the right to the left side of the heart. The existence of the foramen ovale and of the ductus arteriosus gives rise to a peculiar crossing of the streams of blood in passing through the heart, which is characteristic of foetal life, and which may be described as follows : — It will be found upon examination that the two venae cavae, superior and inferior, do not open into the auricular sac on the same plane or in the same direction ; for while the superior vena cava is situated anteriorly, and is directed downward and forward, the inferior is situated quite posteriorly, and passes into the auricle in a direction from right to left, and nearly transverse to the axis of the heart. A nearly vertical curtain or valve at the same time hangs downward behind the orifice of the superior vena cava and in front of the orifice of the inferior. This curtain is formed by the lower edge of the septum of the auricles, which, as we have before stated, is incomplete at this age, and which terminates inferiorly and toward the right in a crescentic border, leaving at that part an oval opening, the foramen ovale. The stream of blood, coming from the superior vena cava, falls accordingly in front of this curtain, and passes directly downward, through the auriculo- ventricular orifice, into the right ventricle. But the inferior vena cava, being situated farther back and directed transversely, opens, properly speaking, not into the right auricle, but into the left ; for its stream of blood, falling behind the curtain above mentioned, passes across through the foramen ovale directly into the cavity of the left auricle. This direction of the current of blood, coming from the inferior vena cava, is further secured by a peculiar mem- branous valve, which exists at this period, termed the Eustachian 582 DEVELOPMENT OF THE CIRCULATORY APPARATUS. valve. This valve, which is very thin and transparent (Fig. 252,/), is attached to the anterior border of the orifice of the inferior vena cava, and terminates by a crescentic edge, directed toward the left ; the valve, in this way, standing Fig. 252. as an incomplete membranous partition between the cavity of the inferior vena cava and that of the right auricle. A bougie, accordingly, placed in the in- ferior vena cava, as shown in Fig. 252, lies naturally quite behind the Eustachian valve, and passes directly through the foramen ovale into the left auricle. The two streams of blood, therefore, coming from the su- perior and inferior venae cavse, cross each other upon entering the heart. This crossing of the streams does not take place, however, as it is sometimes described, in the cavity of the right auricle; but, owing to the peculiar position and direction of the two veins at this period, witb regard to the septum of the auricles, the stream coming from the superior vena cava enters the right auricle exclusively, while that from the inferior passes almost directly into the left auricle. It will also be seen, by examining the positions of the aorta, pul- monary artery, and ductus arteriosus, at this time, that the arteria innominata, together with the left carotid and left subclavian, are given off from the arch of the aorta, before its junction with the ductus arteriosus, and this arrangement causes the blood of the two venae cavse, not only to enter the heart in different directions, but also to be distributed, after leaving the ventricles, to different parts of the body. (Fig. 253 ) For the blood of the superior vena cava passes through the right auricle downward into the right ventricle, thence through the pulmonary artery and ductus arteriosus, into the thoracic aorta, while the blood of the inferior vena cava, enter- Heart of Human Fcetus, attlie end of the sixth month ; from a specimen in tlie author's pos- session. — a. Inferior vena cava. h. Superior vena cava. c. Cavity of right auricle, laid open from the front d. Appendix auricularis. e. Cavity of right ventricle, also laid open. /. Eustachian valve. The bougie, which is placed in the inferior vena cava,, can be seen passing behind the Eustachian valve, just below the point indicated by f, then crossing behind the cavity of the right auricle, and passing through the foramen ovale, to the left side of the heart. DEVELOPMENT OF THE HEART. 583 Fisr. 253. ing the left auricle, passes into the left ventricle, thence into the arch of the aorta, and is distributed to the head and upper extremities, before reaching the situation of the arterial duct. The two streams, therefore, in passing through the heart, cross each other both behind and in front. The venous blood, returning from the head and upper extremities by the superior vena cava, passes through the abdo- minal aorta and the umbilical arte- ries, to the lower part of the body, and to the placenta; while that re- turning from the placenta, by the inferior vena cava, is distributed to the head and upper extremities, through the vessels given off from the arch of the aorta. This division of the streams of blood, during a certain period of foetal life, is so complete that Dr. John Reid,^ on injecting the infe rior vena cava with red, and the superior with yellow, in a seven months' human foetus, found that the red had passed through the foramen ovale into the left auricle and ventricle and arch of the aorta, and had filled the vessels of the head and upper extremities : while the yellow had passed into the right ventricle, pulmonary artery, ductus arteriosus, and thoracic aorta, with only a slight ad- mixture of red at the posterior part of the right auricle. All the branches of the thoracic and abdominal aorta were filled with yel- low, while the whole of the red had passed to the upper part of the body. We have repeated the above experiment several times on the foetal pig, when about one-half and three-quarters grown, first taking the precaution to wash out the heart and large vessels with a wa- tery injection, immediately after the removal of the foetus from the body of the parent, and before the blood had been allowed to coagu- late. The injections used were blue for the superior vena cava, and yellow for the inferior. The two syringes were managed, at the same time, by the right and left hands; their nozzles being firmly held in place by the fingers of an assistant. When the Diagram of CiRcaLATioN through THE FcETAL HEART. — rt. Superior veaa cava. 6. Inferior vena cava. c,c,c,o Arch of aorta and its branches, d. Pulmonary artery. ' Edinburgh Medical and Surgical Journal, vol. xliii. 1835. 584 DEVELOPMENT OF THE CIECULATORY APPARATUS. points of the syringes were introduced into the veins, at equal dis- tances from the heart, and the two injections made with equal force and rapidity, it was found that the admixture of the colors which took place was so slight, that at least nineteen-twentieths of the yellow injection had passed into the left auricle, and nineteen-twentieths of the blue into the right. The pulmonary artery and ductus arteriosus contained a similar proportion of blue, and the arch of the aorta of yellow. In the thoracic and abdominal aorta, however, contrary to what was found by Dr. Eeid, there was always an admixture of the two colors, generally in about equal proportions. This discrepancy may be owing to the smaller size of the head and upper extremities, in the pig, as compared with those of the human subject, which would prevent their receiving all the blood coming from the left ventricle ; or to some differences in the manipulations of these experiments, in which it is not always easy to imitate exactly the force and ra- pidity of the different currents of blood in the living foetus. The above results, however, are such as to leave no doubt of the prin- cipal fact, viz., that up to an advanced stage of foetal life, by far the greater portion of the blood coming from the inferior vena cava passes through the foramen ovale, into the left side of the heart ; while by far the greater portion of that coming from the head and upper extremities passes into the right side of the heart, and thence outward by the pulmonary trunk and ductus arteriosus. Toward the latter periods of gestation, this division of the venous currents becomes less complete, owing to the three following causes: — First, the lungs increasing in size, the two pulmonary arteries, as well as the pulmonary veins, enlarge in proportion; and a greater quantity of the blood, therefore, coming from the right ventricle, instead of going onward through the ductus arteriosus, passes to the lungs, and returning thence by the pulmonary veins to the left auricle and ventricle, joins the stream passing out by the arch of the aorta. Secondly, the Eustachian valve diminishes in size. This valve, which is very large and distinct at the end of the sixth month (Fig. 252), subsequently becomes atrophied to such an extent that, at the end of gestation, it has altogether disappeared, or is at least reduced to the condition of a very narrow, almost imperceptible membranous ridge, which can exert no influence on the direction of the current of blood passing by it. Thus, the cavity of the supe- rior vena cava, at its upper extremity, ceases to be separated from DEVELOPMENT OF THE HEART. 585 that of the right auricle; and a passage of blood from one to the other may, therefore, more readily take place. Thirdly, the foramen ovale becomes partially closed by a valve which passes across its orifice from behind forward. This valve, which begins to be formed at a very early period, is called the valve of the foramen ovale. It consists of a thin, fibrous sheet, which grows from the posterior surface of the auricular cavity, just to the left of the foramen ovale, and projects into the left auricle, its free edge presenting a thin crescentic border, and being attached, by its two extremities, to the auricular septum upon the left side. This valve does not at first interfere at all with the flow of blood from right to left, since its edge hangs freely and loosely into the cavity of the left auricle. It only opposes, therefore, during the early periods, any accidental regurgitation from left to right. But as gestation advances, while the walls of the heart con- tinue to enlarge, and its cavities to expand in every direction, the fibrous bundles, forming the valve, do not elongate in proportion. The valve, accordingly, becomes drawn downward more and more toward the foramen ovale. It thus comes in contact with the edges of the interauricular septum, and unites with its substance; the adhesion taking place first at the lower and posterior portion, and proceeding gradually upward and forward, so as to make the pas- sage, from the right auricle to the left, more and more oblique in direction. At the same time, an alteration takes place in the position of the inferior vena cava. This vessel, which at first looked transversely toward the foramen ovale, becomes directed more obliquely for- ward ; so that, the Eustachian valve having mostly disappeared, a part of the blood of the inferior vena cava enters the right auricle, while the remainder still passes through the equally oblique open- ing of the foramen ovale. At the period of birth a change takes place, by which the foramen ovale is completely occluded, and all the blood coming through tiie inferior vena cava is turned into the right auricle. This change depends upon the commencement of respiration. A much larger quantity of blood than before is then sent to the lungs, and of course returns from them to the left auricle. The left auricle, being then completely filled wath the pulmonary blood, no longer admits a free access from the right auricle through the foramen ovale; and the valve of the foramen, pressed backward more closely against the edges of the septum, becomes after a time 586 DEVELOPMENT OF THE CIRCULATORY APPARATUS. adherent throughout and obliterates the opening altogether. The cutting off of the placental circulation diminishes at the same time the quantity of blood arriving at the heart by the inferior vena cava. It is evident, indeed, that the same quantity of blood which previously returned from the placenta by the inferior cava, on the right side of the auricular septum, now returns from the lungs, by the pulmonary veins upon the left side of the same septum ; and it is owing to all these circumstances combined, that while before birth a portion of the blood always passed from the right auricle to the left through the foramen ovale, no such passage takes place after birth, since the pressure is then equal on both sides of the auricular septum. The foetal circulation, represented in Fig. 253, is then replaced by the adult circulation, represented in Fig. 254. Fig. 254. Diagram of Adult Circulation throuuh the Heart — a, a. Superior and inferior venae cav33. b. Right ventricle, c. Pulmonary artery, dividing into right and left branches, d. Pulmo- nary vein. e. Left ventricle. /. Aorta. That portion of the septum of the auricles, originally occupied by the foramen ovale, is accordingly constituted, in the adult con- dition, by the valve of the foramen ovale, whicb has become adhe- rent to the edges of the septum. The auricular septum in the adult heart is, therefore, thinner at this spot than elsewhere; and presents, on the side of the right auricle, an oval depression, termed the fossa DEVELOPMENT OF THE HEART. 587 ovalis^ which indicates the site of the original foramen ovale. The fossa ovalis is surrounded by a slightly raised ring, the annulus ovalis, representing the curvilinear edge of the original auricular septum. The foramen ovale is sometimes completely obliterated within a few daj^s after birth. It often, however, remains partially pervious for several weeks or months. We have a specimen, taken from a child of one year and nine months, in which the opening is still very distinct; and it is not unfrequent to find a small apertare ex- isting even in adult life. In these instances, however, though the adhesion and solidification of the auricular septum is not complete, no disturbance of the circulation results, and no admixture of blood takes place between the right and left sides of the heart ; since the passage through the auricular septum is always very oblique in its direction, and its valvular arrangement prevents any regurgitation from left to right, while the complete filling of the left auricle with pulmonary blood, as above mentioned, equally opposes any passage from right to left. 588 DEVELOPMENT OF THE BODY AFTER BIRTH. CHAPTER XVIII. DEVELOPMENT OF THE BODY AFTER BIRTH. The newly-born infant is still very far from having arrived at a state of complete development. The changes which it has passed through during intra-uterine life are not more marked than those which are to follow during the periods of infancy, childhood, and adolescence. The anatomy of the organs, both internal and ex- ternal, their physiological functions, and even the morbid derange- ments to which they are subject, continue to undergo gradual and progressive alterations, throughout the entire course of subsequent life. The history of development extends, properly speaking, from the earliest organization of the embryonic tissues to the complete formation of the adult body. The period of birth, accordingly, marks only a single epoch in a constant series of changes, some of which have preceded, while many others are to follow. The weight of the newly-born infant is a little over six pounds. The middle point of the body is nearly at the umbilicus, the head and upper extremities being still very large, in proportion to the lower extremities and pelvis. The abdomen is larger and the chest smaller in proportion than in the adult. The lower extremi- ties are curved inward, as in the foetal condition, so that the soles of the feet look obliquely toward each other, instead of being directed horizontally downward, as at a subsequent period. Both upper and lower extremities are habitually curled upward and forward over the chest and abdomen, and all the joints are constantly in a semi-flexed position. The process of respiration is very imperfectly performed for some time after birth. The expansion of the pulmonary vesicles, and the changes in the circulatory apparatus described in the pre- ceding chapter, far from being sudden and instantaneous, are always more or less gradual in their character, and require an interval of several days for their completion. Respiration, indeed, seems to be accomplished, during this period, to a considerable DEVELOPMENT OF THE BODY AFTER BIRTH. 589 extent through the skin, which is remarkably soft, vascular, and ruddy in color. The animal heat is also less actively generated than in the adult, and requires to be sustained by careful protec- tion, and by contact with the body of the mother. The young infant sleeps during the greater part of the time; and even when awake there are but few manifestations of intelligence or percep- tion. The special senses of sight and hearing are dull and inex- citable, though their organs are perfectly formed ; and even consciousness seems present only to a very limited extent. Volun- tary motion and sensation are nearly absent ; and the almost con- stant irregular movements of the limbs, observable at this time, are evidently of a reflex or automatic character. Nearly all the nervous phenomena, indeed, presented by the newly-born infant, are of a similar nature. The motions of its hands and feet, the act of suckling, and even its cries and the contortions of its face, are reflex in their origin, and do not indicate the existence of any active volition, or any distinct perception of external objects. There is at first but little nervous connection established with the external world, and the system is as yet almost exclusively occu- pied with the functions of nutrition and respiration. This preponderance of the simple reflex actions in the nervous system of the infant, is observable even in the diseases to which it is peculiarly subject for some years after birth. It is at this age that convulsions from indigestion are of most frequent occurrence, and even temporary strabismus and paralysis, resulting from the same cause. It is well known to physicians, moreover, that the effect of various drugs upon the infant is very different from that which they exert upon the adult. Opium, for example, is very much more active, in proportion to the dose, in the infant than in the adult. Mercury, on the other hand, produces salivation with greater difficulty in the former than in the latter. Blisters excite more constitutional irritation in the young than in the old subject; and antimony, when given to children, is proverbially uncertain and dangerous in its operation. The difference in the anatomy of the newly -born infant, and that of the adult, may be represented, to a certain extent, by the fol- lowing list, which gives the relative weight of the most important internal organs at the period of birth and that of adult age; the weight of the entire body being reckoned, in each case, as 1000. The relative weight of the adult organs has been calculated from 590 DEVELOPMENT OF THE BODY AFTER BIRTH. the estimates of Cruveilhier, Solly, Wilson, &c,; that of the organs in the foetus at term from our own observations. FcETDS AT Term. Adult. Weight of the entire body . 1000.00 1000.00 " encephalon . 148.00 23.00 " liver . 37.00 29.00 " heart . 7.77 4.17 " kidneys 6.00 4.00 " renal cap ules l.t)3 0.13 " thyroid gland 0.60 0.51 " thymus gland 3.00 0.00 will be observed that most of the internal orcra ns dimini relative size after birth, owing principally to the increased develop- ment of the osseous and muscular S3'-stems, both of which are in a very imperfect condition throughout intra-uterine life, but which come into activity during childhood and youth. Within the first day after birth the remains of the umbilical cord begin to wither, and become completely desiccated by about the third day. A superficial ulceration then takes place about the point of its attachment, and it is separated and thrown off within the first week. After the separation of the cord, the umbilicus becomes completely cicatrized by the tenth or twelfth day after birth. (Guy.) An exfoliation and renovation of the cuticle also take place over the whole body soon after the birth. According to Kolliker, the eyelashes, and probably all the hairs of the body and head, are thrown off and replaced by new ones, within the first year. The teeth in the newly-born infant are but partially developed, and are still inclosed in their follicles, and concealed beneath the gums. They are twenty in number; viz., two incisors, one canine, and two molars, on each side of each jaw. At birth there is a thin layer of dentine and enamel covering their upper surfaces, but the body of the tooth and its fangs are formed subsequently by progressive elongation and ossification of the tooth-pulp. The fully-formed teeth emerge from the gums in the following order. The central incisors in the seventh month after birth ; the lateral incisors in the eighth month; the anterior molars at the end of the first year; the canines at a year and a half; and the second molars at two years (Kolliker). The eruption of the teeth in the lower jaw generally precedes by a short time that of the corresponding teeth in the upper. During the seventh year a change begins to take place by which DEVELOPMENT OF THE BODY AFTER BIRTH. 591 the first set of teeth are thrown oft' and replaced by a second or permanent set, differing in number, size, and shape from those which preceded. The anterior permanent molar first shows itself just behind the posterior temporary molar, on each side. This happens at about six and a half years after birth. At the end of the seventh year the middle incisors are thrown oft" and replaced by corresponding permanent teeth, of larger size. At the eighth year a similar exchange takes place in the lateral incisors. In the ninth and tenth years, the anterior and second molars are replaced by the anterior and second permanent bicuspids. In the twelfth year, the canine teeth are changed. In the thirteenth year, the second permanent molars show themselves; and from the seven- teenth to the twenty-first year, the third molars, or " wisdom teeth," emerge from the gums, at the posterior extremities of the dental arch. (Wilson.) The jaw, therefore, in the adult condition, contains three teeth on each side more than in childhood, making in all thirty-two permanent teeth ; viz., on each side, above and below, two incisors, one canine, two bicuspids, and three permanent molars. The entire generative apparatus, which is still altogether inactive at birth, begins to enter upon a condition of functional activity from the fifteenth to the twentieth year. The entire configuration of the body alters in a striking manner at this period, and the dis- tinction between the two sexes becomes more complete and well marked. The beard is developed in the male ; and in the female the breasts assume the size and form characteristic of the condition of puberty. The voice, which is shrill and sharp in infancy and childhood, becomes deeper in tone, and the countenance assumes a more sedate and serious expression. After this period, the mus- cular system increases still further in size and strength, and the consolidation of the skeleton also continues; the bony union of its various parts not being entirely accomplished until the twenty-fifth or thirtieth year. Finally, all the different organs of the body arrive at the adult condition, and the entire process of development is then complete. INDEX. Absorption, 128 by bloodvessels, 131 by lacteals, 133 of fat, 137 of oxygen in respiration, 207 by egg during incubation, 508 of calcareous matter by allantois, 508 Absorbent glands, 134 vessels, 133-136 Acid, carbonic, 206-217 lactic, in gastric juice, 107 in souring milk, 277 glyko-cliolic, 146 tauro-cholic, 147 pneumic, 211 uric, 288-296 oxalic, in urine, 301 Acid fermentation of urine, 300 Acidity, of gastric juice, cause of, 107 of urine, 295 Acini, of liver, 279-280 Adipose vesicles, 58 digestion of, 126-127 Adult circulation, 570 establishment of, 585 Aerial respiration, 198-200 Age, influence of, on exhalation of car- bonic acid, 215 on comparative weight of organs, 590 Air, alterations of, in respiration, 206 circulation of, in lungs, 204 Air-cells of lungs, 200 Air-chamber, in fowl's egg, 454 Albumen, 68 of the blood, 189 in saliva, 93 in milk, 276 of the egg, how produced, 453 its liquefaction and absorption dur- ing development of foetus, 504-506 Albuminoid substances, 63 digestion of, 110 Albuminose, 110 interference with Trommer's test, 111 with action of iodine and starch, 112 Alimentary canal, in different animals, 84-88 development of, 546. Alkalies, effect of, on urine, 296 Alkaline chlorides, 39-42 phosphates, 45 carbonates, 44-45 Alkaline fermentation of urine, 302 Alkalescence of blood, due to carbonates, 44 Allantois, 501 formation of, 503 in fowl's egg, 506 function of, 507 in foetal pig, 524 Alligator, brain of, 322 Amnion, 501, formation of, 502 enlargement of, during latter part of pregnancy, 532 contact with chorion, 533 Amniotic folds, 502 Amniotic fluid, 532 its use, 533 contains sugar at a certain period,. 551 Amniotic umbilicus, 502 Analysis, of animal fluids, 32-33 of milk, 80 of wheat flour, 80 of oatmeal, 80 of eggs, 81 of meat, 81 of saliva, 92 of gastric juice, 107 of pancreatic juice, 123 of bile, 142 of blood-globules, 183 of blood-plasma, 188 of mucus, 269 of sebaceous matter, 270 of perspiration, 272 of milk, 275 of butter, 277 of urine, 294 Anoral and Gavakret, production of carbonic acid in respiration, 214 38 594 INDEX. Animal functions, 27 Animal heat, 218 in different species, 220 mode of generation, 222-228 influenced by local causes, 226 in diffei'ent organs, 227 increase of, after section of sympa- thetic nerve, 421-422 Animal and vegetable parasites, 434 Animalcules, infusorial, 431 mode of production, 432 Annulus ovalis, 586 Anterior columns of spinal cord, 321 their excitability, 343 Aorta, action of, 247 development of, 571 Aplysia, nervous system of, 315 Appetite, disturbed by anxiety, &c., 117 necessary to digestion of food, 118 Aquatic respiration, 197-198 Area pellucida, 492 vasculosa, 505-567 Arch of aorta, formation of, 571-572 Arches, cervical, 570 transformation of, 571 Arteries, 246 motion of blood in, 247 pulsation of, 248 elasticity of, 246, 249 rapidity of circulation in, 250-251 omphalo-mesenteric, 567 vertebral, 570 umbilical, 570 Arterial system, development of, 570- 579 Articulata, nervous system of, 316 reflex action in, 317 Articulation of tapeworm, 443 Arytenoid cartilages, 205 movements of, 206 Assimilation, 265 destructive, 282 Auricle, single, of fish, 230 double, of reptiles, birds, and mam- malians, 231-232 contraction of, 245 Auriculo-ventricular valves, action of, 234 Auditory nerves, 386 Axis-cylinder, of nervous filaments, 308-310 Aztec children, 366 Azygous veins, formation of, 575 Beaumont, Dr., experiments on Alexis St. Martin, 103-114 Beknard, on efi'ect of dividing Steno's duct, 99 on digestion of fat in intestine, 123 on formation of liver-sugar, 166, 167, 168 on decomposition of bicarbonates in lung, 212 Bernard, on temperature of blood in dif- ferent organs, 2"i7 Bidder and Schmidt, on daily quantity of bile, 153 on effect of excluding bile from in- testine, 160 on reabsorption of bile, 161 Bile, 141 composition of, 142 tests for, 150 daily quantity of, 153 functions of, 158 reabsorption, 161 mode of secretion, 278 Biliary salts, 143 of human bile, 149 Biliverdine, 71, 142 tests for, 150 passage into the urine, 299 Blastodermic membrane, 490 Blood, 178 red globules of, 179 white globules, 185 plasma, 188 coagulation of, 190 bully coat, 195 entire quantity of, 196 alterations of, in respiration, 207 temperature of, 219 in different organs, 227 circulation of, 229 through the heart, 235 through the arteries, 246 through the veins, 252 through the capillaries, 255 Boussingault, on chloride of sodium in food, 41 on internal production of fat, 61 Brain, 357 of alligator, 322 of rabbit, 323 human, 326 remarkable cases of injury to, 360 size of, in different races, 364 in idiots, 366 development of, 540-541 Branchiae, 197 of meno-branchus, 198 Broad ligaments, formation of, 563 Bronchi, division of, 200 ciliary motion in, 204 Brunner's glands, 120 Buffy coat of the blood, 195 Butter, 276 composition of, 277 condition in milk, 59, 277 Butyrine, 277 Canals of Cuvier, 573 Capillaries, 255 their inosculation, 256 motion of, blood in, 257 Capillary circulation, 256 INDEX. 595 Capillary circiUation, causes of, 258 how modified by inflammation, 259 rapidity of, 260 peculiarities of, in diflferent parts, 262, 264 Caput coli, formation of, 547 Carbonic acid, in the breath, 206 proportion of, to oxygen absorbed, 207 in the blood, 208 origin of, in lungs, 211 in the blood, 212 in the tissues, 212 mode of production, 213 daily quantity of, 214 variations of, 215 exhaled by skin, 217 by egg, during incubation, 508 absorbed by vegetables, 225 Carbonate of lime, 44 of soda, 44 of potass, 45 of ammonia, in putrefying urine, 302 Cardiac circulation, in foetus, 583 in adult, 586 Carnivorous animals, respiration of, 18, 207 urine of, 287, 289 Cartilagine, 70 Caseine, 68 Cat, secretion of bile in, 154 closure of eyelids, after division of sympathetic, 423 Catalytic action, 66 of pepsin, 110 Centipede, nervous system of, 316 Centre, nervous, definition of, 313 Cerebrum, 359. See Hemispheres. Cerebral ganglia, 322-357. See Hemi- spheres. Cerebellum, 370 effects of injury to, 371 removal of, 371, 372 function of, 370-373 development of, 540-541 Cerebro-spinal system, 318-319 development of, 539 Cervix uteri, 456 in fffitus, 564 Cervical arches, 570 transformation of, 571 Changes, in egg, while passing through oviduct, 450, 453, 488 in hepatic circulation, at birth, 578 in comparative size of organs, after birth, 590 Chick, developmfent of, 504-509 Children, Aztec, 366 Chloride of sodium, 39 its proportion in the animal tissues and fluids, 40 importance of, in the food, 41 mode of discharge from the body, 41 Chloride of sodium, partial decomposi- tion of, in the body, 42 Chloride of potassium, 42 Cholesterine, 142 Chorda dorsalis, 493 Chordse vocales, movement of, in respi- ration, 205 action of, in the production of vocal sounds, 403 obstruction of glottis by, after divi- sion of pneumogastric, 405 Chorion, formation of, 510 villosities of, 512 source of vascularity of, 513 union with decidua, 521 Chyle, 121-133 in lacteals, 136 absorption of, 137 by intestinal epithelium, 138 in blood, 139 Ciliary motion, in bronchi, 204 in Fallopian tubes, 475 Ciliary nerves, 414 Circulation, 229 in the heart, 222-235 in the arteries, 246 in the veins, 252 in the capillaries, 255 rapidity of, 261 peculiarities of, in difi'erent parts, 263 in liver, 279 in placenta, 527-581 Circulatory apparatus, development of, 566-587 Civilization, aptitude for, of diflferent races, 364 Classification of cranial nerves, 387 Clot, formation of, 191 separation from serum, 192 bulfed and cupped, 195 Coagulation, 66 "of fibrin, 188 of blood, 190 of white substance of Schwann, in nerve-fibres, 309 Colin, on unilateral mastication, 94 Cold, resistance to, by animals, 218 efi'ect of when long continued, 219 Colostrum, 275 Coloring matters, 70 of blood, 70, 183 of the skin, 171 of bile, 71, 142 of urine, 71 Commissure, of spinal cord, gray, 320 white, 321 transverse, of cerebrum, 327 of cerebellum, 327 Commissures, nervous, 313 olfactory, 322, 385 Congestion, of ear, &c., after division of sympathetic, 422 596 IX TEX. Convolvulus, sexual apparatxis of, 442 Contact, of chorion and amnion, 533 of decidua vera and reflexa, 534 Consentaneons action of muscles, 348 Contraction, of stomacti during digestion, 112 of spleen, 174 of blood-clot, 191 of diaphragm and intercostal mus- cles, 201 of posterior crico arytenoid nmscles, 206 of ventricles, 239 of muscles after death, 328 of sphincter ani, 354 of rectum, 354 of urinary bladder, 355 of pupil, under influence of light, 374, 418 after division of sympathetic, 423 Cookins, effect of, on food, 82 Cord, spinal, 319-340 umbilical, 533 withering and separation of, 590 Corpus callosum, 327 Corpus luteum, 478 of menstruation, 478-482 of pregnancy, 482-487 ■ three weeks after menstruation, 480 four weeks after menstruation, 481 nine weeks after menstruation, 481 at end of second month of preg- nancy, 484 at end of fourth month, 484 at term, 485 disappearance of, after delivery, 486 Corpora Malpighiana, of spleen, 174 Corpora striata, 323, 359 Corpora olivaria, 324 Corpora Wolffiana, 556-563 CosTE, on rupture of Graafian follicle in menstruation, 474, 475 Cranial nerves, 385 classification of, 387 motor, 388 sensitive, 393 Creatine, 287-288 Creatinine, 288 Cremaster musc4e, formation of, 560 function of, in lower animals, 561 Crystals, of stearine, 55 and margarine, 56 of cholesterin, 143 of glyko-cholate of soda, 144, 145 of biliary matters of dog's bile, 148, 155 of urea, 285 of creatine, 287 of creatinine, 288 of urate of soda, 289 of uric acid, 296 of oxalic acid, 302 Crystals, of triple phosphate, 304 Crystallizable substances of organic ori- gin, 47 Crossing of fibres in medulla oblongata, 324 of sensitive fibres in spinal cord. 845 of streams of blood in fcetal heart. 582, 583 Cecikshaxk, rupture of Graafiian folUcle in menstruation, 474 Cumulus proligerus, 469 Cutaneous respiration, 217 perspiration, 271 Cuticle, exfoliation of, during fojtal life, 545 after birth, 590 Cysticercus, 439 transformation of into taenia, 440 production of, from eggs of taenia, 441 Death, a necessary consequence of life, 428 Decidua, 516 vera, 518 reflesa, 519 union with chorion, 521 its discharge in cases of abortion, 520 " at the time of delivery, 535 Decussation of anterior columns of spinal cord, 324 of optic nerves, 375, 376 Degeneration, fatty, of muscular fibres of uterus, after delivery, 537 Deglutition, 100 retarded by division of Steno's duct. 99 by division of pneumogastric, 401 Dentition, first, 590 second, 591 Descent of the testicles, 559 of the ovaries, 562 Destructive assimilation, 282 Development of the impregnated egg, of allantois, 503 of chorion, 510 of villosities of chorion, 511, 512 of decidua, 516 of placenta, 525-531 ot nervous system, 539 of eye, 542 of ear, 543 of skeleton, 543 of limbs, 544 of integument, 545 of alimentary canal, 546-548 of urinary passages, 548 of liver, 550 of pharynx and oesophagus, 551 IKDEX. 597 Development of face, 553 of Wolffian bodies, 556 of kidneys, 557 of internal generative organs, 558 of circulatory apparatus, 56b" of arterial system, 570 of venous system, 573 of hepatic circulation, 576 of heart, 578 of the body after birth, 588 Diabetes, 299 in foetus, 551 Diaphragm, action of in breathing, 201, 202 formation of, 552 Diaphragmatic hernia, 553 Diet, influence of on nutrition, 74-76 on products of respiration, 207, 225 on formation of urea, 286 of urate of soda, 289 Diffusion of gases in lungs, 203 Digestion, 83 of starch, 118 of fats, 121, 123 of sugar, 119 of organic substances, 110 time required for, 114 Digestive apparatus of fowl, 85 of ox, 86 of man, 87 Discharge of eggs from ovary, 450-453 independent of sexual intercourse, 467 mechanism of, 470 during menstruation, 474 Discus proligerus, 469 Distinction between corpora lutea of menstruation and pregnancy, 486-487 Diurnal variations, in exhalation of car- bonic acid, 216 in production of urea, 287 in density and acidity of urine, 293 Division of nerves, 311 of heart, into right and left cavities, 579 DoBSON, on variation in size of spleen, 173 Deafer, John C, on production of urea, 286-287 Drugs, effect of, on newly born infant, 589 Ductus arteriosus, 580 closure of. 580-581 venosus, 577 obliteration of, 578 Duodenal glands, 120 fistuU, 156 DuTKOCHET, on temperature of plants, 221 Ear, muscular apparatus of, 420 development of, 530 Earthy phosphates, 42, 45 in urine, 295 Earthy phosphates, precipitated by addi- tion of an alkali, 296 Ectopia cordis, 553 Egg, 442-446 its contents, 447 where formed, 448 of frog, 450 of fowl, 453-454 changes in, while passing through the oviduct, 450-453 pre-existence of, in ovary, 465 development of, at period of puberty, 466 periodical ripening and discharge, 467 discharge of, from Graafian follicle, 470 impregnation of, how accomplished, 463-464 development of, after impregnation, 488 of fowl, showing area vasculosa, 505 ditto, showing formation of allantois, 506 of fish, showing vitelline circulation, 567 attachment of, to uterine mucous membrane, 521 discharge of from uterus, at the time of delivery, 535 condition of in newly born infant, 564 Elasticity, of spleen, 173 of red globules of blood, 181 of lungs, 200 of costal cartilages, 202 of vocal chords, 206 of arteries, 246 Electrical current, effect of, on muscles, 329 on nerve, 331 different effects of direct and in- verse, 338 Electrical fishes, phenomena of, 337 Electricity, no manifestations of in irri- tated nerve, 338 Elevation of temperature, after division of sympathetic, 421, 422 Elongation of heart in contraction, 240 anatomical causes of, 241 Embryo, formation of, 488 Embryonic spot, 492 Encephalon, 321 ganglia of, 326 Endosmosis, of fatty substances, 137 in capillary circulation, 260 Enlargement of amnion, during preg- nancy, 532, 533 Eutozoa, encysted, 436 mode of production, 438 Epithelium, in saliva, 92 of gastric follicles, 102 of intestine, during digestion, 138 598 INDEX. Epidermis, exfoliation of, in foetal life, 545 after birth, 590 Epididymis, 560 Excretion, 282 nature of, 283 importance to life, 283-284 products of, 284 by placenta, 530 Excrementitious substances, 282 mode of formation of, 283 effect of retention of, 283 Exfoliation of cuticle, during fretal life, 545 after birth, 590 Exhalation of watery vapor, 39 from the lungs, 207 from the skin, 271 from the egg, during incubation, 507 of carbonic acid, 214-217 of nitrogen, 206 of animal vapor, 206 Exhaustion, of muscles, by repeated irri- tation, 330 of nerves, by ditto, 332 Expiration, movements of, 202 after section of pneumogastric, 407 Extractive matters of the blood, 190 Eye, protection of, by movements of pupil, 374, 418 by two sets of muscles, 419 Eyeball, inflammation of, after division of 5th pair, 397 Eyelids, formation of, 543 Face, sensitive nerves of, 395 motor nerve, 390 development of, 553 Facial nerve, 390 sensibility of, 391 influence of, on muscular apparatus of eye, 419 of nose, 420 of ear, 420 paralysis of, 391 Fallopian tubes, 455 formation of, 559, 562 Farinaceous substances, 47. in food, 48 digestion of, 118 Fat, decomposition of, in the blood, 137 Fats, 54 proportion of, in dififerent kinds of food, 56 condition, in the various tissues and fluids, 57 internal source of, 61 decomposed in the body, 62 indispensable as ingredients of the food, 74 Fatty matters of the blood, 189 Fatty degeneration of decidua, 536 of muscular fibres of uterus, after delivery, 537 Female generative organs, 446 of frog, 449 of fowl, 453 of sow, 455 of human species, 456 development of, 562 Fermentation, 67 of sugar, 53 acid, of urine, 300 alkaline, of ditto, 302 Fibrin, 68 of the blood, 188 coagulation of, 188 varying quantity of, in blood of dif- ferent veins, 189 Fifth pair of cranial nerves, 394 its distribution, 395 division of, paralyzes sensibility of face, 395 and of nasal passages, 396 produces inflammation of eye- ball, 397 lingual branch of, 398 small root of, 389 Fish, circulation of, 230 mode of progression, 370 formation of umbilical vesicle in, 498 vitelline circulation, in embryo of, 567 Fish, electrical, phenomena of, 337 Fissure, longitudinal, of brain and spinal cord, 319 formation of, 541 Fissure of palate, 555 Fistula, gastric, Dr. Beaumont's case of, 103 mode of operating for, 104 duodenal, 156 Foetal circulation, first form of, 566 . second form of, 568 Follicles, of stomach, 101 of Lieberkiihn, 119 of Brunner's glands, 120 Graafian, 448, 470 of uterus, 517 Food, 73 composition of, 80, 81 daily quantity required, 81 effect of cooking, on, 82 Foramen ovale, 581 valve of, 585 closure of, 585 Force, nervous, nature of, 335 Formation of sugar in liver, 165 in foetus, 551 Fossa ovalis, 586 Functions, animal, 27 vegetative, 27 of teeth, 89 of saliva, 98 of gastric j,uice, 109 of intestinal juices, 118 INDEX. 599 Functions, of pancreatic juice, 123 of bile. 158 of spleen, 175 of nuicus, 2G9 of sebaceous matter, 270 of perspiration, 272 of the tears, 274 Galvanism, action of, on muscles, 329 on nerves, 331 Ganglion, of spinal cord, 348 of tuber annulai-e, 377 of mediilla oblongata, 378 Casserian, 393 of Andersch, 393, 398 pneumogastric, 393, 400 ophthalmic, 414 spheno-palatine, 414 submaxillary, 414 otic, 415 semilunar, 416 impar, 416 Ganglionic system of nerves, 319,414 Ganglia, nervous, 312 of radiata, 313 of moUusca, 315 of articulata, 316 of posterior roots of spinal nerves, 320 of alligator's brain, 322 of rabbit's brain, 323 of medulla oblongata, 324, 378 of human brain, 326 of great sympathetic, 414 Gases, diffusion of, in lungs, 203 absorption and exhalation of, by lungs, 208 by the tissues, 212 Gastric follicles, 101 Gastric juice, mode of obtaining, 105 composition of, 107 action on food, 109 interference with Trommer's test. Ill interference with action of starch and iodine, 112 daily quantity of, 115 solvent action of, on stomach, after death, 117 Gelatine, how produced, 32 effect of feeding animals on, 77 Generation, 429 spontaneous, 429 of infusoria, 432 of parasites, 434 of encysted entozoa, 436 of tsenia, 438 sexual, by germs, 442 Germ, nature of, 442 Germination, heat produced in, 221 Germinative vesicle, 447 disappearance of, in mature egg, 488 Germinative spot, 447 Gills, of fish, 197 Gills, of menobranchus, 198 Glands, of Brunner, 120 mesenteric, 134 vascular, 175 Meibomian, 270 perspiratory, 271 action of, in secretion, 265 Glandulse, solitariae and agminatse, 128 Globules, of blood, 178 red, 179 different appearances of, under microscope, 179, 180 mutual adhesion of, 180 color, consistency, and structure of, 181 action of water on, 182 composition of, 183 size, &c., in different animals, 184,185 white, 185 action of acetic acid on, 186 red and white, movement of, in circulation, 257 Globuline, 68, 183. Glomeruli, of Wolffian bodies, 557 Glosso -pharyngeal nerve, 398 action of, in swallowing, 399 Glottis, movements of, in respiration, 204 in formation of voice, 402 closure of, after section of pneumo- gastrics, 405 Glycine, 146 Glyco-cholic acid, 146 Glyco-cholate of soda, 146 its crystallization, 144 Glycogenic function of liver. 165 in foetus, 551 Glycogenic matter, 169 its conversion into sugar, 170 Graafian follicles, 448 structure of, 469 rupture of, and discharge of egg, 470 ruptured during menstruation, 474 condition in foetus at term, 564 Gray substance, of nervous system, 312 of spinal cord, 320 of brain, 326 its want of irritability, 360 Great sympathetic, 319-414 anatomy of, 415 sensibility and excitability of, 417 connection of, with special senses, 418, 421 division of, influence on animal heat, 422 on pupil and eyelids, 423 reflex actions of, 424 Gubernaculum testis, 560 function of, in lower animals, 561 Gustatory nerve, 398 Hammoxd, Dr. Wm. A., on effects of non-nitrogenous diet, 76 600 INDEX. Hammoxd, on production of urea, 2^6 Hsematine, 70, 183 Hairs, formation of, in embryo, 545 Hare-lip, 554 Harvey, on motions of heart. 238, 240, 245 Heart, 230 of fish, 230 of reptiles, 231 of mammalians, 232 of man, 233 circulation of blood through, 235 sounds of, 235 movements of, 238 impulse, 244 development of, 578 Heat, vital, of animals, 218 of plants, 221 how produced, 222 increased by division of sympathetic nerve, 422 Hemispheres, cerebral, 322, 359 remarkable cases of injury to, 360 eifect of removal, on pigeons, 361- 362 effect of disease, in man, 363 comparative size of, in different races , 364 functions of, 365 development of, 540 Hemorrhage, from placenta, in parturi- tion, 535 Hepatic circulation, 279 development of, 576 Herbivorous animals, respiration of, 18, 207 urine of, 287, 289 Hernia, congenital, diaphragmatic, 553 umbilical, 548 Hernia, inguinal, 562 Hippurate of soda, 289 Hunger and thirst, continue after divi- sion of pneumogastric, 411 Hydrogen, displacement of gases in blood by, 209 exhalation of carbonic acid in an atmosphere of, 213 Hygroscopic property of organic sub- stances, 65 Hypoglossal nerve, 392. See Sublingual. Impulse, of heart, 244 Infant, newly-born, characteristics of, 588 Inflammation, changes of capillary cir- culation in, 259 of eyeball, after division of 5th pair, 397 Infusoria, 431 different kinds of 432 conditions of their production, 432 Schultze's experiment on generation of, 434 Inguinal hernia, congenital, 562 Injection of placental sinuses from ves- sels of uterus, 529 Inorganic substances, as proximate prin- ciples, 37 their source and destination, 46 Inosculation, of veins, 253 of capillaries, 256-257 of nerves, 312 Insalivation, 92 importance of, 99 function of, 100 Inspiration, how accomplished, 201 movements of glottis in, 205 Instinct, nature of, 382, 383 Integument, respiration by, 217 development of, 545 Intellectual powers, 364-383 in animals, 384 Intestine, of fowl, 85 of man, 87 juices of, 118 digestion in, 118, 127 epithelium of, 138 disappearance of bile in, 158 development of, 494, 495, 499, 547 Intestinal digestion, 125 Intestinal juices, 118 action of, on starch, 119 Involution of uterus after delivery, 537 Iris, movements of, 374, 418 after division of sympathetic, 423 Irritability, of gastric mucous membrane, 105 of the heart, 241 of muscles, 328 of nerves, 330 Jackson, Prof. Samuel, on digestion of fat in intestine, 122 Jaundice, 142 yellow color of urine in, 299 Kidneys, peculiarity of circulation in, 263 elimination of medicinal substances by, 298 formation of, 557 KiJCHENMEiSTER, experiments on produc- tion of tcenia from cysticercus, 440 of cysticercus from eggs of tjenia, 441 Lachrymal secretion, 273 its function, 274 Lactation, 274 variations in composition of milk during, 275-278 Lacteals, 134-135 and lymphatics, 136 Larynx, action of, in respiration, 205 in formation of voice, 402 nerves of, 400 INDEX. 601 Larynx, protective action of, 404 movements in respiration, 205,404 Layers, external and internal, of blasto- dermic membrane, 490 Lead, salts of, action in distinguishing the biliary matters, 145-14(3 Lehmann, on formation of carbonates in blood, 45 on total quantity of blood, 196 on effects of non-nitrogenous diet, 76 Ledckakt, on production of cysticercus, 441 Ligament of the ovary, formation of, 563 Limbs, formation of, in frog, 496 in human eraWyo, 544 Liver, vascularity of, 279 lobules of, 280 secreting cells, 281 formation of sugar in, 165 congestion of, after feeding, 172 development of, 560, 576 Liver cells, 280 their action in secretion, 281 Liver-sugar, formation of, 165 after death, 168 in foetus, 551 Lobules, of lung, 200 of liver, 279 Local production of carbonic acid, 212 of animal heat, 226 Local variations of cii'culation, 263 LoNGET, on interference of albuminose with Trommer's test. Ill on sensibility of glosso-pharyngeal nerve, 398 on irritability of anterior spinal roots, 343 LoNGET AND Matteucci, experiment on signs of electricity in an irritated nerve, 338 Lungs, structure of, in reptiles, 199 in man, 200 alteration of, after division of pneu- mogastrics, 407 Lymph, 135 Lymphatics, 133 Magnus, on proportions of oxygen and carbonic acid in blood, 209 Male organs of generation, 458 development of, 558 Malpighian bodies of spleen, 174 Mammalians, circulation in, 231 Mammary gland, structure of, 274 secretion of, 275 Mastication, 89 unilateral, in ruminating animals, 94 retarded by suppressing saliva, 99 Meconium, 549 Medulla oblongata, 324, 378 ganglia of, 325-326 Medulla oblongata, reflex action of, 379- 382 effect of destroying, 381 development of, 540 Meibomian glands, 270 Melanine, 71 Membrane, blastodermic, 490 Membrana granulosa, 469 Membrana tympani, action of, 421 Memory, connection of, with cerebral hemispheres, 362 Menobranchus, size of blood-globules in, 185 gills of, 198 spermatozoa of, 459 Menstruation, 472 commencement and duration of, 473 phenomena of, 473 rupture of Graafian follicles in, 474 suspended during pregnancy, 473, 486 Mesenteric glands, 134 Michel, Dr. Myddleton, rupture of Graaf- ian follicle in menstruation, 474 Milk, 274 composition and properties of, 275 microscopic characters, 276 souring and coagulation of, 277 variations in, during lactation, 278 Milk-sugar, 51, 52 converted into lactic acid, 277 Mollusca, nervous system of, 315 Moore and Pennock, experiments on movements of heart, 240 Motor cranial nerves, 388 Motor nervous fibres, 315 Motor oculi communis, 389 externus, 389 Movements, of stomach, 112 of intestine, 130 of heart, 238 of chest in respiration, 201 of glottis, 204 associated, 348 of foetus, 533 Mucosine, 69 Mucous follicles, 268 Mucous membrane, of stomach, 101 of intestine, 119, 128, 129 of uterus, 516 Mucus, 268 composition and properties of, 269 of mouth, 93 of cervix uteri, 456 Muscles, irritability of, 328 directly paralyzed by sulpho-cya- nide of potassium, 330 consentaneous action of, 348 of respiration, 201-202 Muscular fibres, of spleen, 173 of heart, spiral and circular, 242 Muscular irritability, 328 duration after death, 329 602 INDEX. Muscular irritability, esliausted by re- peated irritation, 330 Masculine, 70 Nails, formation of, in embryo, 545 NtGEiEE, on rupture of Graafian follicle, in menstruation, 474 Nerve-cells, 312 Nerves, division of, 311 inosculation of, 312 irritability of. 32S spinal, 319, 320, 342 cranial, 385 olfactory, 385 optic, 386 auditory, 386 oculo-motorius, 389 patheticus, 389 motor externus, 3S9 masticator, 389 facial, 390 sublingual, 392 spinal accessory, 392 trifacial (oth pair), 394 glosso-pbaryngeal, 398 pneumogastric, 399 superior and inferior laryngeal, 400 great sympathetic, 414 Nervous filaments, 308 of brain, 309 of sciatic nerve, 310 motor and sensitive, 343 Nervous force, tow excited, 330 nature of, 335 mode of transmission, 336-339 Nervous tissue, two kinds of, 307 Nervous irritability, 328 how shown, 331 duration of, after death, 331 exhausted by excitement, 332 destroyed by woorara, 333 distinct from muscular, 335 nature of, 335-339 Nervous system, 305 general structure and functions of, 305-327 of radiata, 313 of moUusca, 315 of articulata, 316 of vertebrata, 318 reflex action of, 314 Network, capillary, in Peyer's glands, 128 in web of frog's foot, 256 in lobule of liver, 280 Newly-born infant, weight of, 558 respiration in, 558 nervous phenomena of, 589 comparative size of organs in, 590 Newport, on temperature of insects, 221 Nitric acid, action of, on bile-pigment, 150 precipitation of uric acid by, 295 Nitrogen, exhalation of. in respiration, 206"' Nutrition, 26-29 Obliteration, of ductus venosus, 578 of ductus arteriosus, 580 Oculo-motorius nerve, 389 CEsophagus, paralysis of, after division of pneumogastric, 401 development of, 552 CEstruation, phenomena of, 471 Oleaginous substances, 54 in difi'erent kinds of food, 56 condition of, in the tissues and fluids, 57 partly pirodTTced in the body, 61 decomposed in the body, 62 in the blood, 137 indispensable as ingredients of the food, 74 insuiEcient for nutrition, 76 Olfactory apparatus, protected by two sets of muscles, 420 commissures, 322, 323, 385 Olfactory ganglia, 322 their function, 358 Olfactory nerves, 385 Olivary bodies, 324 Omphalo-mesenterio vessels, 567-569 Ophthalmic ganglion, 414 Optic ganglia, 322, 373 Optic nerves, 386 decussation of, 375-376 Optic thalami, 323, 359 development of, 540 Organs of special sense, development of, 542 Organic substances, 63 indefinite chemical composition of, 64 hygroscopic properties, 65 coagulation of, 66 catalytic action, 66 putrefaction, 67 source and destination, 72 digestion of, 110 Origin, of plants and animals, 429 of infusoria, 431 of animal and vegetable parasites, 434 of encysted entozoa, 436 Ossification of skeleton, 544 Osteine, 70 Otic ganglion, 415 Ovary, 443 of tfenia, 443 of frog, 449 of fowl, 453 of human female, 455, 456 Ovaries, descent of, in foetus, 562 condition at birth, 564 Oviparous and viviparous animals, dis- tinction between, 465 INDEX. 603 Oxalic acid, produced in urine, 302 Oxygen, absorbed in respiration, 207 state of solution in blood, 209 dissolved by blood-globules, 210 absorbed by tbe tissue-, 212 exhaled by plants, 225 Palate, formation of, 555 Pancreatic juice, 121 mode of obtaining, 122 composition of, 123 action on fat, 123 daily quantity of, 123 Pancreatine, 68 in pancreatic juice, 123 Panizza, experiment on absorption by bloodvessels, 131 Paralysis, after division of anterior root of spinal nerve, 341 direct, after lateral injury of spinal cord, 344 crossed, after lateral injury of brain, 345 facial, 341, 391 of muscles, by sulplio-cyanide of po- tassium, 330, 352 of motor nerves, by woorara, 353 of sensitive nerves, by strychnine, 353 of voluntary motion and sensation, after destroying tuber annulare, 377 of pharynx and oesophagus, after section of pneumogastrics, 401 of larynx, 402-412 of muscular coat of stomach, 411 Paraplegia, reflex action of spinal cord, in, 351 Parasites, 434 conditions of development of, 435 mode of introduction into body, 436 sexless, reproduction of, 436-441 Parotid saliva, 93 Parturition, 534-535 Par vagum, 399. See Pneumogastric Patheticus nerve, 389 Pelvis, development of, 554 Pennock and Moore, experiments on movements of heart, 240 Pepsine, 69 in gastric jxiice, 108 Perception of sensations, afterremoval of hemispheres, 362 destroyed, after removal of tuber annulare, 377 Periodical ovulation, 465 Peristaltic motion, of stomach, 112-113 of intestine, 130 of oviduct, 450-452 of Fallopian tubes, 475 Perspiration, 271 daily quantity of, 272 Perspiration, composition and properties of, 272 function, in regulating temperature, 273 Pettenkofer's test for bile, 151 Peyer's glands, 128 Pharynx, action of, in swallowing, 40l formation of, 551 Phosphate of lime, its proportion in the animal tissues and fluids, 42 in the urine, 295 precipitated by alkalies, 296 Phosphate, triple, in putrefying urine, 304 Phosphates, alkaline, 45 in urine, 295 earthy, 42-45 in urine, 295 of magnesia, soda, and potass, 45 Phosphorus, not a proximate principle, 31 Physiology, definition of, 17 Phrenology, 367 objections to, 368 practical difficulties of, 368-369 Pigeon, after removal of cerebrum, 362 of cerebellum, 372 Placenta, 523 comparative anatomy of, 524 formation of, in human species, 525 . foetal tufts of, 527 maternal sinuses of, 527 injection of, from uterine vessels, 529 function of, 530 separation of, in delivery, 535 Placental circulation, 568, 570 Plants, vital heat of, 221 generative apparatus of, 442 Plasma of the blood, 187 Pneumic acid, 21] Pneumogastric nerve, 399 its distribution, 400 action of, on pharynx and oesopha- gus, 401 on larynx, 402 in formation of voice, 403 in respiration, 404 effect of its division, on respiratory movements, 406-407 cause of death after division of, 409-410 influence of, on oesophagus and sto- mach, 411-412 Pneumogastric ganglion, 400 PoGGiALE, on glycogenic matter in but- cher's meat, 170 Pons Varolii, 324, 327 Portal blood, quantity of fibrin in, 189 temperature of, 227 Portal vein, in liver, 279 development of, 576 Posterior columns of spinal cord, 321 604 INDEX. Primitive trace, 492 Production, of sugar in liver, 165 of carbonic acid, 214 of animal heat, 218 of urea in blood, 285 of infusorial animalcules, 431 of animal and vegetable parasites, 434 Proximate principles, 29 definition of, 31 mode of extraction, 32 manner of their association, 33 varying proportions of, 34 three distinct classes of, 35 Proximate principles of the first class (inorganic), 37 of the second class (crystallizable substances of organic origin), 47 of the third class (organic sub- stances), QS Ptyaline, 93 Puberty, period of, 467 signs of, in female, 472 Pulsation, of heart, 235 in living animal, 239 of arteries, 247, 248 Pupil, action of, 374, 418 contraction of, after division of sym- pathetic, 423 Pupillary memi)rane, 542 Putrefaction, 67 of the urine, 300 Pyramids, anterior, of medulla oblon- gata, 324 Quantity, daily, of water exhaled, 39 of food, 81 of saliva, 96 of gastric juice, 114 of pancreatic juice, 123 of bile, 153 of carbonic acid exhaled, 214 of perspiration, 272 of urine, 291 of urea, 286 of urate of soda, 289 Quantity, entire, of blood in body, 196 Rabbit, brain of, 323 Races of men, different capacity of, for civilization, 364 Radiata, nervous system of, 313 Rapidity of circulation, 261 of transmission of nervous force, 336-337 Reactions, of starch, 50 of sugar, 52 of fat, 54 of saliva, 93 of gastric juice, 107-108 of intestinal juice, 121 of pancreatic juice, 123 of bile, 150 Reactions, of mucus, 269 of milk, 276 of urine, 295 Reasoning powers, 363-364 in animals, 383 Red globules of blood, 179 Reflex action, 314 in centipede, 317 of spinal cord, 348, 382 of medulla oblongata, 379, 382 of tuber annulare, 378, 382 of brain, 383 of optic tubercles, 384 in newly born infant, 589 Regeneration, of uterine mucaus mem- brane after pregnancy, 536 of walls of uterus, 537-538 Regnault and Reiset, on absorption of oxygen, 225 Reid, Dr. John, experiment on crossing of streams in foetal heart, 583 Reproduction, 427 nature and object of, 427-429 of parasites, 434 of tasnia, 438 by germs, 442 Reptiles, circulation of, 231 Respiration, 197 by gills, 198 by lungs, 199 by skin, 217 changes in air during, 206 changes in blood, 207 of newly born infant, 588 Respiratory movements of chest, 201 of glottis, 204 after section of pneumogastrics,404- 406 after injury of spinal cord, 380 Restiform bodies, 325 Rhythm of heart's movements, 244 Rotation of heart during contraction, 243 Round ligament of the uterus, formation of, 563 of liver, 578 Rumination, movements of, 94 Rupture of Graafian follicle, 470 in menstruation, 474 Rutting condition, in lower animals, 471 Saccharine substances, 51, 52, 54 in stomach and intestine, 118 in liver, 165 in blood, 171 in urine, 299 Saliva, 92 different kinds of, 93 daily quantity of, 96 action on boiled starch, 96 variable, 97 does not take place in stomach, 98 physical function of saliva, 98 INDEX. 605 Saliva, quantity absorbed by difTerent kinds of food, 100 Salivary glands, 93 Salts, biliary, 143 of the blood, 190 of urine, 288 Saponification, of fats, 55 ScuAKLiNG, on diurnal variations in exha- lation of carbonic acid, 216 ScHULTZE, experiment on generation of infusoria, 434 Scolopendra, nervous system of, 316 Sebaceous matter, 269 composition and properties of, 270 function of, 271 in foetus, 545 Secretion, 265 varying activity of, 267 of saliva, 93 of gastric juice, 105 of intestinal juice, 120 of pancreatic juice, 122 of bile, 153, 278 of sugar in liver, 165 of mucus, 268 of sebaceous matter, 269 of perspiration, 271 of the tears, 273 of bile in foetus, 551 Segmentation of the vitellus, 489 Seminal fluid, 458 mixed constitution of, 462 Sensation, remains after destruction of hemispheres, 362 lost after removal of tuber annulare, 377 special, conveyed by pneumogastric nerve, 379, 406 Sensation and motion, distinct seat of, in nervous system, 340 in spinal cord, 343 Sensibility, of nerves to electric current, 337 and excitability, definition of, 341 seat of, in spinal cord, 343 in brain, 357 of facial nerve, 391 of sublingual nerve, 392 of spinal accessory, 393 of great sympathetic, 417 Sensibility, special, of olfactory nerves, 385 of optic nerves, 386 of auditory nerves, 386 of lingual branch of 5th pair, 398 of glosso-pharyngeal, 399 of pneumogastric, 408 Sensitive nervous filaments, 315 Sensitive fibres, crossing of, in spinal cord, 345 of facial nerve, source of, 390 Sensitive cranial nerves, 393 Septa, inter-auricular and inter- ventricu- lar, formation of, 581 S£quakd, on crossing of sensitive fibres in spinal cord, 345 Serum, of the blood, 192 Sexes, distinctive characters of, 444 Sexless entozoa, 436 Sexual generation, 442 Shock, effect of, in destroying nervous irritability, 330 SiEBOLD, on production of tienia from cysticercus, 440 Sinus terminalis, of area vasculosa, 505 Sinuses, placental, 527 Skeleton, development of, 543 Skin, respiration by, 217 sebaceous glands of, 269 perspiratory glands of, 271 development of, 545 Smell, ganglia of, 358 nerves of, 385 injured by division of 5tli pair, 396 Smith, Dr. Southwood, on cutaneous and pulmonary exhalation, 272 Solar plexus of sympathetic nerve, 416 Sounds, of heart, 235 how produced, 236 vocal, how produced, 420 destroyed by section of inferior la- ryngeal nerves, 403 of spinal accessory, 404 Sounds, acute and grave, transmitted by memljrana tympani, 421 Species, mode of continuation, 429 Spermatic fluid, 458 mixed constitution of, 462 Spermatozoa, 458-459 movements of, 460 formation of, 461 Spina bifida, 543 Spinal accessory, 392 sensibility of, 393 communication of, with pneumogas- tric, 400 Influence of, on larynx, 404 Spinal column, formation of, 493, 543 Spinal cord, 319-340 commissures of, 320 anterior and posterior columns, 321 origin of nerves from, 319, 320 sensibility and excitability of, 343 crossed action of, 344 reflex action of, 348 protective action of, 353 influence on sphincters, 354 effect of injury to, 355 on respiration, 380 formation of, in embryo, 493, 543 Spinal nerves, origin of, 319, 321 Spleen, 173 Malpighian bodies of, 174 extirpation of, 176 Spontaneous generation, 429 606 INDEX. Starch, 47 proportion of, iu different kinds of food, 48 varieties of, 48 reactions of, 50 action of saliva on, 96 digestion of, 118 Starfish, nervous system of, 313 St. Martin, case of gastric fistula in, 103 Strabismus, after division of motor oculi communis, 389 of motor externus, 389 Striated bodies, 359 Sublingual gland, secretion of, 93-94 nerve, 392 Submaxillary ganglion, 414 gland, secretion of, 93-94 Sudoriparous glands, 271 Sugar, 51 varieties of, 51 composition of, 52 tests for, 52 fermentation of, 52 proportion in diflferent kinds of food, 54 source and destination, 54 discharged by urine in disease, 299 Sugar in liver, formation of, 165 percentage of, 167 produced in hepatic tissue, 168 from glycogenic matter, 169 absorbed by hepatic blood, 171 decomposed in circulation, 171 Sulphates, alkaline, in urine, 296 Sulphur of the bile, 147 not discharged with the feces, 161- 162 Swallowing, 100 retarded by suppression of saliva, 99 by division of pneumogastic, 411 Sympathetic nerve, 414 its distribution, 415 sensibility and excitability of, 417 influence of, on special senses, 418 onpupil, 418, 419, 423 on nutrition of eyeball, 397-398 on nasal passages, 420 on ear, 421 on temperature of particular parts, 421-422 reflex actions of, 424 Tadpole, development of, 494-495 transformation into frog, 496 Taenia, 438 produced by metamorphosis of cys- ticercus, 440 single articulation of, 443 Tapeworm, 438 mode of generation, 439 Taste, nerves of, 398-399 of alimentary substances, developed by cooking, 82 Taurine, 147 Tauro-cholate of soda, 146 microscopic characters of, 144-145 Tauro-cholic acid, 147 Tears, 273 their function, 274 Teeth, of serpent, 89 of polar bear, 90 of horse, 90 of man, 91 first and second sets of, 590-591 Temperature, of the blood, 219 of diff'erent species of animals, 220 of the blood in diflferent organs, 227 elevation of, after section of sympa- thetic nerve, 422 Tensor tympani, action of, 421 Tests, for starch, 50 for sugar, 52 for bile, 150 Pettenkofer's, 151 Testicles, 461 periodical activity of, in flsh, 433 development of, 558 descent of, 559 Tetanus, pathology of, 350 Thalami, optic, in rabbit, 323 in man, 326 function of, 359. Thoracic duct, 134 Thoracic respiration, 380 Tongue, motor nerve of, 392 sensitive, 398-399 Trichina spiralis, 436 Tricuspid valve, 234. See Auriculo- ventricular. Triple phosphate, in putrefying urine, 304 Trommer's test for sugar, 52 interfered with by gastric juice, 111 Tuber annulare, 325 efi'ect of destroying, 377 action of, 378 Tubercula quadrigemina, 322, 323, 373 reflex action of, 374 crossed action of, 375 development of, 540 Tubules of uterine mucous membrane, 517 Tufts, placental, 527 Tunica vaginalis testis, formation of, 561 Tympanum, function of, in hearing, 421 Umbilical cord, formation of, 533 withering and separation of, 590 Umbilical hernia, 548 Umbilical vesicle, 498 in human embryo, 499 in chick, 506 disappearance of, 533 Umbilical vein, formation of, 570-577 obliteration of, 578 Umbilicus, abdominal, 494 INDEX. 607 Umbilicus, amniotic, 502 decidual, 519 Unilateral mastication, in ruminating animals, 94 Urate of soda, 288 its properties, source, daily quantity, kc, 289 Urates of potass and ammonia, 289 Urachus, 549 Urea, 284 source of, 285 mode of obtaining, 285 conversion into carbonate of am- monia, 285 daily quantity of, 286 diurnal variations in, 287 decomposed in putrefaction of urine, 302 Uric acid, 288, 296 Urine, 290 general character and properties of, 291 quantity and specific gravity, 292 diurnal variations of, 293 composition of, 294 reactions, 295-296 interference with Trommer's test, 297 accidental ingredients of, 297 acid fermentation of, 300-301 alkaline fermentation of, 302 final decomposition of, 304 Urinary bladder, paralysis and inflam- mation of, after injury to spinal cord, 355 formation of, in embryo, 548 Urosacine, 71 Uterus, of lower animals, 455 of human female, 456 mucous membrane of, 517 changes in, after impregnation, 518 involution of, after delivery, 537 development of, in foetus, 562 position of, at birth, 564 Uterine mucous membrane, 516 tubules of, 517 conversion into decidua, 519 exfoliation of, at the time of delivery, 535 its renovation, 536 Valve, Eustachian, 581-582 of foramen ovale, 585 Valves, cardiac, action of, 234 cause of heart's sounds, 236 Vasa deferentia, formation of. 559-560 Vapor, watery, exhalation of, 39 from lungs, 207 from the skin, 271 Variation, in quantity of bile in differ- ent animals, 154-157 in production of liver-sugar, 171-172 in size of spleen, 173 Variation, in rapidity of coagulation of blood, 192 in size of glottis in respiration, 205 in exhalation of carbonic acid, 214- 216 in temperature of blood in different parts, 227 in composition of milk during lac- tation, 278 in quantity of urea, 286-287 in density and acidity of urine, 291 -293 Varieties of starch, 48 of sugar, 51 of fat, 54 of biliary salts in diiferent animals, 158 Vegetable food, necessary to man, 74 Vegetables, production of heat in, 221 absorption of carbonic acid and ex- halation of oxygen by, 17, 225 Vegetable parasites, 434-435 Vegetative functions, 27 Veins, 252 al3Sorption by, 131 action of valves in, 253 motion of blood through, 252-254 rapidity of circulation in, 254, 255 omphalo-mesenteric, 567 umbilical, 570 vertebral, 573 Vense cavse, formation of, 574-575 position of, in foetus, 581 Vena azygos, superior and inferior, formation of, 575 Venous system, development of, 573 Ventricles of heart, single in fish and reptiles, 230-231 double in birds and mammalians, 232 situation of, 233 contraction and relaxation of, 239 elongation during contraction, 240 muscular fibres of, 242 Vernix caseosa, 545 Vertebrata, nervous system of, 318 Vertebrje, formation of, 493, 543 Vesicles, adipose, 58 pulmonary, 200 seminal, 462, 561 Vesiculse seminales, 462 formation of, 561 Vicarious secretion, non-existence of, 266 Vicarious menstruation, nature of, 266 Villi, of intestine, 129 absorption by, 130 of chorion, 512 Vision, ganglia of, 322, 323, 326, 373 nerves of, 386 Vital phenomena, their nature and pecu- liarities, 22 Vitellus, 447 608 INDEX. Vitellus, segmentation of, 489 formation of, in ovary of foetus, 564, 565 Vitelline circulation, 566, 567 Vitelline membrane, 446 Vitelline spheres, 489 Vocal sounds, how produced, 402 Voice, formation of, in larynx, 402 lost, after division of spinal acces- sory nerve, 403 Volition, seat of, in tuber annulare, 377 Vomiting, peculiar, after division of pneumogastrics, 411 Water, as a proximate principle, 37 its proportion in the animal tissues and fluids, 38 its source, 38 mode of discharge from the body, 39 Weight of organs, comparative, in newly born infant and in adult, 590 White globules of the blood, 185 White globules of the blood, action of acetic acid on, 186 sluggish movement of, in circulation, 257 White substance, of nervous system, 308 of Schwann, 308 of spinal cord, 320-321 of brain, insensible and inexcitable, 360 Withering and separation of umbilical cord, after birth, 590 Wolffian bodies, 556 structure of, 557 atrophy and disappearance of, 560 vestiges of, in adult female, 563 WvMAN, Prof. Jeffries, on cranial nerves of Rana pipiens, 388 Yellow color, of urine in jaundice, 299 of corpus luteum, 481 Zona pellucida, 446 THE END, CATALOGUE OP BIANCHARD AND LEA'S MEDICAL AND SCIENTIFIC PUBLICATIONS. TO THE MEDICAL PROFESSION. The prices on the present catalogue are those at which our books can gene- rally be furnished by booksellers throughout the United States, who can readily procure any which they may not have on hand. To physicians who have not convenient access to bookstores, we will forward them by mail, at these prices, free of postage (as long as the existing facilities are afforded by the post office), for any distance under 3,000 miles. As we open accounts only with booksell- ers, the amount must in every case, without exception, accompany the order, and we assume no risks of the mail, either on the money or on the books; and as we deal only in our own publications, we can supply no others. Gentlemen desirous of purchasing will, therefore, find it more advantageous to deal with the nearest booksellers whenever practicable BLANCHARD & LEA. Philadelphia, July, 1859. THE AMERICAIf MEDICAL JOURJfAL. TWO MEDICAL PERIODICALS, FREE OF POSTAGE, FOR FIVE DOLLARS PER ANNUM. 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"With Illustrations on Wood, (of which a more detailed account will be found on p. 42.) Commenced in the News for January, 1858, the work will be completed during 1859. Twenty pages (so arranged as to be readily detached for binding on completion) will generally be devoted to it in every number, while the remainder of the periodical will contain, as hereto- fore, a choice selection of important clinical reports, together with topics of immediate interest, domestic and foreign. It will thus be seen that for the small sum of FIVE DOLLARS, paid in advance, the sub- scriber will obtain a Quarterly and a Monthly periodical, EMBRACING ABOUT FIFTEEN HUNDRED LARGE OCTAVO PAGES. mailed to any part of the United States, free of postage. Those subscribers who do not pay in advance will bear in mind that their subscription of Five Dollars will entitle them to the Journal only, without the News, and that they will be at the expense of their own postage on the receipt of each number. The advantage of a remittance when ordering the Journal will be thus apparent. As the Medical News and Library is in no case sent without advance payment, its subscribers will always receive it free of postage. It should also be borne in mind that the publishers will n6w take the risk of remittances by mail, only requiring, in cases of loss, a certificate from the subscriber's Postmaster, that the money was duly mailed and forwarded. ^f Funds at par at the subscriber's place of residence received in payment of subscriptions. Address BLANCHARD & LEA, Philadelphia. MEDICAL AND SCIENTIFIC PUBLICATIONS. NEW AND ENLARGED EDITION— Now Ready (1858). MEDICAL LEXICON; A DICTIONARY OF MEDICAL SCIENCE, CONTAINING A Concise Explanation of the Various Subjects and Terms of ANATOMY, PHYSIOLOGY, PATHOLOGY, HYGIENE, THERAPEUTICS, PHARMA- COLOGY, PHARMACY, SURGERY, OBSTETRICS, MEDICAL JURIS- PRUDENCE, DENTISTRY, &c., NOTICES OF CLIMATE AND OF MINERAL WATERS; FORMULAE FOR OFFICINAL. EMPIRIGM. AND DIETETIC PREPARATIONS, ETC, WITH FRENCH AND OTHER SYNONYMES. By ROBLEf DUNGLISON, M. D., Professor of Institutes of Medicine in tlie Jefferson Medical College, Philadelphia. Fifteenth Edition, Revised and very greatly Enlarged, III one very large and handsome octavo volume, of 992 douhle-columned pages, in small type; strongly hoiond in leather, with raised bands. Price $4. Especial care has been devoted in the preparation of this edition to render it in every respect worthy a continuance of the very remarkable favor which it has hitherto enjoyed. The rapid sale of Fifteen large editions, and the constantly increasing demand, show that it is regarded by the profession as the standard authority. Stimulated by this fact, the author has endeavored in the present revision to introduce whatever might be necessary "to make it a satisfactory and desirable — if not indispensable — lexicon, in which the student may search without disap- pointment for every term that has been legitimated in the nomenclature of the science " To accomplish this, large additions have been found requisite, and the extent of the author's labors may be estimated from the fact that about Six Thousand subjects and terms have been intro- duced throughout, rendering the whole number of definitions about Sixty Thousand, to ac- commodate which, the number of pages has been increased by nearly a hundred, notwithstand- ing an enlargement in the size of the page. The medical press, both in this countrj^ and in England, has pronounced the work indispensable to all medical students and practitioners, and the present improved edition will not lose that enviable reputation. The publishers have endeavored to render the mechanical execution worthy of a volume of such universal use in daily reference. The greatest care has been exercised to obtain the typo- graphical accuracy so necessary in a work of the kind. By the small but exceedingly clear type employed, an immense amount of matter is condensed in its thousand ample pages, while the binding will be found strong and durable. With all thete improvement* and enlargements, the price has been kept at the former very moderate rate, placing it within the reacii of all. This work, the appearance of the fifteenth edition of which it has become our duty and pleasure to an- nounce, is perhaps the most stupendous monument of labor and erudition in medical literature. One would hardlj' suppose after constant use of the pre- ceding; editions, where we have never failed to find a sufiSciently full explanation of every medical term, that in this edition "about six thousand subjects and terms have been added" with a careful revision and correction of the entire work. It is only necessary to announce the advent of this edition to make it occupy the place of the preceding one on the table of every medical man. as it is without doubt the best and most comprehensive work of the kind which has ever ap- peared. — Buffalo Med. Journ., Jan. 1S58. The work is a monument of patient research, skilful judgment, and vast physical labor, that will perpetu- ate the name of the author more effectually than any possible device of stone or metal. Dr. Dunglison de- serves the thanks not only of the American profession, but of the whole medical world. — JVorth Am. Mtdico- Chir. Review, Jan. 1858. A Medical Dictionary better adapted for the wants of the profession than any other with which we are acquainted, and of a character whicli places it far above comparison and competition. — American Jour- nal of the Medical Sciences, Jan. 1858. AVe need only say, that the addition of 6,000 new terms, with their accompanying definitions, may be said to constitute a new woric by itself. We have ex- amined the Dictionary attentively, and are most happy to pronounce it unrivalled of its kind. The erudition displayed, and the extraordinary industry which must have been demanded, in its preparation and perfec- tion, redound to the lasting credit of its author, and have furnished us with a volume indispensable at the present day, to all who would find themselves au ni- veau with the highest standards of medical informa- tion.— jBostore Med. and Surg. Journal, Dec. 31, 1867. Good lexicons and encyclopedic works generally, are the most labor-saving contrivances which literary men enjoy: and the labor which is required to produce them in the perfect manner of this example is some- thing appalling to contemplate. The author tells us in his preface that he has added about six thousand terms and subjects to this edition, which, before, was considered universally as the best work of the kind in any language. — Silliman's Journal, March, 1858. A very perfect work of the kind, undoubtedly the most perfect in the English language.— i/ed. awd Surg. Reporter, Jan. 1858. The most complete authority on the subject to be found in any language. — Va. Med. Journal, Feb. 185 8. BLANCHARU AND LEA'S New and Improved Edition— Just Issued. XN ANALYTICAL COMPEIDIDM OF THE VARIOUS BRANCHES OF MEDICAL SCIENCE. £ov tlje ilse arib (£?famincition of Stubcnts. BY JOHN NEILL, M. R, StlKGEON TO THE PENNSYLVANIA HOSPITAL, ETC., AND FRANCIS GURNEY SMITH, M. D., PROFESSOR OF INSTITUTES OF MEDICINE IN THE PENNSYLVANIA MEDICAL COLLEGE, ETC. A NEW EDITION, REVISED AND IMPROVED, With Three Hnndred and Seventy-four Illnstrations. /« one, very large and handsome royal l2mo. volume of vmrly 1000 pages, strongly honnd in leather, with raised bands. $3. This work presents a complete and systematic- outline of the whole range of medical science, di- vided under the headings of Anatomy, Physiolo GY, Surgery, Midwife- ry, Chemistry, Mate- ria Medica and The- rapeutics, and Prac- tice or Medicine. — Each portion is thorough- ly and appropriately il- lustrated, rendering tlie volume one of the cheap- est as yet placed befure the profession. Lateral Operation of Lithotomy The very flattering reception which has been accorded to this work, and the high estimate placed upon it by the profession, as evinced by the constant and increasing demand which has rapidly exhausted two large editions, have stimulated the authors to render the volume in its present revision more worthy of the success which has attended it. It has accordingly been thoroughly examined, and such errors as had on former occasions escaped observation have been corrected, and whatever additions were necessary to maintain it on a level with the advance of science have been introduced. The extended series of illustrations has been still further in- creased and much improved, while, by a slight enlargement of the page, these various additions have been incorporated without increasing the hulk of the volume. The work is therefore again presented as eminently worthy of the favor with which it has hitherto been received. As a book for daily reference by the student requiring a guide to his more elaborate text-books, as a manual for preceptors desiring to stimulate their students by frequent and accurate examination, or as a source from which the practitioners of older date may easily and cheaply acquire a knowledge of the changes and improvement in professional science, its reputation is permanently established. The Compend of Drs. Neill and Smith is inoompara bly the most valuable Tvork of its class ever published in this country. Attempts have been made in various- quarters to squeeze Anatomy, Physiolosiy, Surgery, the Practice of Medicine, Obstetrics, Materia Medica, and Chemistry into a single manual ; but the operation has signally failed in the hands of all up to the advent of " Neill and Smith's" volume, which is quite a mira- cle of success. The outlines of the whole are admirably drawn and illustrated, and the authors are eminently entitled to the grateful consideration of the student of every class. — N. 0. Med. and Surg. Joum., May, 1856. This popular favorite with the student is so well known that it requires no more at the hands of a medical editor than the annunciation of a new and improved edition. There is no sort of comparison be- tween this work and any other on a similar plan, and for a similar object. — Nashville Journal, of Medicine, Sept. 1856. There are but few students or practitioners of medi- cine unacquainted with the former editions of this unassuming though highly instructive work. The whole science of medicine appears to have been sifted, as the gold-bearing sands of El Dorado, and the pre- cious facts treasured up in this little volume. A com- plete portable library so condensed that the student may make it his constant pocket companion. — Western Lancet, May, 1856. To compress the whole science of medicine in less than 1,000 pases is an impossibility, but we think that the book before us Approaches as near to it as is possi- ble. Altogether, it is the best of its class, and has met with a deserved success. As an elementary text-book for students, it has been useful, and will continue to be employed in the examination of private classes, whilst it will often be referred to by the country prac- titioner. — Va. Med. Journal, May, 1856. MEDICAL AND SCIENTIFIC PUBLICATIONS. NEW AND MUCH ENLARGED EDITION— (Just Issued.) A MANUAL OF EXAMmATIONS UPON ANATOMY, PHYSIOLOGY, SURGERY, PRACTICE OF MEDICINE, CHEMISTRY, OBSTETRICS, MATERIA MEDICA, PHAR- MACY, AND THERAPEUTICS: TO WHICH IS ADDED A MEDICAL FORMULARY. BY J. L. LUDLOW, M. D. A new Edition, thoroughly modified, and greatly extended and enlarged. With three hundred and seventy wood engravings, 1)1 one large and handsome royal VZtno. vol. of over S^Q closely printed pages, strongly bound in leather., price S2 50. Crura Cerebri, ij-c. Ampulalion of Hand. The great popularity which this volume has always enjoyed, has stimulated the author in his revision to render "it in every respect worthy of the confidence of the profession, giving rise to the delay which has caused it to remain out of print for so long a time Every portion has been sedulously examined, and the most recent observations and investigations introduced, rendering it an accurate resume of the most improve! condition of medi^^al science. A con- siderable portion has been rewritten, and entire sections on Physiology and Organic Chemistry have been added, while a very complete series of illustrations has been introduced, elucidating the text wherever such assistance appeared necessary or desirable. Notwithstanding an en- largement of the page, these improvements have increased the size of the volume to over eight hundred pages, and with the greatly improved style of mechanical execution, it may in almost every respect Ise regarded rather as a new work than a new edition. The arrangement of the volume in the form of question and answer renders it especially suit- able for the office examination of students, and for those preparing for graduation. We would recommend this book as one of the best of its kind, and to every right-minded student a most valuable aid in acquiring the Facts of Medical Science. — South. Med. and Surg. Journ., Nov. 1S57. In the main, we regard his work as well adapted for its professed object, and that the author's claims are valid. Abundant illustrations of the text are fur- nished by means of wood cuts, well executed; a de- cided advantage and help to the student. — Penin Jour. of Med; Aug. 1857. For the purpose for which it is intended, we do not see but Dr Ludlow's work is as good as any other of the kind. The peculiarity which distinguishes it from some others of the same class, consists in the form of question and answer in which it is written. We com- mend it to the notice of the student who feels that he must rely on such a work before being confidentially closeted with his friends, the professors, previous to a final departure. — A^. J. Med. Reporter, Aug. 1857. The illustrations are good ; and the system of ques- tions and answers are as well arranged as the stu- dent seeking this kind of help will probably desire. — Charleston Med. Jour., July, 1857. BLANCHARD AND LEA'S New and Enlarged Edition— Just Issued. A DICTIONARY OF TERMS USED IN MEDICINE, AND THE COLLATERAL SCIENCES. BY RICHARD D. HOBLYN. A NEW AMERICAN FROM THE LAST LONDON EDITION. Revised, with numerous Additions, BY ISAAC HAYS, M. D., Editor of the American Journal of the Medical Sciences. In one large royal Vlmo. volume £>/522 closely printed douhle-cohcmn pages, leather. %\ 50. In this volume the object of the author and editor has been to produce a work which, at an exceedingly moderate price, and in a portable and convenient form, should prcf^ent all the assist- ance requisite to the medical student and ordinary scientific reader. All obsolete terms have been carefully excluded, and it will be found a complete and concise manual of definitions, em- bodying the terms employed in medicine and its allied sciences in their present advanced condi- tion. By the employment of a small but clear type, the amount of an ordinary octavo volume has been condensed into its pages. If the 'frequency with which we have referred to this -volume since its reception from the publisher, two or three weeks ago, tie any criterion for the future, the binding will soon have to be renewed, even with careful handling. We find that Dr. Hays has done the profession great service by his careful and indus- trious labors. The Dictionary has thus become emi- nently suited to our medical brethren in this country. The additions by Dr. Hays are in brackets, and we believe there is not a single page but bears these in- signia; in every instance which we have thus far noticed, the additions are really needed and exceed- ingly valuable. We heartily commend the work to alfwho wish to be au courant in medical terminology. — Boston Med. and Surg. Journal. To supply the want of the medical reader arising from this cause, we know of no dictionary better ar- ranged and adapted than the one bearing the above title. It is not encumbered with the ob.solete terms of a by-gone age, but it contains all that are now in use; emliracing every department of medical science down to the very latest date. The volume is of a con- venient size to be used by the medical student, and yet large enough to make a respectable appearance in the library of a physician. — yVesta-n Lancet. Hoblyn's Dictionary has long-been a favorite with us. It is the best book of definitions we have, and ought always to be upon the studeat'a table. — South- ern Med. mid Surg. Journal. WILSON'S DISSECTOR— New Edition, just issued. THE DISSECTOR'S MANUAL OF PRACTICAL AND SURGICAL ANATOMY. BY ERASMUS WILSOX, F. R. S., Author of "A System of Human Anatomy." Third American from the last and Revised Lon- don Edition. ILLUSTR.^TED WITH 144 Engravings on Wood. EDITED BY WILLIAM HUNT. M.D., Demonstrator of Anatomy in the University of Penn'a In one large and handsome royal 12tno. volume of 58-4 pages. Bound in leather, $2, The modifications and additions which this work has received in again passing through the hands of the author are sufficiently indicated by the fact that the present edition contains nearly one- half more matter than the preceding, while the series of illustrations ha* been increased in extent and greatly improved in character. By the em- ployment of a smaller type, these addi- tions have been accommodated with- out a correspondnig enlargement in the size and price of the volume, and it is again presented as fully worthy a continuance of the favor which it has heretofore enjoyed as a sound practical guide to the study of anatomy. MEDICAL AND SCIENTIFIC PUBLICATIONS. A NE-W AMERICAN DISSECTOR— (Just Issued). THE PEACTIOAL ANATOMIST; OR, THE STUDENT'S GUIDE IN THE DISSECTING-ROOM. BY J. M. ALLEN, M. D., Late Professor of Anatomy in the Pennsylvania Medical College. SMitlj 26*6^ Illustrations. In 07ie large and very hatidsomi', royal 12mo. volume of 632 pages, leather, $2 25. The very large number of elaborate illusira- lions with which this woric abounds, serves to render the verbal details easy of comprehen- sion, showing the student what to examine, and where and how to look for it, while the long experience of the author as a teacher of anatomy, has given him a familiarity with the wants of students, and has shown him the best modes of obviating or relieving the difli- cullies which present themselves in the pro- gress of dissection. As adapted to the course pursued in our colleges, and as containing am- ple practical directions and instructions, in ad- dition to the anatomical details presented, it, therefore, possesses claims to the immediate attention of teachers and students. It is a very convenient manual, and by its ar- rangement, is well adapted to the study of practical anatomy. — N'. Y. Journ. of Medicine. We think it admirably arranged for rendering aid to the student in prosecuting dissections. Its illus- trations are generally accurately drawn and highly useful. It will, doubtless, rank among the best guides in the study of practical anatomy that are now before the profession. — N. W. Med. and Surg. Journal, Jan. 1857. We are greatly pleased with Prof Allen's little work. We handed our copy to a friend, who is more particularly engaged in Practical Anatomy, for his careful examination and opinion ; he reports it " the very best dissector yet produced." The arrangement is clear, concise, and convenient; its size is satisfac- tory — full enough for all the purposes of the dissect- ins-rorm, and yet, not prolix or bulky. It is beautifully illustrated— perhaps much the most so of any dissector yet published. — Cincinnati Med. Observer. Arteries in the Groin. SPECIAL ANATOMY AND HISTOLOGY BY WILLIAM E. HORNER, M. D , Late Professor of Anatomy in the University of Pennsylvania 3£i5tlt anij EmproiitlJ B&ition. I?i two large and handsome octavo volumes of over one thousand pages, With more than 300 beautiful Illustrations. Extra Cloth, $6. This work has so long occupied the position ot a standard text-book and work of reference among anatomists, that the present edition, fully revised and thoroughly brought up by the author shortly before his death, cannot fail to maintain its distin- guished reputation. Os Femoris macerated in acid. BLANCHAUD AND LEA'S THE STUDENT'S TEXT-BOOK OF ANATOMY. New and much enlarged edition — Just Ready (1858). In one large and exquisitely printed octavo Yolume, ■with three hundred and ninety-seven beautiful engravings on wood, and more than 600 large pages. Price, in leather, $3 25. Saphenous opening m the Fascia Lata. A SYSTEM OF HUMAN ANATOMY. GENERAL AND SPECIAL. BY ERASMUS WILSON, F. R. S, Author of " The Dissector's Manual," " A Treatise on Diseases of the Skin," &c. A New and Revised American, from the last and enlarged English Edition. Edited by W. H. GOBRECHT, M. P., Professor of Anatomy in the Philadelphia College of Medicine, &c. The publishers trust that the well-earned reputation so long enjoyed by this work will be more than maintained by the pre- sent edition. Besides a very thorough revision by the author, it has been most carefully examined by the editor, and the efforts of both have been directed to introducing everything which increased experience in its use has suggested as desira- ble to render it a complete text- book for those seeking to obtain or to renew an acquamtance with Human Anatomy. The amount of additions which it has thus received may be esti- mated from the fact that the present edition contains over one- fourth more matter than the last, rendering a smaller type and an enlarged page requisite to keep the volume within a con- venient size. The author has not only thus added largely to the work, but he has also made alterations throughout, where- ver there appeared the opportunity of improving the arrange- ment or style, so as to present every fact in its most appropri- ate manner, and to render the whole as clear and intelligible as possible. The editor has exercised the utmost caution to obtain eniire accuracy in the text, and has largely increased the number of illustrations, of which there are about one hundred and fifty more in this edition than in the last, thus bringing distinctly before the eye of the student everything of interest or importance. The publishers have felt that neither care nor expense should be spared to render the external finish of the volume worthy of the universal favor with which it has been received by the Ame- rican profession, and they have endeavored, consequently, to produce in its mechanical execu- tion, an improvement corresponding with that which the text has enjoyed. It will therefore be found one of the handsomest specimens of typography as yet produced in this country, and in all respects suited to the office table of the practitioner, notwithstanding the exceedingly low price at which it has been placed. Portion of one of Peyer''s Glands. It is therefore at the expense of some struggle with our predilections that we find ourselves called upon to recognize the merits of a successor to our earlier companion and guide. The struggle over, we are con- strainea to declare that the edition of 1858 is a vast improvement upon all others. This is, in one sen- tence, the best edition of the best teaching anatomy now extant. — Nashville Mrmthly Record, Nov. '58. The great practical value of Wilson's Anatomy, as a manual for the student, the practitioner, and for all who desire to become acquainted with the subject, ia too well attested by the unprecedented success of the work, and the universal verdict in its favor, to render recommendation necessary. We have ever commend- ed Wilson's Anatomy, without hesitation or reserve, to students of medicine, and the present edition only increases our approbation. — Southern Med. and Surg. Journal, Nov. 1858. MEDICAL AND SCIENTIFIC PUBLICATIONS NEW AND MAGNIFICENT ANATOMICAL TEXT-BOOK— (Now Ready.) ANATOMY, DESCRIPTIVE AND SURGICAL. BY HENRY GRAY, P. K. S., Lecturer on Anatomy at St. George's Hospital, Loudon, &c. The Drawings by H. V. CARTER, M. D., Late Demonstrator of Anatomy at St. George's Hospital. THE DISSECTIONS JOINTLY BY THE AUTHOR AND DR. CARTER. In 07ie splendid imperial octavo volume of about 800 very large pages, with 363 large and elabo- rate engravings on wood. Price in, extra cloth, $6 25; in leather, raised bands, $7. The author has endeavored in this work to cover a more extended range of subjects than is customary in the ordinary text-books, by giving not only the details necessary for the student, but also the application of those details in the practice of medicine and surgery, thus rendering it not only a guide for the learner, but an admirable work of reference for the active practitioner. The engravings form a special feature in the work, many of them being the size of nature, nearly all original, and having the names of the various parts printed on the body of the cut, in place of figures of reference with descriptions at the foot. They thus form a complete and splendid series, which will greatly assist the student in obtaining a clear idt a of Anatomy, and will also serve to refresh the memory of those who may find in the exigencies of practice the necessity of recalling the details of the dissecting-room; while, combining as it does a complete Anatomical Atlas with a thorough treatise on descriptive, practical, and applied anatomy, the work will be found of essential service to physicians who receive students in their offices, assist- ing both teacher and pupil in laying the groundwork of a thorough medical education. *^* The large size of many of the illustrations prevents the selection of a favorable average specimen. A small one, however, is inserted to show the manner in which the lettering ac- companies the cuts. It will be observed that the attachments of the muscles are indicated by dotted lines. Occipital Bone — outer surface. As a full, systematic, and advanced treatise on anatomy, combining the various merits of the vol- umes of many countries, scientifically excellent, and adapted to all the ■wants of the student, we are not acquainted with any work in any language which can take equal rank with the one before us. — London Lancet, Sept. 11, ISoS. Mr. Gray's book, in excellency of arrangement and completeness of execution, exceeds any work on anatomy hitherto published in the English lan- guage, afi'ording a complete view of the structure of the human body, with especial reference to practi- cal surgery. Thus the volume constitutes a perfect book of reference for the practitioner, demanding a place in even the most limited library of the phy- sician or surgeon, and a work of necessity for the student to fix in his mind what he. has learned by the dissecting knife from the book of nature. — The Dublin Qvxirterly Journal of Med. Sei., JNov. 1S58. 10 BLAN CHARD AND LEA'S A LIBRARY ON HUMAN ANATOMY. HUMAli AXiTOMY, BY JONES QUAIN, M. D. EDITED BY RICHARD QUAIN, M. D., WM. SHARPEY, M.D., F.R.S., Professors of Anatomy and Physiology in University College, London. View of the Left Nasal Fossa. jFtrst fCmwuan, from Hi jfiitl ^Lonbon JEiilioir. , Edited by JOSEPH LEIDY, M. D., Professor of Anatomy in the University of Pennsylvania. Li two large arid Jiandsome octavo volumes, containing thirteen hundred pages, and five hun- dred and eleven beautiful engravings on wood, bound in leather. Price %%. It is indeed a work calculated to make an era in ana- tomical study, by placing before the student every de- partment of his science, with a view to the relative im- portance of each; and so skilfully have the different parts been interwoven, that no one who makes this work the basis of his studies, will hereafter have any excuse for neglecting or undervaluing any important parti- culars connected with the structure of the human frame; and whether the bias of his mind lead him in a more especial manner to surgery, physic, or physio- logy, he will find here a work at once so comprehen- sive and practical as to defend him from exclusiveness on the one hand, and pedantry on the other. — Monthly Journal and Retrospect of the Med. Sciences. TVe have no hesitation in recommending this treatise on anatomy as the most complete on that subject in the English language: and the only one. perhaps, in any language, which brings the state of knowledge forward to the most recent discoveries. — The Edinburgh Med. and Surg. Journal. Admirably calculated to fulfil the object for which it is intended. — Prminciol Medical Journal. The most complete Treatise on Anatomy in the English language. — Edinburgh Medical Journal. There is no work in the English language to be pre- ferred to Dr. Quain's Elements of Anatomy. — London Journal of Medicine. CARPENTER'S MANUAL OE PHYSIOLOGY. ELEMENTS OF physiology: INCLUDING PHYSIOLOGICM ANATOMY. BY AV. B. Carpenter, M.D.,F.R.S. AUTHOR OF " Human Physiology," " Compara- tive Physiology," &c. r\ Second American, from a late J'-^' and Revised London edition. -^ ' With 190 Illustrations. In one handsome octavo volume of 566 pages, leather, $3. Distribution of Olfactory Nerve. MEDICAL AND SCIENTIFIC PUBLICATIONS. SMITH AND HORNER'S ANATOMICAL ATLAS. ANATOMICAL ATLAS, ILLUSTRATIYE OF THE STRUGTUEE OF THE HUMAN BODY. Nerves of Ampulla of the Ear. By henry H. smith, M. D., Professor of Surgery in the University of Pennsylvania, UNDER THE SUPERVISION OF WM. E. HORNER, M. D., Late Professor of Anatomy in the University of Pennsylvania. With about Six Hundred and Fifty exquisite Illustrations on Wool. In one imperial octavo volume, extra cloth. Price S3. The great advantages possessed by vrood- engraving for scientific illustration, in the clearness and distinctness of its minute details, are fully shown in this work, containing as it does, in the compass of a single convenient volume, the general and special anatomy of all the component parts of the body. Commencing with the Bones and Ligaments, iffollows with the Muscular and Dermoid Systems, the Or- gans of Digestion and Generation, Respiration and Ciiculation, and concludes with the Nerv- ous System and the Senses. Not only is every separate organ and portion of the human frame thus biought distinctly and separately before the eye o( the student, but he is also presented with llie results of the recent microscopical investigations into the minute anatomy of the various tissues ; while the plan adopted, giving the plate and references on the same page, en- ables him to obtain a more definite impression of the objects, by avoiding the annoyance and interruption of turning backwards and for- wards. As a specimen of art, nothing superior to it has yet been produced, while the exceed- ingly low rate at which it is offered, places it within the reach of ever}'' member of the pro- fession. To country practitioners and students it will be found especially useful, as supplying in a great measure the place of skeletons and subjects. The plan of this Atlas is admirable, and its execu- tion superior to anything of the kind before published in this country. It is a real labor-saving affair, and we regard its publication as the greatest boon that could be conferred on the student of anatomy. It "ivill be equally valuable to the practitioner, by affording him an easy means of recalling the details learned in the dissecting-room, and which are soon forgotten. — American Medical Journal. Deep stated Muscles of the Hip. THE PRINCIPAL FORMS OF THE SKELETON AND OF THE TEETH. By Professor R. Owen, F. R. S , Author of "Lectures on Comparative Anatomy," &c. With seventy-six beautiful illustrations, fn one handsome volume, royal 12mo., of three hundred and thirty pages, extra cloth, $1 25. 12 BLANC HARD AND LEA'S Just Issued. IN ITS RELATIONS TO DESCRIPTIYE ANATOMY, PHYSIOLOGY, AND PATHOLOGY. WITH FOUR HUNDRED AND THIRTY-EIGHT ILLUSTRATIONS ON WOOD. BY E. R. PEASLEE, M. D., Professor of Physiology and Pathology in the New York Medical College, etc. .^v^^Tf^ ^\^^ C-J .--L - In one .-c beautifully printed octavo volume of 616 pages. Price, ^^ -A in leather. ) $3 75. '7 Insertion of Tendo Achillis hito Calcaneutn. It embraces a library upon the topics discussed within itself, and is just what the teacher and learner need Ano- ther advantage, by no means to be overlooked, everything of real value in the wide range which it embraces, is with great skill compressed into an been caused by the determination of the author to spare no pains in rendering it complete on every point : it is now, however, proceeding rapidly through the press, and the publisliers expect to have it in readiness lor the fall sei^sions of the medical schools. A COMPLETE ENCYCLOPEDIA OP MATERIA MEDICA. THE ELEMENTS OF MATERIA MEDICA AM THERAPEUTICS. BY JONATHAN PEREIRA, M. D., F. R. S., &c. 2[f)trTi gimerfcan HDitioit, lenlavfletr anU fimprobeTi Ijb Hje Slutljor. INCLUDING NOTICES OF MOST OF THE MEDICINAL SUBSTANCES IN USE IN THE CIVILIZED WORLD, AND FORMING AN ENCYCLOPEDIA OF MATERIA MEDICA. Edited by JOSEPH CARSON, M. D., Professor of Materia Medica and Pharmacy in the University of Pennsylvania, &c. hi two very large octavo volumes q/"2100 pages, hi small type, with over 450 illustrations, strongly hound in leather, with raised hands, price §9. The Second Volume will no longer be sold separate. MEDICAL BOTANY; OR, DESCRIPTIONS OF THE MORE IMPORTANT PLANTS USED IN MEDICINE, With their History, Properties, and Modes of Administration. By R. EGLESFELD GRIFFITH, M. D., Author of "A Universal Formulary," &c. WITH UPWARDS OF THREE HUNDRED ILLUSTRATIONS. In one large and very handsome octavo volume of 100 pages ; extra cloth, price $3. 30 BLANCHARD AND LEA'S MATERIA MEDICA AND THERAPEUTICS. INCLUDING THE PREPARATIONS OF THE PHARMACOPCEIAS OF LONDON, EDINBURGH, DUBLIN, AND OF THE UNITED STATES. WITH MANY NEW MEDICINES. By J. FORBES BOYLE, M.D.,F.R.S., &c. Edited BY JOSEPH CARSON, M.D., Professor of Materia Medica and Pharmacy in the University of Pennsylvania, &c. In one very handsome octavo volume of nearly seven hundred pages, with about one hundred beautiful illustrations on wood. Extra cloth, price $3. Senna and its Adulterations. This work is, indeed, a most valuable one, and will fill up an important vacancy that existed between Dr. Pereira's most learned and complete system of Mate- ria Medica, and the class of productions on the other extreme, which are necessarily imperfect from their small extent. — British and Foreign Medical Review. Ne-vr and Revised Edition. SYNOPSIS OF THE COURSE OF LECTURES ON MATERIA MEDICA AND PHAR- MACY, delivered in the University of Pennsylvania. By Joseph Carson, M. D., Professor of Materia Medica and Pharmacy in the University of Pennsylvania. Second edition, revised. In one very neat octavo volume, of 208 pages, cloth, f 1 50. THE THREE KINDS OF COD-LIVER OIL, comparatively considered, with their Chemical and Therapeutic properties. By D. L. De Jongh, M. D. 12mo., extra cloth, 75 cents. CABPENTER ON THE USE OF ALCOHOLIC LIQUORS IN HEALTH AND DISEASE. 'New edition, with a Preface by D. F. Condie, M. D. In one neat 12mo. volume, extra cloth, pp 178. {Just Issued.) 50 cents. THE MEDICAL FORMULARY; being a collection of Prescriptions, derived from the Writings and Practice of many of the most eminent Physicians of America and Europe : together with the usual Dietetic Preparations and Antidotes for Poisons. To which is added an Appendix, on the Endermic Use of Medicines, and on the uses of Ether and Chlo- roform. The whole accompanied with a few brief Pharmaceutic and Medical observations. By Benjamin Ellis, M. D. Tenth edition, revised and much extended, by Robert P. Tho- mas. M. D., Professor of Materia Medica in the Philadelphia College of Pharmacy, &c. In one neat octavo volume, extra cloth, of three hundred pages, price $1 75. After an examination of the new matter and the alterations, we believe the reputation of the work built up by the author, and the late distinguished editor, will continue to flourish under the auspices of the present editor, who has the industry and accuracy, and, we would say, conscientiousness requisite for the responsible task, — Ameriwn Journal of Pharmacy. It will prove particularly useful to students and young practitioners, ss the most important pre.scrip- tions employed in modern practice, which lie scattered through our medical literature, are here collected and conveniently arranged for reference. — Charleston Med. Journal and Review. MEDICAL AND SCIENTIFIC PUBLICATIONS. 31 New and Enlarged Edition — (Just Issued.) IST E W^ R EM E D I E S : WITH FORMUIJ] FOR THEIR PREPARATION AND ADMINISTRATION. BY ROBLEY DUNOLISON, M. D., Professor of the InslitiUes of Medicine, &c. ia the Jefferson Medical College, Philadelphia. Seventh Edition, with extensive Additions. In one very targe octavo volume of IIQ pages, leather, $3 75. As the value of a work sucih as the present is greatly enhanced by the thoroughness with which all the latest improvements and discoveries are embodied in its pages, the author has spared no pains in preparing the present edition to render it a complete exposition of the most recent aspect of therapeutic science. A large number of additional articles have been introduced, and many of the older ones have been rewritten to adapt them to the present condition of the subject. A very considerable increase in the size of the page has accommodated these additions without unduly swelling the bulk of the volume, and it is confidently presented as in every respect worthy a continuance of the very great favor which it has thus far received. This new and much enlarged edition of Dr. Dungli- pon's very valuable work will be welcomed by the pro- fession already well acquainted with its merits. The Tast research and thorough information which the author has brought to the execution of his labors, aie evident upon every page; and he is entitled to the hearty thanks of his brethren for this and the many other good things which he has done. Few would willingly be without this excellent volume who desire not only to know " new remedies" thoroughly, but also to test their efficacy. — Boston Med. and Surg. Journal, Aug. 1856. It may be considered almost a work of supereroga- tion to enter Into an elaborate criticism of a work ■which has reached its se.re.nth edition. The public has pronounced in the most authoritative manner its ver- dict, and we are cerlainly not disposed in the present instance to dispute its decision. In truth, such books a.« this will always be favorably received by the pro- fession of our country. They are labor-saving pro- ductions, which, at the expense of much research and reading to the author, condense in a convenient space the novelties and discoveries of the age. The present edition of this work is considerably enlarged and im- proved. The author, with his accustomed accuracy, has elaborated and ampliiied many of the articles but casually or imperfectly treated of in the former edi- tions ; and he has also added considerably to the list of new remedies. Abiut thirty new agents or novel applications of old remedies are introduced to the notice of the readerln this edition. — Va. Med. o.nd Surg. Journal, Sept. 1S56. As a work of reference upon all new remedies, this is one of the most complete with which we are ac- quainted. The quotations of authorities are extensive, minute, and carefully given; and we are satisfied that no medical man would regret following our advice to acquire it, as daily opportunities will occur in which he may both test its value and increase his own know- ledge, in searching for the practical information it affords. — British and Foreign Med.-Chirurgical Review, July, 1857. A. n3ISI^ENS.A.TOIl Y; Commentary on the Plmrmacopceias of Great Britain and the United States : COMPRISING THE Natural History, Description, Chemistry, Phar- macy, Action, Uses, and Doses of the Articles of the Materia Medica. BY R.V.CHRISTISON.M.D.,V.P.R.S.E.,&c. SECOND EDITION, REVISED AND IMPROVED. WITH A SUPPLEMENT Containing the most important New Remedies, WITH COPIOUS ADDITIONS BY R. EGLESFELD GRIFFITH, M.D. In one very latge and handsome octavo volume of over 1000 pages, with 213 engravings on wood; leather, price $3 50. It is not needful that we should compare it with the other pharmacopoeias extant, which enjoj' and merit the confidence of the profession : it is enough to say that it appears to us as perfect as a Dispensatory, in the present state of pharmaceutical science, could he made. If it omits any details pertaining to this branch of knowledge which the student has a right to expect in such a work, we confess the omission has escaped our scrutiny. We cordially recommend this work to such of our readers as are in need of a Dispensatory. They cannot make choice of a better. — Western Jour- nal of Medicine and Surgery. There is not in any language a more complete and perfect Treatise. — iVew Ycrrk Annalist. Aronitum napeltux. 32 BLANCHARD AND LEA'S THE GREAT LIBRARY OF PATHOLOGY— (Just Issued.) Complete in four handsome octavo volumes bound in two, containing about twelve hundred and fifty large pages; extra cloth, price $5 50. Medullary Carcinu?na. A MANUAL OF PATHOLOGICAL ANATOMY. BY CARL ROKITANSKY, M. D., Curator of the Imperial Pathological Museum and Professor at ihe University of Vienna. Vol. I —Manual of General Pathological Anatomy. Translated by W. E. Swaine. Vol. II.— Pathological Anatomy of the Abdominal Viscera. Translated by Edward Sieveking, M.D. Vol. III.— Pathological Anatomy of the Bones, Cartilages, Muscles, and Skin, Cellular and Fibrous Tissues, Serous and Mucous Membrane, and Nervous System. Translated by C. H. MooRE. Vol. IV. — Pathological Anatomy of the Organs of Respiration and Circulation. Translated by G. E. Day, M. D. To render this large and important work more easy of reference, and at the same time less cum- brous and costly, the four volumes have been arranged in two, retaining, however, the sepa- rate paging, &c. The publishers feel much pleasure in presenting to the profession of the United Slates the great work of Prof. Rokitansky, which is universally referred to as the standard of authority by the pathologists of all nations. Under the auspices of the Sydenham Society of London,' the combined labor of four translators has at length overcome the almost insuperable difficulties which have so long prevented the appearance of the work in an English dress, while the addi- tions made from various papers and essays of the author present his views on all the topics em- braced, in their latest published form. To a work so widely known, eulogy is unnecessary, and the publishers would merely state that it is said to contain the results of not less than thirty THOVSAND post-morte7n. examinations made by the author, diligently compared, generalized, and wrought into one complete and harmonious system. The profession is too well acquainted with the re- putation of Roliitansky's work to need our assurance that this is one of the most profound, thorough, and valuable hooks ever issued from the medical press. It is SMI fifeneris, and has no standard of comparison. It is only necessary to announce that it is issued in a form as cheap as is compatible with its size and pre- servation, and its sale follows as a matter of course. No library can be called complete without it. — Bvffalo Med. Journal. An attempt to give our readers any adequate idea of the vast amount of instruction accumulated in these volumes, would be feeble and hopeless. The effort of the distinguished author to concentrate into a small space his great fund of knowledge, has so charged his text with valuable truths, that any at- tempt of a reviewer to epitomize is at once paralyzed, and must end in a failure. — Western Lancet. As a hook of reference, therefore, this work must prove of inestimable value, and we cannot too highly recommend it to the profession. — Charleston Med. Journal and Review. This book is a necessity to every practitioner. — Am. Med. Monthly, The attempt to give our readers any adequate idea of the vast storehouse of knowledge contained in these volumes we feel to be utterlj' hopeless. The patient labors of nearly thirty years in a field so vast as almost to pass beyond our comprehension ; the accurate dis- sections made on more than thirty thousand human bodies are here carefully noted ; and the whole power of a vigorous and practical mind has been concentrated upon its execution. To the student of disease in its very formation, and in its endless variety of develop- ment; to the investigators of the true causes of symp- toms, and the many phenomena displayed at the bed- side; to all who strive to build up a sound and dis- creet method of treatment, based, as it must always be, on an accurate acquaintance with the pathological conditions of each case: we recommend this noble work as containing in itself everything which is needed to the fullest comprehension of the subject. — Va. Med. and Surg. Journal. It is, everywhere, in fact, acknowledged to be fore- most and unrivalled in its department. Only the most earnest professional ardor, the most persevering labor, could have enabled its author to amass the immense amount of information contained in its pages. — Medi- cal Examiner. MEDICAL AND SCIENTIFIC PUBLICATIONS. 33 New and thoroughly Revised Edition — (Just Issued.) ELEMENTS OF PATHOLOGICAL ANATOMY. BY SAMUEL D. GROSS, M. D., Professor of Surgery in the Jefferson Medical College, Philadelphia. Third Edition, Modified and thoroughly Revised, ILLUSTRATED BY THREE HUNDRED AND FORTY-TWO ENGRAVINGS ON WOOD. In one large and very handsome octavo volume, of nearly 800 pages. Price, in extra cloth, $4 75; leather, raised bands, $5 25. Keloid growth of the Face. The very rapid advances in the Science of Pathological Anatomy during the last few years have rendered essential a thorough modification of this work, with a view of making it a correct exponent of the present state of the subject. The very careful manner in which this task has been executed, and the amount of alteration which it has undergone, have enabled the author to say that "with the many changes and improvements now introduced, the work majr be re- garded almost as a new treatise," while the efforts of the author have been seconded as regards the mechanical execution of the volume, rendering it one of the handsomest productions of the American press. A very large number of new and beautiful original illustrations have been introduced, and the work, it is hoped, will fully maintain the reputation hitherto enjoyed by it of a complete and practical exposition of its difficult and important subject. We most sincerely congratulate the author on the successful manner in which he has accomplished his proposed object. His book is most admirably calcu- lated to fill up a blank which has long been felt to exist in this department of medical literature, and as such must become very widely circulated amongst all classes of the profession. — Dublin Quarterly Journ. of Med. Science, Nov. 1857. We have been favorably impressed with the general manner in which Dr. Gross has executed his task of affording a comprehensive digest of the present state of the literature of Pathological Anatomy, and have much pleasure in recommending his work to our read- ers, as we believe one well deserving of diligent pe- rusal and careful study. — Montreal Med. Chron., Sept. 1857. ATLAS OF PATHOLOGICAL HISTOLOGY. BY GOTTLEIB GLtJGB, M. D., Professor of Physiology and Pathological Anatomy in the University of Brussels, &c. TRANSLATED, WITH NOTES AND ADDITIONS, BY JOSEPH LEIDY, M. D., Professor of Anatomy in the University of Pennsylvania, &c. In one handsoine volume, large imperial quarto, with 320 Figures, plain and colored, on twelve copperplate engravings, price $5 00. This being, as far as we know, the only work in which pathological histology is separately treated of in a comprehensive manner, it will, we think, for this reason, be of infinite service to those who desire to investigate the subject systematically, and who have felt the difficulty of arranging in their minds the un- 3 connected observations of a great number of authors. The development of the morbid tissues, and the format tion of abnormal products, may now be followed and studied with the same ease and satisfaction as the best arranged system of Physiology. — American Medical Journal. 34 BLANCHARD AND LEA'S PATHOLOGICAL MANUAL- (Lately Published.) Pyelitis. Fibrinous Deposits in Granular Kidney. A MANUAL OF PATHOLOGICAL ANATOMY. BY C. HANDFIELD JONES, M. D., F. E. S., Lecturer on Physiology at St. Mary's Hospital, &c. AND EDWARD H. SIEVEKING, M. D., Lecturer on Materia Medica at St. Mary's Hospital. FIRST AMERICAN EDITION, REVISED. With three hundred and ninety-seven Engravings on "Wood. Li one large and very handsome octavo volume of seven hundred and thirty-four pages, leather, $3 75. As a concise text-book, containing, in a condensed form, a complete outline of what is known in the domain of Pathological Anatomy, it is perhaps the hest work in the English language. Its great merit consists in its completeness and brevity, and in this respect it supplies a great desideratum in our literature. Heretofore the student of pathology was obliged to glean from a great number of monographs, and the field was so extensive that but few cultivated it with any de- gree of success. The authors of the present work have sought to correct this defect by placing before the reader a summary of ascer- tained facts, together with the opinions of the most eminent patho- logists both of the Old and New World. As a simple work of reference, therefore, it is of great value to the student of pathological anatomy, and should be in every physician's library. — Western Lancet. AYe urge upon our readers and the profession generally the import- ance of informing themselves in regard to modern views of pathology, and recommend to them to procure the work before us as the best means of obtaining this information. — Stethoscope. In offering the above titled work to the public, the authors have not attempted to intrude new views on their professional brethren, but simply to lay before them, what has long been wanted, an outline of the present condition of pathological anatomy. In this they have been completely successful. The work is one of the best compila- tions which we have ever perused. The opinions and discoveries of all the leading pathologists and physiologists are engrossed, so that by reading any subject treated in the book you have a .synopsis of the views of the most approved authors. — Charleston Medical Journal and Review. We have no hesitation in recommending it as worthy of careful and thorough study by every member of the profession, old or young. — N. W Med. and Surg. Journal. to be largely useful, as it suits itself to those busy m en who have little time for minute investigation, and prefer a summary to an elaborate treatise. — Buffalo Medical Journal. Osteophytes in lower end of Femur. From the casual examination we have given we are inclined to regard it as a text-book, plain, rational, and intelligible, such a book as the practical man needs for daily reference. For this reason it will be likely GENERAL PATHOLOGY; As Conducive to the Establishment of Rational Principles for the Prevention and Cure of Dis- ease. A Course of Lectures delivered at St. Thomas's Hospital. By John Simon, F. R. S &c. In one neat octavo volume, extra cloth, $1 25. MEDICAL AND SCIENTIFIC PUBLICATIONS. 35 BARCLAY ON DIAGNOSIS— A New Work, Now Ready (1858). A MANUAL OF MEDICAL DIAGNOSIS; be™ an analysis of the signs and symptoms of disease. By a. W. BARCLAY, M. D., Assistant Physician to St. George's Hospital, &o. Ill one neat octavo volume of 424 pages ; extra cloth, price f 2. Of works exclusively devoted to this important branch, our profession has at command, comparative- ly, hut few, and therefore, in the publication of the present work, Messrs. Blanchard & Lea have conferred a great favor upon us. Dr. Barclay, from having occu- pied, for a long period, the position of Medical Regis- trar at St. George's Hospital, possessed advantages for correct observation and reliable conclusions, as to the significance of symptoms, which have fallen to the lot of but few, either in his own or any other country. He has carefully systematized the results of his ob- servation of over twelve thousand patients, and by his diligence and judicious classification, the profession has been presented with the most convenient and re- liable work on the subject of Diagnosis that it has been our good fortune ever to examine; we can, there- fore, say of Dr. Barclay's work, that from his system- atic manner of arrangement, his work is one of the best works " for reference" in the daily emergencies of the practitioner, with which we are acquainted; but, at the same time, we would recommend our read- ers, especially the younger ones, to read thoroughly and study diligently the whole work, and the '•' emer- gencies" will not occur so often. — Southern Med. and Surg. Journal, March, 1S58. To give this information, to supply this admitted deficiency, is the object of Dr. Barclay's Manual. The task of composing such a work is neither an easy nor a light one: but Dr. Barclay has performed it in a manner which meets our most unqualified approba- tion. He is no mere theorist; he knows his work tho- roughly, and in attempting to perform it, has not ex- ceeded his powers. — British Med. Journal, Dec. 5, 1867. We venture to predict that the work will be deserv- edly popular, and soon become, like Watson's Practice, an indispensable necessity to the practitioner. — iV". O. Med. Journal, April, 1858. An inestimable work of reference for the young practitioner and student. — Nashville Medical Journal, May, 1858. We hope the volume will have an extensive circula- tion, not among students of medicine only, but prac- titioners also. They will never regret a faithful study of its pages. — Cincinnati Lancet, March, 1858. This Manual of Medical Diagnosis is one of the most scientific, useful, and instructive works of its kind that we have ever read, and Dr. Barclay has done good ser- vice to medical science in collecting, arranging, and analyzing the signs and symptoms of so many dis- eases.— iV. J. Med. and Surg. Reporter, March, 1858. We hail the appearance of this valuable book, com- ing to us in its present exceedingly neat style, as an important acquisition to medical literattire. It is a work of high merit, both from the vast importance of the subject upon which it treats, and also from the real ability displayed in its elaboration. In conclu- sion, let us bespeak for this volume that attention of every student of our art which it so richly deserves — that place in every medical library which it can so well adorn. — T!ie Peninsular and Independent Medical Journal, Sept. 1858. Restricted as we are to certain limits, we can hardly do more than give this brief synopsis, which will serve, however, to convey to the reader an idea of the scope and the number and variety of diagnostic details of Dr. Barclay's volume. It was much wanted, and is full of instruction on a branch of pathology which furnishes we will not say the only, but certainly the chief and the safest, indications for the treatment of disease. — N. A. Medico-Chir. Review, May, 1858. We conclude by assuring the honest and earnest student who has acquainted himself sufficiently with the principles of physiology and the details of anato- my, that he could not better aid or advance his clinical investigations, than by carrying this work with him to the bedside of his patients, and the junior practi- tioner will find much in its small compass to repay a careful and perhaps repeated perusal. — Charleston, Med. Journal and Review, May, 1858. The author writes with a confidence which severe and careful study can alone justify. There is a full table of contents, and a most copious index, which in such a work is invaluable. In conclusion, we can ho- nestly recommend this work to the profession — to the junior members of it as a handbook of study; to the seniors, as a useful compendium of things to be re- membered in cases of obscurity. — Medical Times and Gazette, London, Nov. 14, 1857. CYCLOP J]DIA OF PRACTICAL MEDICIM; COMPRISING Treatises on the Nature and Treatment of Diseases, Materia ledica and Therapeutics, Diseases of Women and Children, Medical Jurisprudence, &c. &c. EDITED BY JOHN FORBES, M.D., F. R. S., ALEXANDER TWEEDIE, M. D., F. R. S., AND JOHN CONNOLLY, M. D. Revised, with Additions, BY ROBLEY DUNGLISON, M. D. COMPLETE, IN FOUR LARGE SUPER-ROYAL OCTAVO VOLUMES. Containing thirty-two hundred and fifty-four unusually large pages, in doiible cokimiis, printed on good paper, vnth a new and clear type. • THE WHOLE WELL AND STRONGLY BOUND IN LEATHER, WITH RAISED BANDS AND DOUBLE TITLES. Price $12 00. This work contains no less than TOUR HUNDRED AND EIGHTEEN DISTINCT TREATISES, By Sixty-eight Distinguished Physicians. The most complete work on Practical Medicine ex- tant ; or, at least, in our language. — Buffalo Medical and Surgical Journal. For reference, it is above all price to every practi- tioner. — Western Lancet. 36 BLANCHARD AND LEA'S New and Much Improved Edition. PRINCIPLES OF MEDICINE. AN ELEMENTARY VIEW OF THE CAUSES, NATURE, TREATMENT, DIAGNOSIS AND PROGNOSIS OF DISEASE, WITH BRIEF REMARKS ON HYGIENICS, OR THE PRESERVATION OF HEALTH. BY CHARLES J. B. WILLIAMS, M.D.,F.R.S. A new American, from the Third and Revised Liondon Edition. In 0716 neat octavo volume, of about five hundred large pages, leather, $2 50. The very recent and thorough revision which this work has enjoyed at the hands of (he au- thor has: brought it so completely up to the present stale of the subject that in reproducing it no additions have been found neces^sary. The success which the work has heretofore met shows that its importance has been appreciated, and in its present form it will be found eminentl}' wor- thy a continuance of the same favor, possessing as it does the strongest claims to the attention of the medical student and practitioner, from the admirable manner in which the various inqui- ries in the different branches of pathology are investigated, combined, and generalized by an experienced practical physician, and directly applied to the investigation and treatment of disease. We find that the deeply-interesting matter and style of this book have so far fascinated us, that we have unconsciously hung upon its pages, not too long, in- deed, for our own profit, but longer than reviewers can be permitted to indulge. We leave the further analysis to the student and practitioner. Our judg- ment of the work has already been sufficiently ex- pressed. Itis a judgment of almost unqualified praise. The work is not of a controversial, but of a didactic character ; and, as such, we hail it, and recommend it for a text-book, guide, and constant companion to every practitioner and every student who wishes to extricate himself from the well-worn ruts of empiri- cism, and to base his practice of medicine upon prin- ciples — London Lancet, Dec. 27, 1856. A text-book to which no other in oxxT language is comparable. — Charleston Me.dical Journal. No work has ever achieved or maintained a more deserved reputation. — Virginia Med. and Surg. Journ. The Principles of Medicine of Dr. Williams has, by common consent, become one of the classics of our profession. Few works have done more towards ac- complishing that union of the science and practice of medicine so indispensable for its perfection, and which are too apt,to be found separate from each other — a separation, the inevitable tendency of which must ever be to favor empiricism. The rapid sale of three edi- tions of this work in our country we regard as a marked tribute to its value, and as no less compli- mentary to the discrimination and appreciation of the profession here in giving rise to such a demand. — jV. Y. Medical Times. The work as now presented to the public, is perhaps the most perfect of any other treating on similar sub- jects; it combines the science and the art, the theory and the practice, in a most masterly manner, and we feel confident that as knowledge of the practical views and scientific principles laid down in the book become generally known, medicine — practical medicine— will advance, in the same proportion, to a greater perfec- tion and certainty. — N. Orleans Med. and Su7-g. Journ. There is no work in medical literature which can fill the place of this one. It is the Primer of the young practitioner, the Koran of the scientific one. Three large editions of it have already been exhausted in the United States, and now the fourth is presented. It must have, so long as the size of the volume remains uncumbersome, the first place among pathological au- thorities. We feel warranted in saying that no medi- cal book has yet been written which contains so much in the small number of pages which compose this one, and yet it is complete. It takes up disease at its very foundation, and treats of its fundamental nature in a logical and inductive style. — The Stethoscope. This exceedingly valuable work is the best, we be- lieve, in the whole round of medical literature. The division of the different subjects is excellent. The au- thor's method of investigation and mode of expression, in our judgment, are faultless. We can most cheer- fully commend the work as the best that has ever appeared on the principles of medicine, and we would advise young practitioners especially to furnish them- selves with a copy, as well for the value of the informa- tion it contains as for the facilities it will afford them in the prosecutionof their own investigations. — Soutli- ern Journal of the Med. and Phys Sciences. The best exposition in our language, or, we believe, in any language, of rational medicine, in its present improved and rapidly improving state. — British and Foreign Medico-Chirurgical Review. WHAT TO OBSERVE AT THE BEDSIDE AND AFTER DEATH, IN MEDICAL CASES. Published under the authority of the London Society for Medical Observation. A new American from the Se- cond and Revised London edition. In one very handsome volume, royal 12mo , extra cloth, A MANUAL OF CLINICAL MEDICINE AND PHYSICAL DIAGNOSIS. By Thomas H. Tanner, M. D., Physician to the Hospital for Women, &c. Second American edition. In one neat volume, small 12mo., extra cloth, 88 cts. ; or in flexible cloth for the pocket, 80 cts. DUNGLISON'S PRACTICE. THE PRACTICE OF MEDICINE: A Treatise on Special Pathology and Thera- peutics. By RoBLEY DuNGLisoN, M. D., Professor of the Institutes of Medicine in Jefferson Medical College, Philadelphia. Third and revised edition. In two large octavo volumes of about fifteen hundred pages; leather, f6 25. MEDICAL AND SCIENTIFIC PUBLICATIONS. 37 New and much enlarged edition of "WATSON'S PKACTICE"— Just Ready (185S). LECTURES ON THE PRKfCIPLES AND PRACTICE OF PHYSIC. DELIVERED AT KING'S COLLEGE, LONDON. BY THOMAS WATSON, M. D., Late Physician to the Middlesex Hospital, &c. ^ mbs ^nt^ritait, from t^e last llebiseb nnb inlargcb ^itgltsl^ ©iritioit. With Additions, by D. FRANCIS CONDIE, M.D., Author of "A Practical Treatise on the Diseases of Children," &c. WITH ONE HUNDRED AND EIGHTY-FIVE ILLUSTRATIONS ON WOOD. Bi one very large and handsome vohtme, imperial octavo^ of over 1200 large and closely printed pages, in sm,all type; the whole strongly botind in leather, with raised bands. Price $4 25. The publishers feel that they are rendering a service to the American profession in presenting at so very mode- rate a price this vast body of sound practical information. Whether as a guide for the student entering on a course of instruction, or as a book of reference for daily con- sultation by the practitioner, "Watson's Practice" has long been regarded as second to none; the soundness and fulness of its teachings, the breadth and liberality of its views, and the easy and flowing style in which it is writ- ten having won for it the position of a general favorite. That this high reputation might be fully maintained, the author has subjected it to a thorough revision ; every portion has been examined with the aid of the most re- cent researches in pathology, and the results of modern investigations in both theoretical and practical subjects have been carefully weighed and embodied throughout its pages. The watchful scrutiny of the editor has likewise introduced whatever possesses immediate importance to the American physician in relation to diseases incident to our climate which are little known in England, as well as those points in which experience here has led to different modes of practice ; and he has also added largely to the series of illustrations, believing that in this manner valu- able assistance may be conveyed to the student in eluci- dating the text. The work will, therefore, be found thoroughly on a level with the most advanced state of medical science on both sides of the Atlantic. The additions which the work has received are shown by the fact that notwithstanding an enlargement in the size of the page, more than two hundred additional pages have been necessary to accommodate the two large vo- lumes of the London edition (which sells at ten dollars) within the compass of a single volume, and in its present form it contains the matter of at least three ordinary oc- tavos. Believing it to be a work which should lie on the table of every physician, and be in the hands ot every student, the publishers have put it at a price within the reach of all, making it one of the cheapest books as yet presented to the American profession, while at the same time the beauty of its mechanical execution renders it an exceedingly attractive volume. Stomach contracted from chronic ulctraaon. 38 BLANCHARD AND LEA'S New and Improved Edition — Now Ready (June, 1859). ELEMENTS OE MEDICINE: A COMPENDIOUS VIEW OF PATHOLOGY AND THERAPEUTICS; OR, THE HISTORY AND TREATMENT OF DISEASES. BY SAMUEL H. DICKSON, M. D., Professor of the Practice of Medicine in the Jefferson Medical College of Philadelphia. In one large and handsome octavo volume of 750 pages ; leather, $3 75. The steady demand which has so soon exhausted the first edition of this work, sufficiently shows that the author was not mistaken in supposing that a volume of this character was need- ed — an elementary manual of practice, which should present the leading principles of medicine with the practical results, in a condensed and perspicuous manner. Disencumbered of unneces- sary detail and fruitless speculations, it embodies what is most requisite for the student to learn, and at the same time what the active practitioner wants when obliged, in the daily calls of his profession, to refresh his memory on special points. The clear and attractive style of the au- thor renders the whole easy of comprehension, while his long experience gives to his teachings an authority everywhere acknowledged. Few physicians, indeed, have had wider opportunities for observation and experience, and few, perhaps, have used them to better purpose. As the result of a long life devoted to study and practice, the present edition, revised and brought up to the date of publication, will doubtless maintain the reputation already acquired as a condensed and convenient American text-book on the Practice of Medicine. A few notices of the first edition are appended. This book is eminently what it professes to be; a dis- tinguished merit in these days. Desisned for " Teachers and Students of Medicine," and admirably suited to their wants, we think it will be received, on its own merits, with a hearty welcome. — Boston Med. and Surg. Journal. The volume is admirably adapted to supply a want long since felt by the American student and young practitioner of medicine, with reference to whom it has evidently been prepared. This class will find it a ju- dicious and valuable compend of the elements of medicine. — N. T. Journal of Medicine, Sept. 1856. Indited by one of the most accomplished writers of our country, as well as by one who has long held a high position among teachers and practitioners of medicine, this work is entitled to patronage and care- ful study. The learned author has endeavored to con- dense in this volume most of the practical matter con- tained in his former productions, so as to adapt it to the use of those who have not time to devote to more extensive works. — Southern Med. and Surg. Journal. We can strongly recommend Dr. Dickson's work to our readers as one of interest and practical utility, well deserving of a place in their libraries as a book of reference; and we especially commend the first part as presenting an admirable outline of the principles of medicine. — Dublin Quarterly Journal. This volume is designed as a text-book for teachers and students; but its merits extend far beyond its modest dedication; it is a complete treatise upon me- dicine, and one that will stand the test of years. The arrangement is simple, a feature oftentimes obscured in otherwise excellent works. This Treatise is a valu- able addition to our medical literature, and in the clear and accurate descriptions, purity, and simplicity of style, and soundness of precept, the reader will find much to admire and adopt, and not a little that calls for deep reflection. We cordially recommend this volume to our readers, whether old practitioners or students, for we take it that the physician should al- ways be a student. — American Lancet. Prof Dickson's work supplies, to a great extent, a desideratum long felt in American medicine. — iV. O. Med. and Surg. Journal. Estimating this work according to the purpose for which it is designed, we must think highly of its me- rits, and we have no hesitation in predicting for it a favorable reception by both students and teachers. Not professing to be a complete and comprehensive treatise, it will not be found full in detail, nor filled with discussions of theories and opinions, but em- bracing all that is essential in theory and practice, it is admirably adapted to the wants of the American student. Avoiding all that is uncertain, it presents more clearly to the mind of the reader that which i.s established and verified by experience. The varied and extensive reading of the author is conspicuously apparent, and all the recent improvements and dis- coveries in therapeutics and pathology are chronicled in its pages. — Charleston Med. Journal. A New Text-Book on Practice— (Lately Issued.) A MANUAL OE THE PRACTICE OE MEDICINE. BY GEORGE H. BARLOW, M. D., Physician to Guy's Hospital, London, &c. With Additions by D. E. CONDIE, M.D., Author of "A Practical Treatise on the Diseases of Children," &c. I7i one handsome octavo volume of over six Mmdred pages; leather, $2 75. We recommend Dr. Barlow's Manual in the warm- est manner as a most valuable vade-mecum. We have had frequent occasion to consult it, and have found it clear, concise, practical, and sound. It is eminently a practical work, containing all that is essential, and avoiding useless theoretical discussion. The work supplies what has been for some time wanting, a manual of practice based upon modern discoveries in pathology and rational views of treatment of disease It is especiiilly intended for the use of students and junior practitioners, but it will be found hardly less useful to the experienced physician. The American editor has added to the work three chapters— on Cholera Infantum, Yellow Fever, and Cerebro-spinal Meningitis. These additions, the two first of which are indispensable to a work on practice destined for the profession in this countr}', are executed with great judgment and fidelity, by Dr. Condie, who has also succeeded happily in imitating the conciseness and clearness of style which are such agreeable character- istics of the originial 'hook.— Boston Med. and Surg. Journal. MEDICAL AND SCIENTIFIC PUBLICATIONS. 39 LA ROCHE ON YELLOW FEVER— (Just Issued. YELLOW FEYER, CONSIDERED IN ITS HISTORICAL, PATHOLOGICAL, ETIOLOGICAL, AND THERAPEUTICAL RELATIONS: INCLUDING A Sketch, of the Disease as it has occurred in Philadelphia from 1699 to 1854. With an Bzamination of the Connedions between it and the Fevers known under the same name in other parts of Temperate as well as in Tropical Regions, BY R. LA ROCHE, M. D. In two large and handsome octavo volumes of nearly 1500 pages ; extra cloth, $7. The publishers are happy in being able at length to present to ihe profession this great work, which they are assured will be regarded as an honor to the medical literature of the country. As the result of many years of personal observation and study, as embodying an intelligent re- sume of all that has been written regarding the disease, and as exhausting the subject in all its various aspects, these volumes must at once talce the position of the standard authority and work of reference on the many important questions brought into consideration. From Professor S. H. Diclcson, Ckarlestmi, S. C. A monument of intelligent and well-applied re- search, almost without example. It is, indeed, in it- self, a large lilirary, and is destined to constitute the special resort as a book of reference, on the subject of which it treats, to all future time. We have not time at present, engaged as we are, by day and by night, in the work of combating this very disease, now prevailing in our city, to do more than give this cursory notice of what we consider as un- doubtedly the most able and erudite medical publica- tion our country has yet produced. But in view of the startling fact, that this, the most malignant and unmanageable disease of modern times, has for several years been prevailing in our country to a greater ex- tent than ever before; that it is no longer confined to either large or small cities, but penetrates country villages, plantations, and farm-houses; that it is treated with scarcely better success now than thirty or forty years ago ; that there is vast mischief done by ignorant pretenders to knowledge in regard to the disease, and in view of the probability that a majority of southern physicians will be called upon to treat the disease, we trust that this able and comprehensive treatise will be very generally read in the south. — Memphis Med. Recorder. This is decidedly the great American medical work of the day — a full, complete, and systematic treatise, unequalled by any other upon the all-important sub- ject of Yellow Fever. The laborious, indefatigable, and learned author has devoted to it many years of arduous research and careful study, and the result is such as will reflect the highest honor upon the author and our country. — Southern Med. and Surg. Journal. The genius and scholarship of this great physician could not have been better employed than in the erec- tion of this towering monument to his own fame, and to the glory of the medical literature of his own coun- try. It is destined to remain the great authority upon the subject of Yellow Fever. The student and phy- sician will find in these volumes a risumi of the sum total of the knowledge of the world upon the awful scourge which they so elaborately discuss. The style is so soft and so pure as to refresh and invigorate the mind while absorbing the thoughts of the gifted author, while the publishers have succeeded in bringing the externals into a most felicitous harmony with the inspiration that dwells within. Take it all in all, it is a book we have often dreamed of, but dreamed not that it would ever meet our waking eye as a tangible reality. — Nashville Journal nf Medicine. We deem it fortunate that the splendid work of Dr. La Roche should have been issued from the press at this particular time. The want of a reliable digest of all that is known in relation to this frightful malady has long been felt — a want very satisfactorily met in the work before us. We deem it but faint praise to say that Dr. La Roche has succeeded in presenting the profession with an able and complete monograph, one which will find its way into every well ordered library. — Va. Stethoscope. Although we have no doubt that controversial trea- tises on the mode of origin and propagation of the fever in question will, as heretofore, occasionally appear, yet it must be some time before another systematic work can arise in the face of so admirable and care- fully executed a one as the present. It is a mine of information, quite an encyclopedia of references, and resume of knowledge relative to what has been re- corded upon the subject. — Loivlon Lancet. A miracle of industry and research, constitviting a complete library of reference on the disease of which it treats. — Dublin Quarterly Journal. Dr. La Roche's work embodies all that is wanted. It is a compendium of the whole vast literature of Yellow Fever. Thanks to his labors, the medical scholar who desires to be profoundly conversant with all that pertains to the subject need not go beyond these two portly volumes. As embodying whatever is important in all that has been hitherto written on the subject, it will be a work for reference not less valuable in ages to come than now. Its merit, how- ever, by no means consists solely in its completeness as an encyclopa^dian work. The author presents the conclusions to which he is led by a philosophical in- vestigation of the facts and opinions gathered from past and contemporaneous publications. Of the sound- ness of the conclusions the reader can judge from the data which are spread before him. — Buffalo Med. Jour- nal, Sept. 1856. By the same Author. PNEUMONIA; its Supposed Connectiot], Pathological and Etiological, with Autumnal Fevers, including an Inquiry into the Existence and Morbid Agency of Malaria. In one handsome octavo volume, extra cloth, of 500 pages. $3. A more simple, clear, and forcible exposition of the groundless nature and dangerous tendency of certain pathological and etiological heresies, has seldom been presented to our notice. — N. Y. Journal of Medicine and Collateral Science. This work should be carefully studied by Southern physicians, embodying as it does the reflections of an original thinker and close observer on a subject pecu- liarly their own. — Virginia Medical and Surgical Jour- nal. 40 BLANCHARD AND LEA'S New and Improved Edition— Just Issued. A. 9 IN ONE VERY HANDSOME OCTAVO VOLUME, OF FIVE HUNDRED PAGES. ■WITH Four exquisite COLOEED PLATES, AITD NUMEROUS WOOD-CUTS. Extra cloth, S3 00. Suction of small Portal Vein and Canal. ON DISEASES OF THE LIVER. BY GEORGE BCDD, M.D., F. E. S., Professor of Medicine in King's College, London, &c. Spirit g^merkEit, from t^£ Cljirb anb finlarg^b l^onbon 6btlion, Has fairly established for itself a place among the classical medical literature of England. — Brilish and Foreign Medico-Chir. Review, July, 1857. Dr. Budd's Treatise on Diseases of the Liver is now a standard work in Medical literature, and during the intervals which have elapsed between the successive editions, the author has incorporated into the text the most striking novelties which have characterized the recent progress of hepatic physiology and pathology ; 80 that although the size of the book is not percepti- bly changed, the history of liver diseases is made more complete, and is kept upon a level with the progress of modern science. It is the best work on diseases of the liver in any language. — London Med. Times and Gazette., June 27, 1857. This work, now the standard book of reference on the disea.ses of which it treats, has been carefully re- vised, and many new illustrations of the views of the learned author added in the present edition. — Dublin Quarterly Journal, Aug. 1857. By the same Author— (Just Issued.) ON THE ORGANIC DISEASES AND FUNCTIONAL DISORDERS OF THE STOIVI^CH. In one neat octavo voluTne, of two hundred and fifty pages, extra cloth, $1 50. LALLEMAND AND WILSON ON SPERMATORRHCBA— (Now Ready). A PRACTICAL TREATISE ON THE CAUSES, SYMPTOMS, AND TREATMENT OF SPERMATORRHCBA. By M. Lallemand. Translated and edited by Henry J. McDougall Third American edition. To whicli is added— ON DISEASES OF THE VESICUL.'E SEMINALES, and their associated organs. With special re- ference to the Morbid Secretions of the Prostatic and Urethral Mucous Membrane. By Mae- Ris WiLSONj M. D. Ill one neat octavo volume, of about 400 pp., extra cloth. $2. FEVERS, THEIR DIAGNOSIS, PATHOLOGY, AND TREATMENT. Prepared and edited, with large Additions, from the Essays on Fever in Tweedie's Library. By Meredith Clymer, M. D. In one octavo volume, of six hundred pages, $1 50. HUGHES' CLINICAL INTRODUCTION TO THE PRACTICE OF AUSCULTATION AND OTHER MODES OF PHYSICAL DIAGNOSIS, IN DIS- EASES OF THE LUNGS AND HEART. Second American, from the second London edition. 1 vol. royal 12mo., extra cloth, pp. 304. $1. COPLAND ON PALSY.— Of the Causes, Nature, and Treatment of Palsy and Apoplexy. In one volume, royal 12mo., 80 cents. BLAKISTON'S PRACTICAL OBSERVATIONS ON CERTAIN DISEASES OF THE CHEST, and on the Principles of Auscultation. In one vol., cloth, 8vo., pp. 384. $1 25. BUCKLER ON THE ETIOLOGY, PATHOLOGY, AND TREATMENT OF FIBRO-BRONCHITIS AND RHEUMATIC PNEU.MONIA. In one Svo. vol., ex- tra cloth, pp. 150. $1 25. MEDICAL AND SCIENTIFIC PUBLICATIONS. 41 FLINT ON RESPIRATORY ORGANS— (Just Issued.) PHYSICAL EXPLORATION AND DIAGNOSIS OF DISEASES AEFECTING THE RESPIRATORY ORGANS. BY AUSTIN FLINT, M. D., Professor of Clinical Medicine and Pathologj' in the University of Buffalo, &c. I7i one large and handsome octavo volume of six hundred and thirty-six pages ; extra cloth, $3. Dr. Flint is one of the most industrious and ener- p;etio men in the medical profession of this country. His previous contributions to our medical literature have won for him both American and European repu- tation, and we assure our readers that the present volume is full of valuable and interesting matter. We unhesitatingly commend the book to all who wish to become well acquainted with thoracic diseases and the signs by which they may be distinguished. — N. W. Med. and Surg. Journal, Nov. 1856. We have selected these points in the physical ex- ploration of the chest not only from their importance, but to show the manner in which Dr. Flint handles his subject. Our readers will, we doubt not, agree with us in the opinion that he has done this carefully, thoroughly, and judiciously. — Charleston Med, Jour- nal, Nov. 1856. We can only state our general impression of the A work of original observation of the highest merit. We recommend the treatise to every one who wishes to become a correct auscultator. Based to a very large extent upon cases numerically examined, it carries the evidences of careful study and discrimination upon every page. It does credit to the author, and. through him, to the profession in this country. It is, what we cannot call every book upon aiiscultation, a readable book. — Am. Journal Med. Sciences. A work, of which we cannot but admire the spirit that has presided over its composition. There is an evident accuracy aimed at throughout by means of the carefully noted cases, and a searching after truth which recommends the volume highly to the attention of the profession. — Med. Examiner. This is the most elaborate work devoted exclusively to the physical exploration of diseases of the lungs, high value of this work, and cordially recommend it i with which we are acquainted in the English lan- to all. We regard it, in point both of arrangement \ guage. From the high standing of the author as a and of the marked ability of its treatment of the sub- jects, aS destined to take the first rank in works of this class. So far as our information extends, it has at present no equal. To the practitioner, as well as the student, it will be invaluable in clearing up the diag- nosis of doubtful cases, and in shedding light upon difficult phenomena. — Buffalo Med. Journal. clinical teacher, and his known devotion, during many years, to the study of thoracic diseases, much was to be expected from the announcement of his determin- ation to embody in the form of a treatise, the results of his study and experience. — Boston Med. and Surg. Journal. By the same Author— (In Press.^ THE DIAGNOSIS, PATHOLOGY, AND TREATMENT OF DISEASES OE THE HE^HT. In one neat octavo volume, of nearly 500 pages. This work is now so far advanced that the publishers can promise it with confidence for the autumn of 1859. The reputation of the author, and the attention which he lias long paid to this department of pracical medicine, are sufficient guarantee that the present volume will supply the want which has long been felt of a complete and authoritative treatise on the subject. Just Issued, MEDICAL NOTES AND REFLECTIONS. BY SIK HENRY HOLLAND, Bart., F. E. S., Physician in Ordinary to the Queen, &c. FRON/1 -THE THIRD AND ENt-ARGED ENQI-ISH EDITION. 1)1 one handsome octavo volume of about five hundred pages, extra doth, $3 GO. Just Issued. CLINICAL LECTURES ON CERTAIN DISEASES OF THE URINARY ORGANS, AND ON DROPSIES, BY ROBERT BENTLEY TODD, M. D., F. R. S., Physician to King's College Hospital, &c. In one handsome octavo volume, of 270 pages, extra cloth, $1 50. By the same Author — (In Press.) CLINIC AiTlECTURES ON OEKTAIN ACUTE DISEASES. In one neat octavo volume, extra cloth. 42 BLANCHARD & LEA'S Publishing in the "Medical News and Library," 1858 and 1859. PATHOLOGICAL AND PRACTICAL OBSERVATIONS OX DISEASES OF THE ALIMENTARY CAXAL, (ESOPHAGUS, STOMACH, CECUM, AND INTESTINES. By S. 0. HABERSHOX, M. D., Assistant Physician to and Lecturer on Materia Medica and Tiieraiieutics at Guy's Hospital. WITH HANDSOME ILLUSTRATIONS ON WOOD. |^° B}^ reference to the Terms of the "Ame- rican Journal of the Medical Sciences," pp. 3 and 4, it will be seen that advance-pay- ing subscribers obtain this valuable practical work without charg'e. He takes us well through all the diseases, structural and functional (as they are called), of the alimentary canal. He begins at the pharynx, and carefully and honestly follows the tube through all its constrictions, dilatations, divarications, and appendices. He tells us what misfortunes happen at every part of it: he de- scribes the particular seat of the disease at each con- ventional division of the apparatus; their anatomical characters (when they have any); their general patho- logy: the symptoms they excite; the conditions of the system with which they are coincident; the particular general disorder with which they are associated : and he also tells us what aids medicine offers man to help him through these numerous dangers and difficulties; and how those aids are to be wisely used to their ends. The practitioner will find it a valuable work of refer- ence. — London Med. Times and Gaz.. Nov. 7, 1857. Dr. Habershon abstains, as a rule, from speculations, and confines himself mainly to the record of facts re- lating to symptoms, morbid chanses, and treatment. His remarks are illustrated by the histories of one I THE PATHOLOGY AND TREATMENT OF PULMOMRT TUBERCULOSIS, AND ON THE LOCAL MEDICATION OF PHARYNGEAL AND LARYNGEAL DISEASES FREQUENTLY MIS- TAKEN FOR OR ASSOCIATED WITH PHTHISIS. Chronic Ulcer of Stomach. hundred and sixty-three cases, recorded in the ease- books of Guy's Hospital, descriptive of the numerous forms of disea.«e of the alimentary canal. The book is therefore essentially practical; and on this ground, and bearing evidence of being the work of a careful observer, it forms an addition of value to the already existing literature of Diseases of the Alimentary Canal. British Med. Journal, Nov. 21, 1857. Amongst the valuable treatises we have pointed out there yet existed room for a work which should deal but little with theory and scientific analyses, but con- fine itself to the record of direct bedside and post- mortem, room results. Dr. Habershon"s treatise sup- plies this deficiency. We recommend Dr. Habershon's treatise as a valuable repertory of clinical experience, of very trustworthy character. — London Lancet, Nov. 21, 1857. We believe that this book will be read with interest. It is calculated to impart instruction to all of us. — Brit, and For. Medico-Chirurg . Meview, April, 1858. J. HUGHES BENNETT, M. D., F.R.S.E., Professor of Clinical Medicine in the ITniversily of Edinburgh, &e. Ill one small octavo volume, extra cloth, with beau- tiful illustrations. $1 '25. Tubercular Cavity in the act of Healing. MEDICAL AND SCIENTIFIC PUBLICATIONS. 43 RICORD AND HUNTER ON VENEREAL— New Edition, Now Ready (1869). A TREATISE 01 THE"yENEREAL DISEASE. BY JOHN HUNTER, F. K S., &c. With Copious Additions by PH. RICORD, M. D. Second Edition, containing a resume of Ricord's recent Lectures on Chancre. Translated and Edited, witu Notes, by FREEMAN J. BUMSTEAD, M.D., Lecturer on Venereal at the College of Physicians and Surgeons, New York. Ill 07ie large and handsome octavo volume of 550 pages, with eight plates ; extra cloth, $3 25. "M. Ricord's annotations to Hunters Treatise on the Venereal Disease were first published at Paris, in 1840, in connection with Dr. G. Richelot's translation of the work, including '.he contributions of Sir Everard Home and Mr. Babington. In a second edition, which has recently appeared, M. Ricord has thoroughly revised his part of the work, bringing it up to the knowledge of the present day, and so materially increasing it that it now constitutes full one-third of the voltirae. This publication has been received with great favor by the French, both because it has placed within their reach an important work of Hunter, and also because it is the only recent practical work which M. Ricord has published, no edition of his Traite des Maladies Veneri- ennes having appeared for the last fifteen years." — Translator'' s Preface. The addition, in the present edition, of the material contained in the "Lectures on Chancre," published a few months since by M. Ricord, renders this work the most complete embodiment of the views of the great French syphilographer that has 'vet been given to the profession, while the editor has further endeavored to present all other matter of interest that has appeared since the publication of the first edition. The volume may therefore be regarded as a complete work of reference on the subject in which the practitioner may at all times be certain of finding an elucidation of doubtful questions either of theory or practice. Every one will recognize the attractiveness and value which this work derives from thus presenting the opinions of these two masters side by side. But, it must be admitted, what has made the fortune of the book, is the fact that it contains the " most com- plete embodiment of the veritable doctrines of the Hopital du Midi," which has ever been made public. The doctrinal ideas of M. Ricord, ideas which, if not universally adopted, are incontestably dominant, have heretofore only been interpreted by more or less skilful secretaries, sometimes accredited and sometimes not. Tn the notes to Hunter, the master substitutes himself for his interpreters, and gives his original thoughts to the world, in a summary form it is true, but in a lucid and perfectly intelligible manner. In conclusion, we can say that this is incontestably the best treatise on syphilis with which we are acquainted, and, as we do not often employ the phrase, we may be excused for expressing the hope that it may find a place in the library of every physician. — Va. Med. and Surg. Journ. Also — HUNTER'S COMPLETE WORKS. numerous Illustrations; leather, $10 00. In four octavo volumes, with RICORD'S LETTERS ON SYPHILIS, Addressed to the Chief Editor of the Union Medicale. With an Introduction by Amedee Latour. Translated by W. P. Lattimore, M. D. In one neat octavo volume, extra cloth, $2. Lately Published. THE MODERN TRE/VTMENFoF SYPHILITIC DISEASES, BOTH PRIMARY AND SECONDARY. COMPRISING THE Treatment of Constitutional and confirmed Syphilis by a safe and successful Method. WITH NUMEROUS CASES, FORMULA, AND CLINICAL OBSERVATIONS. BY LANGSTON PARKER, Surgeon to the Queen's Hospital, Birmingham. Erom the Third and entirely rewritten London Edition. Ill one neat octavo vohime of over 3QQ pages, extra cloth, $1 75. The third edition of Mr. Parker's work constitutes, I has contrived to emhody in it everything of import- we must say, an excellent practical treatise, the suh- auce respecting syphilis and its treatment. — Dublin ject is remarkably well arranged; indeed, the author | Medical Press. LECTURES ON THE PRINCIPLES AND METHODS OF MEDICAL OB- SERVATION AND RESEARCH. For the use of Advanced Students and Junior Practi- tioners. By Thomas Laycock, M. D., F. R. S. E., Professor of Practical and Clinical Medi- cine in the University of Edinburgh, etc. In one very neat royal 12mo. vol., extra cloth, $1. 44 BLANCHARD AND LEA'S NEW AND MUCH IMPROVED EDITION— (Just Issued.) THE HISTORY, DIAGNOSIS, AND TREATMENT OP THE FEYERS OF THE UNITED STATES. BY ELISHA BARTLETT, M. D., Late Professor of Materia Medica, &c., in the College of Physicians and Surgeons, New York. J^ourtlj ibttion, ^Itbxscb BY ALONZO CLARK, M. D., Professor of Pathology and Practical Medicine in the College of Physicians and Surgeons, New York. In one large and handsome octavo volume, of over six hundred pages ; extra cloth, $3 00. From the Editor'' s Preface. " The question maybe fairly raised whether any book in our profession illustrates more clearly the beauties of sound reasoning, and the advantages of vigorous generalization from carefully selected facts. Certainly no author ever brought to his labor a more high-minded purpose of representing the truth, in its simplicity and m its fulness, w^hile (ew have been possessed of higher gifts to discern, and gracefully to exhibit it. Had I been prepared by previous reading for the duty which the partiality of Dr. Bartlett assigned to me, of preparing this edition for the press, the labor would have been inconsiderable. As it is, I have read extensively, to learn how little that the book contains can be advantageously altered. Considerable matter has been added, it is true, because new facts have been observed, and new opinions have been ex- pressed, which both add to our knowledge, and suggest new topics for investigation. This 1 have endeavored to select, and so far as it is original to write, with the same re>pect for truth, and desire for usefulness, which influenced the mind of my endeared friend, the accomplished author." Carefully revised by Professor Clark, who has introduced whatever is new in the literature of this branch of medical science since the appearance of the last edition, embodying the results of the observations and researches of Drake, La Roche, Flint, Barton, Dickson, Fenner, Peacocke, and others, this volume will be found fully brought up to the present day, and emi- nently worthy a continuance of the confidence of the profession. The masterly and elegant treatise by Dr. Bartlett is invaluable to the American student and practitioner. — Dr. Holmes's Report to the Nat. Med. Association. We regard it, from the examination we have made of it, the best work on fevers extant in our language, and as such cordially recommend it to the medical public. — St. Louis Medical and Surgical Journal. Take it altogether, it is the most complete history of our fevers which has yet been published, and every practitioner should avail himself of its contents. — The Western Lancet. Of the value and importance of such a work, it is needless here to speak; the profession of the United States owe much to the author for the very able volume which he has presented to them, and for the careful and judicious manner in which he has executed his task. No one volume with which we are acquainted contains so complete a history of our fevers as this. To Dr. Bartlett we owe our best thanks for the very able volume he has given us, as embodying certainly the most complete, methodical, and satisfactory ac- count of our fevers anywhere to be met with. — The Charleston Med. Journal and Review. THE HTJ]M:A.N BHA-IN; WITH A DESCRIPTION OF THE TYPICAL FORMS OF BRAIN IN THE ANIMAL KINGDOM. BY SAMUEL SOLLY, F. R. S., Senior Assistant Surgeon to St. Thomas's Hospital, &c. jFrom i\t snav.'ii I/onlion jcIJittDit. WITH ONE HUNDRED AND EIGHTEEN ELABORATE ILLUSTRATIONS ON WOOD. In one handsome octavo volume, of about five hundred pages, extra cloth, %2 00. Inferior Surfaceofthe Cerebellum. MEDICAL AND SCIENTIFIC PUBLICATIONS. 45 Just Issued. ATLAS or CUTANEOUS DISEASES. BY J. MOORE NELIGAN, M. D., M. R. I. A., &c. Willi Splendid Colored Plates, presenting nearly one hundred elaborate representations of Disease, colored after nature. Ill one very handsome quarto volume., extra doth., price $4 50. Also now ready, by the same Author. A PRACTICAL TREATISE ON DISEASES OF THE SKIN. SECOND AMERICAN EDITION. In one neat royal 12mo. volume, extra cloth, of 334 pages, price $1. These two volumes, constituting- together a complete work on the diagnos^is. pathology, and treatment of cutaneous affections, will be forwarded by mail on receipt of $5. A compend which will very much aid the practi- tioner in this difficult branch of diagnosis. Taken with the beautiful plates of the Atlas, which are re- markable for their accuracy and beauty of coloring, it constitutes a very valuable addition to the library of a practical man. — Buffalo Med. Journal. The lithographs are so colored as to be true and faithful representations of these ninety varieties of a class of diseases whose exact diagnosis is thus made plain and easy, and which, in the absence of such aid or a long and attentive study, is so difficult that very few praotitioners seriously attempt it. The work is cheap, and no practitioner ambitious of a high profes- sional status can afford to dispense with such helps. — JVashviUe Journal of Medicine. Neligan's Atlas of Cutaneous Diseases supplies a long existent desideratum much felt by the largest class of our profession. It presents, in quarto size, 16 plates, each containing from 3 to 6 figures, and form- ing in all a total of 90 distinct representations of the different species of skin affections, grouped together in genera or families. The illustrations have been taken from nature, and have been copied with such fidelity that they present a striking picture of life; in which the reduced scale aptly serves to give, at a coup d'ceil, the remarkablepeculiaritiesofeach individual variety. And while thus the disease is rendered more definable, there is yet no loss of proportion incurred by the ne- cessary concentration. Each figure is highly colored, and so truthful has the artist been that the most fas- tidious observer could not justly take exception to the correctness of the execution of the pictures under his scrutiny. — Montreal Med. Chronicle. Ne-w and enlarged Edition — No^w Ready (June, 1859). UEINARYT3EP0SITS; THEIR DIAGNOSIS, PATHOLOGY, AND THERAPEUTICAL INDICATIONS. BY GOLDING BIRD, M. D., F. R. S., &c. Edited bt EDMUND LLOYD BIRKETT, M. D., &c. A New American, from the Fifth London Edition. WITH EIGHTY ILLUSTRATIONS. In one handsoTne octavo volume, extra cloth, of 382 pages, $2 00. The death of Dr. Bird has rendered it necessary to entrust the revision of (he present edition to other hands, and in his performance of the duty thus devolving on him, Dr. Birkett has sedu- lou^ly endeavored to carry out the author's plan by introducing such new matter and modifica- tions of the text as the progress of science has called for. Notwithstanding the utmost care to keep the work within a reasonable compass, these additions have resulted in a considerable enlargement. It is, therefore, hoped that it will be found fulJy up to the present condition of the subject, and that the reputation of the volume as a clear, complete, and compendious manual, will be fully maintained. It can scarcely be necessary for us to say anything of the merits of this well-known Treatise, which so ad- mirably brings into practical application the results of those microscopical and chemical researches regard- ing the physiology and pathology of the urinary se- cretion, which have contributed so much to the increase of our diagnostic powers, and to the extension and satisfactory employment of our therapeutic re- sources. In the preparation of this new edition of his work, it is obvious that Dr. Golding Bird has spared no pains to render it a faithful representation of the present state of scientific knowledge on the subject it embraces. — British and Foreign Med.-Chir. Review. Crystals of Phosphate of Soda. MANUALS ON THE BLOOD AND URINE.— By John | FRICK ON THE URINE.— Renal Affections, their William Griffith, M. D,, G. Owen Reese, M. D., and | Diagnosis and Pathology. In one handsome volume, Alfred Markwick. In one large 12mo. volume, extra I royal 12mo., with illustrations, 75 cents, cloth, of 460 pages, with Plates, $1 25. I 46 BLANCHARD AND LEA'S The Standard work on the Skin— (Just Issued.) ON i:>isea-Se:s~of the skidst. BY ERASMUS WILSON, F. R. S., Author of "A System of Human Anatomy," &c. S;ije Joitrtlj anb ^nlargeb '^mmcmx, from tljc last anb |mpra&cb ITonbon (Bbition. In, one large octavo vohime, of 650 pages, extra cloth, $2 75. This volume, in passing for tlie fourth time through the hands of the author, has received a careful revision, and has been greatly enlarged and improved. About one hundred and fifty pages have been added, including new chapters on Classification, on General Pathology, on General Therapeutics, on Furuncular Eruptions, and on Diseases of the Nails, besides extensive •additions throughout the text, wherever they have seemed desirable, either from former omis- sions or from the progress of science and the increased experience of the author. Appended to the volume will also now be found a collection of Selected FoemuLjE, consisting for the most part of prescriptions of which the author has tested the value. Thus perfected and brought up to the latest moment, this work cannot fail to maintain its character as the standard authority on this important and perplexing class of affections. When the first edition of this work appeared, about fourteen years ago, Mr. Erasmus Wilson had already given some years to the study of Diseases of the Skin, and he then expressed his intention of devoting his future life to the elucidation of this particular branch of Medical Science. In the present edition Mr. Wil- son presents us with the results of his matured ex- perience gained after an extensive acquaintance with the pathology and treatment of cutaneous affections ; and we have now before us not merely a reprint of his former publications, but an entirely new and re- written volume. Thus, the whole history of the dis- eases affecting the skin, whether they originate in that structure or are the mere manifestations of de- rangement of internal organs, is brought under notice, and the book includes a mass of information which is spread over a great part of the domain of Medical and Surgical Pathology We can safely recommend it to the profession as the best work on the subject now in existence in the English language. — London Med. Times and Gazette, March 28, 1857. The " Diseases of the Skin," by Mr. Erasmus Wilson, may now be regarded as the standard work in tliat department of medical literature. — Medico-Chirurg . Review. Also, Now Ready. A SERIES OF PLATES IllUSTRATING "WILSON ON DISEASES OF THE SKIN." CONSISTING OF NINETEEN BEAUTIFULLY EXECUTED PLATES, OF WHICH TWELVE AEE EXQUISITELY COLORED. Presenting the Normal Anatomy and Pathology of the S/dn, and embracing accurate repre- sentations of about one hundred varieties of Disease, m,ost of them, the size of Nature. Price, in extra cloth, $4 25. For beauty of drawing, and accuracy and finish of colorinff, these plates are confidently pre- sented as superior to anything of the kind as yet issued in this country. The plates by which this edition is accompanied leave nothing to be desired, so far as excellence of delineation and perfect accuracy of illustration are concerned. — Medicc-Chirurgical Review. Of these plates it is impossible to speak too highly. The representations of the various forms of cutaneous disease are singularly accurate, and the coloring ex- ceeds almost anything we have met with in point of delicacy and finish." — British and Foreign Medical Review. Epidermal horn, caused by Disease of the Oil Glands. By the same Author— (Lately Issued.) HEALTHY SKIN: A Popular Treatise on the Skin and Hair, their Preservation and Management. SECOND AMERICAN, PROM THE POUKTH AND REVISED LONDON EDITION. J>i o?ie neat royal 12mo. volum,e, of about three htm- dred pages, with numerous Illustrations ; extra cloth, or flexible cloth, $1 00 ; paper covers, 75 ce7it . The student will be delighted to find his labors so much facilitated; and a few hours of agreeable society with a moat pleasantly-written book will do more to make him acquainted with a class of obscure diseases than all that has been pre- viously written on the subject. — London Lancet. By the same Author. ON CONSTITUTIONAL AND HEREDITARY SYPHILIS, AND ON SYPHILITIC EHUPTIONS. In one beautifully printed octavo volume, with four exquisite colored plates, presenting more than thirty varieties of Syphilitic Eruptions ; extra cloth, $2 25. MEDICAL AND SCIENTIFIC PUBLICATIONS. 47 New and inipioved Edition — Now Ready (1859), A PRACTICAL TREATISE ON THE DISEASES OF CHILDREN. BY D. FRANCIS CONDIE, M. D. FIFTH EDITION, REVISED AND ENLARGED. In one large octavo volume of nearly eight hundred pages ; leather, $3 25. From the Author's Preface. To present a complete and faithful exposition of the pathology and therapeutics of the mala- dies incident to the earlier stages of existence — a full and exact account of the diseases of infancy and childhood — has been the aim of the author of the present treatise. For the furtherance of this object, in the preparation of a fifth edition, the entire work has been subjected to a careful and thorough revision; a considerable portion of it has been entirely rewritten, and several new chapters have been added. In the different sections will be found incorporated every important observation in reference to the diseases of which they treat, that lias been recorded since the appearance of the last edi- tion ; and in the several new chapters, an account of some affections omitted in former editions, and for the accurate description and satisfactory management of which we are indebted mainly to the labors of recent observers. Tlie value of works by native authors on the diseases which the physician is called upon to combat, will be appreciated by all ; and the work of Dr. Condie has gained for itself the character of a safe guide for stu- dents, and a \iseful work for consultation by those en- gaged in practice. — Nnu York Medical Times. In the department of infantile therapeutics, the work of Dr. Condie is considered one of the best which has been published in the English language. — The Stetho- scope. As we said before, we do not know of a better book on Diseases of Children, and to a large part of its re- commendations we yield an unhesitating concurrence. — Buffalo Medical Journal. The work of Dr. Condie is unquestionably a very able one. It is practical in its character, as its title imports; but the practical precepts recommended in it are based, as all practice should be, upon a familiar knowledge of disease. The opportunities of Dr. Con- die for the practical study of the diseases of children have been great, and his work is a proof that they have not been thrown away. He has read much, but ob- served more; and we think that we may safely say that the American student cannot tind, in his own language, a better book upon the subject of which it treats. — Am. Journal Medical Sciences. To call the attention of the medical fraternity to a new edition of this American medical classic, seems almost a work of supererogation. The name of Dr. Condie is too well known, and his influence too gene- rally felt and acknowledged in the department of Pa- thology to which the present work refers, to require other than a very brief notice of this new edition. — The Peninsxdar Journal of Medicine. Taken as a whole, in our judgment. Dr. Condie's Treatise is one from the perusal of which the practi- tioner in this country will rise with the greatest sa- tisfaction. — Western Journal of Medicitie and Surgery. One of the best works upon the Diseases of Children in the English language. — Western Lancet. We feel assured from actual experience that no phy- sician's library can be complete without a copy of this work. — iV". T. Jounial of Medicine. Perhaps the most full and complete work now before the profession of the United States; indeed, we may say in the English language. It is vastly superior to most of its predecessors. — Transylvania Med. Journal. A veritable ptediatric encyclopedia, and an honor to American medical literature. — Ohio Medical and Surgical Journal. The author's great experience, extensive reading, and familiarity with foreign languages, peculiarly fit him for the execution of such a work ; and its value is abundantly proved by the numerous editions which it has reached in a very short period. It is character- ized by great clearness and lucidity of style and ar- rangement, sound pathological views, and precise and plain therapeutical directions. — Medical Examiner. To the American practitioner. Dr. Condie's remarks on the di.^eases of children will be invaluable, and we accordingly advise those who have failed to read this work to procure a copy, and make themselves familiar with its sound principles. — The New Orleans Medical and Surgical Journal. LECTUKES ON THE DISEASES OF INFANCY AND CHILDHOOD. BY CHARLES WEST, M. D., Physician to the Hospital for Sick Children, &c. Second American, from the Second and Enlarged London Edition. In one neat octavo volume of nearly 500 large pages; extra cloth, $2. In taking leave of Dr. West, we can scarcely do more than reiterate our former praise of him. We have given, we fear, but a very faint notion of the scope of his work, and of its excellent execution. It is one standing by itself upon its important subject in our language — unapproached, unrivalled His knowledge of what others have done is equalled only by his own extensive experience ; and the results of both are com- bined in his valuable practical lectures now offered for the guidance of others. — Brit, and For. Med.-Ckirurg. Jteview. The book has about it that practical common sense character which is always acceptable to the practition- er of medicine, whilst the immense experience of Dr. West, derived from his connection with the London Hospital for Sick Children, gives to him opportunities for the minute observation of the diseases incident to childhood, such as no private practice can offer. We would especially recommend the careful study of these lectures to the medical student who is preparing him- self for general practice. — Va Med. and Surg. Journal. 48 BLANCHARD AND LEA'S New and Enlarged Edition — (Just Issued.) ON THE DISEASES OF INFANTS AND CHILDREN. BY FLEETWOOD CHURCHILL, M. D., M. R. I. A., Author of " Theory and Practice of Midwifery," '• Diseases of Females," &e. &c. Second American Edition, Enlarged and Revised by the Anthor. Edited, ttith Notes, by W. V. KEATING, M. D. In one large and handsome octavo volume of over 100 pages ; leather, $3 25; extra doth, $3. The thorough manner in which Dr. Churchill has revised this work may be estimated from the fact that not only has a considerable portion been rewritten, but that in all the present edition contains about one-third more matter than the former. Besides an enlargement in the size of the page, the volume has been increased by about seventy-five pages, notwithstanding which the price has been maintained at the former very moderate rate. Few -works devoted to the same subject will be comes to us with its value greatly enhanced, not found, in fact, to excel the one before us, in extent of merely by bein<; carefully revi.-ed throughout, but research, copiousness of reference, and fulness and also by the addition of several entire chapters ; on accuracy of detail. It will constitute a valuable and Atelectasis Pulmonum, Pulmonary Phthisis, Tabes reliable guide to the knowledge of the several morbid Mesenterica. (Edema of the Cellular Tissue, Typhoid conditions incident to infancy and childhood, their Fever, and Infantile Syphilis. The volume is a most causes, semeiology, seats, character, and progress, and valuable addition to this department of medical sci- the means best adapted for their alleviation and cure, ence. — Boston Med. and Surg. Jowtial, Sept. 1856. equally suitable to the student, as to the practitioner . . •. , . •. who would render himself familiar with the lights It is with pleasure we notice the appearance of the which recent investigation and discoveries have second edition of this excellent work on the diseases thrown upon each and all of these particulars.— J»i. , of Children. It has been carefully revised by the au- Journ. Med. Sciences. Oct. 1856. thor. and all the information added which has been de- I rived from recent researches. Every paragraph, says Dr. Churchill is well known as the author of various ' the author, has been carefully gone over with a view works on obstetrics and the diseases of women. Like to correction, and suggestions made by reviewers have them, this contribution to infantile therapeutics is been carefully weighed, and adopted when correct, especially useful for the careful and extensive collec- ^ Several entire new chapters, and portions of chapters, tion of material bearing upon the subject. As a book ' have been added, and especial reference has been had of reference, it will be often sought for, whilst its emi- to American authorities so as to adapt it to the wants nently practical character will make it an admirable of this country. We feel that it is no more than jus- manual for the student. We are compelled for want tice to add our recommendation, most cordially, of this of space, to refrain from making as many extracts as work, as being among the best we have on this sub- we would like to do, and will close our notice of the I ject. — Cincinno.ti Medical Observer, Oct. 1856. work by heartilv recommending it both to the student I „, , , . ^ ,, , , ^ -,, ^ ^ , and the practitioner.— Fu. Med. Journal, Sept. 1856. ^he book is fully posted up and will be found a va- luable addition to the practitioner s library. — Southern This new edition of the work of a favorite author | Med. and Surg. Journal, Nov. 1856. A TREATISE ON THE PHYSICAL AXD MEDICAL TPiEAT3IENT OF CHEDREN. BY WILLIAM P. DEWEES, M. D. Tenth Edition. In one octavo volume of 548 pages ; extra cloth, $2 80. Tenth Edition. A TREATISE ON THE DISEASES OF FEMALES. BY WILLIAM P. DEWEES, M. D. hi one octavo volume of five hundred and thirty-two pages, vAth plates, S'i 00. Just Issued. ON THE CONSTITUTIONAL TREATMENT OF FEM:A.LE I3ISEA.SES. BY EDWARD RIGBY, M. D., Senior Physician to the General Lying-in Hospital, &c.; Author of "A System of Midwifery," &c. In one neat royal 12mo. volume of 250 pages; extra cloth, $1. MEDICAL AND SCIENTIFIC PUBLICATIONS. 49 New axid Improved Edition— Now Ready (June, 1859). ¥OMA¥; HER DISEASES AND REMEDIES. A SERIES OF LETTERS TO HIS CLASS. BY CHARLES D. MEIGS, M. D., Professor of Midwifery, and Diseases of Women and Cliildren, in the Jefferson Medical College, Philadelphia. FOURTH EDITION, REVISED. hi one large and liandsome octavo volume of over seven hundred fages ; leather^ $3 60. The gratifying appreciation of his labors, as evinced by the exhaustion of three large impres- sions of this work, has not been lost upon the author, wtio has endeavored in everj' way to ren- der it worthy of the favor with which it has been received. The opportunity thus afforded for another revision has been improved, and the work is now presented as in every way superior to its predecessors, additions and alterations having been made wherever the advance of science has rendered them desirable. The typographical execution of the work will also be found to have undergone a similar improvement, and the volume, it is hoped, will be found in all respects worthy to maintain the position it has acquired as the standard American text-book on the Dis- eases oi Females. A few notices of the previous editions are appended. On its first appearance we had occasion to speak of it in terms of merited commendation. In announcing the third edition, it is only necessary to say that it has been thoroughly revised and much enlarged by its distinguished author so as greatly to enhance its value and usefulness. — St. Louis Med. and Surg. Journ. Keplete with sound practical views, and evidently the production of a man of vast experience and tho- roughly conversant with the subject. — Medical Chro- nicle. He is a bold thinker, and possesses more originality of thought and style than almost any American writer on medical subjects. If he is not an elegant writer, there is at least a freshness — a raciness in his mode of expressing himself— that cannot fail to draw the reader after him even to the close of his work ; j'ou cannot nod over his pages; he stimulates rather than narcotises your senses, and the reader cannot lay aside these letters when once he enters into their merits. This the second edition is much amended and enlarged, and affords abundant evidence of the author's talents and industrj'. — N. 0. Medical and Surgical Journal. The practical writings of Dr. Meigs are second to none. — Ttie N. T. Journal of Medicine. The excellent practical directions contained in this volume, give it great utility, which we trust will not be lost upon our older colleagues: with some conden- sation, indeed, we should tliink it well adapted for translation into German. — Zeitschriftfur die Gesammte Medecin. The merits of the first edition of this work were so generally appreciated, and with such a high degree of favor by the medical profession throughout the Union, that we are not surprised in seeing a second edition of it. It is a standard work on the diseases of females, and in many respects is one of the very best of its kind in the English language. Upon the appearance of the first edition, we gave the work a cordial recep- tion, and spoke of it in the warmest terms of commen- dation. Time has not changed the favorable estimate we placed upon it. but has rather increased our con- victions of its superlative merits. But we do not now deem it necessary to say more than to commend this work, on the diseases of women, and the remedies for them, to the attention of those practitioners who have not supplied themselves with it. The most select library would be imperfect without it. — The Western Journal of Medicine and Surgery. There is an off-hand fervor, a glow and a warm- heartedness infecting the effort of Dr. Meigs, which is entirely captivating, and which absolutely hurries the reader through from beginning to end. Besides, the book teems with solid instruction, and it shows the very highest evidence of ability, viz., the clearness with which the information is presented. We know of no better test of one's understanding a subject than the evidence of the power of lucidly explaining it. The most elementary, as well as the obscurest subjects, under the pencil of Prof. Meigs, are isolated and made to stand out in such bold relief, as to produce distinct impressions upon the mind and memory of the reader. — The Charleston Medical Journal. That he has succeeded in writing a readable work, in our estimation, we have the best practical evidence in the fact, that, in spite of a deliberate resolve to postpone its perusal, having other more pressing matters on hand, we found ourselves taking it up day after day, at our leisure moments, until the volume was finished. If we may judge of others by our own experience, the student and practitioner will find it an attractive book. The character of the work is infi- nitely more pleasing to us than if written in a tame, spiritless manner, albeit, in the latter case, it were less obnoxious to criticism. We like to see the man in the book. It shows the author to be a scholar, a man of genius, and as regards matter, when it is considered that Prof M. ranks among the most talented and learned of the profession of this country, and, we may safely add, of any country, and, moreover, that his experience in his particular province has been im- mense, it can hardly appear otherwise than that what he may write on the diseases of females shall be de- serving of most respectful attention. — Buffalo Medical Journal. By the same Author. A TREATISE ON ACUTeTnD CHROKIC DISEASES OF THE NECK OF THE UTERUS. With numerous Plates, drawn and colored from Nature in the highest style of art. In one very handsorae octavo volume ; extra doth., $4 50. A TREATISE ON THE DISEASES AND SPECIAL HYGIENE OE EEMALES. By CoLOMBAT De L'Isere. Translated from the French, with additions, by Charles D. Meigs, M. D., Professor of Midwifery, and Diseases of Women and Children, in the Jefferson Medi- cal College, Philadelphia. A new and revised edition. In one octavo volume of over 700 pages, with numerous illustrations ; extra cloth, $3 50. 4 50 BLANCHARD AND I.EA'S Enlarged and Illustrated Edition— Just Issued. VERY HANDSOME I OCTAVO VOLUME, OF OVER SEVEN HUNDRED AND FIFTY LARGE PAGES. WITH Numerous Illustrations. LEATHER, Price $3 00. Prolapsus Vkri. ON THE DISEASES OF WOMEN, mCLUDI^^G DISEASES OF PREGNA^^CY AND CHILDBED. BY FLEETWOOD CHURCHILL, M. D., M. R. L A., Author of "Diseases of Infants and Children," "Theory and Practice of Midwifery," &c &c. Sixth American Edition, extensively Enlarged and Revised by the Author. W.iX\ NotfS anlJ gLtilJitions BY D. FRANCIS CONDIE, M. D , Author of "A Practical Treatise on Diseases of Children," &c. On no former edition has the author devoted more labor to render the work a complete and satisfactory resume of the existing state of knowledge on the important questions under con- sideration. To effect this, extensive additions have been required, some additional chapters inserted, and many portions rewritten A handsome series of original illustrations of the more important pathological conditions has been prepared, while the mechanical execution of the volume will be found greatly superior to any former edition. Former editions of this work have been noticed in previous numbers of the Journal. The sentiments of high commendation expressed in those notices, have only to be repeated in this : not from the fact that the profession at large are not aware of the high merits which this work really possesses, but from a desire to see the principles and doctrines therein contained more generally recognized, and more universally car- ried out in practice. — N. Y. Journal of Medicine. We know of no author who deserves that approba- tion, on " the diseases of females," to the same extent that Dr. Oh archill does. His, indeed, is the only tho- rough treatise we know of on the subject; and it may be commended to practitioners and students as a mas- terpiece in its particular department. The former editions of this work have been commended strongly in this journal, and they have won their way to an extended, and a well-deserved popularity. This fifth edition, before us, is well calculated to maintain Dr. Churchill's high reputation. It was revised and en- larged by the author, for his American publishers, and it seems to us that there is scarcely any species of desirable information on its subjects that may not be found in this work. — The We^te,rn Journal of Medicine and Surgery. "We are gratified to announce a new and revised edi- tion of Dr. Churchill's valuable work on the diseases of females. We have ever regarded it as one of the very best works on the subjects embraced within its scope, in the English language; and the present edi- tion, enlarged and revised by the author, renders it still more entitled to the confidence of the profession. — The Western Lo.ncet. As a comprehensive manual for students, or a work of reference for practitioners, we only speak with com- mon justice when we say that it surpasses any other that has ever issued on the same subject from the British press. — Ttie Dublin Quarterly Journal. By the same Author. ESSAYS ON THE PUERPERAL FEVER, AND OTHER DISEASES PECULIAR TO WOMEN. Selected from the writings of Briti-h Authors previous to the close of the Eight- eenth Century. In one neat octavo volume, of about 450 pages, extra cloth, $2 50. MEDICAL AND SCIENTIFIC PUBLICATIONS. 51 MEIGS ON PUERPERAL FEVER— (Lately Published.) ON THE NATURE, SIGNS, AND TREATMENT OF CHILDBED FEVERS: IN A SERIES OF LETTERS ADDRESSED TO THE STUDENTS OF HIS CLASS. BY CHARLES D. MEIGS, M. D., Professor of Midwifery and Diseases of Women and Children in the Jefferson Medical College, Philadelphia. hi 07ie very liandsome octavo volume of 362 pages ; extra cloth, $2 50. The author, a practitioner of more than forty years, has collected the best and most reliable opinions of many writers on this disease, and applying to them the test of clinical experience, has produced a volume superior in many respects to any which has heretofore come from his pen As the results of a critical inquiry into the literature of the disease and a large and long experience in practice, this volume will be received by the profession with distinguished favor, and we feel warranted in commending it to the favorable attention of our readers. — N. Y. Journal of Medicine. The present is upon one of the most diflBcult subjects in obstetrics, and we feel bound to say that as a whole it is superior to any other work upon the same subject. — Edinburgh Med. Journal. We cordially recommend it to our readers as a work abounding in practical good sense, and sound patho- logical and therapeutical views. — Si. Louis Med. and Surg. Journal. The instructive and interesting author of this work, whose previous labors in the department of medicine which he so sedulously cultivates, have placed his countrymen under deep and abiding obligations, again challenges their admiration in the fresh and vigorous, attractive and racy pages before us. It is a delectable book. * * * This treatise upon childbed fevers will have an extensive sale, being destined, as it de- serves, to find a place in the library of every practi- tioner who scorns to lag in the rear of his brethren. — Nashville Journal of Medicine and Surgery. Just Issued. ON SOME DISEASES OF WOMEN ADMITTING OF SURGICAL TREATMENT. BY ISAAC BAKER BROWN, Surgeon Accoucheur to St. Mary's Hospital, &c. WITH HANDSOME ILLUSTRATIONS. In 07ie neat octavo volume, extra cloth, of two hundred and seventy-six pages, $1 60. During the appearance of this work in the "Medical News and Li- brary" for 1855 and 1856, it commanded the warmest approbation of the profession. To the practitioner the work possesses peculiar import- ance as treating- fully and completely of a class of accidents and diseases of frequent occurrence, which have hitherto received but little attention from writers of systematic works. Occupying a middle ground be- tween surgery and obstetrics, the authorities on each subject have well- nigh abandoned them to the consideration of the other, and thus they are nowhere to be found treated with the attention to which their import- ance is entitled. The subjects considered in extenso are Ruptured Pe- rineum—Prolapse of the Vagina — Prolapse of the Uterus — Vesico-Vagi- nal Fistula — Recto- Vaginal Fistula — Lacerated Vagina — Polypus of the Uterus — Stone in the Female Bladder — Vascular Tumor of the Meatus Urinarius — Imperforate Hymen — Encysted Tumor of the Labia — Dis- eases of the Rectum arising from the Uterus — Ovarian Dropsy. Needle for Ruptured Peri7ieum . A PRACTICAL TREATISE ON THE DISEASES PECULIAR TO WOMEN. Illustrated by Cases derived from Hospital and Private Practice. By Samuel Ashwell, M- D., Obstetric Physician and Lecturer to Guy's Hospital, London. Third American, from the Ihird and Revised London edition. In one octavo volume of 528 pages; extra cloth, $3. The most useful practical work on the subject in the English language. — Boston Med. and Surg. Journal. The most able, and certainly the most standard and practical, work on female diseases that we have yet seen. — Medico-Chirurgical Review. The young practitioner will find it invaluable, while those who have had most experience will yet find something to learn, and much to commend, in a book which shows so much patient observation, practical skill, and sound sense. — British and Foreign Med. Re- view. We commend it to our readers as the best practical treatise on the subject which has yet appeared. — Lon- doisoisrs, IN RELATION TO MEDICAL JURISPRUDENCE AND MEDICINE. Second American, from the Second and Revised English Edition. In one large octavo volume of Ibb pages, leather, $3 50. The length of time which has elapsed since the first appearance of this work has wrought so great a change in the subject as to require a very thorough revision to adapt the volume to the present wants of the profession. The rapid advance of Chemistry has introduced into use many new substances which may become fatal through accident, carelessness, or design — while at the same time it has likewise designated new and more exact modes of counteraciing or detecting those previously treated of. Mr. Taylor's position as the leading medical jurist of England, has during this period conferred on him extraordinary advantages in acquiring experience in all that relates to this department, nearly all cases of moment being referred to him for examination, as an expert whose testimony is generally accepted as final. The results of his labors, therefore, as gathered together in this volume, carefully weighed and sified, and presented in the clear and intelligible style for which he is noted, may be received as an acknowledged authority, and as a guide to be followed with implicit confidence. In his Preface the author says : — " My space has been limited, and I have endeavored to fill it with materials which may be of profit to the practitioners of law and medicine, for whose especial use this volume is intended. Under this view the plan of the former edition has been entirely changed. Many chapters have been struck out, and an equal number of new chapters introduced. The requirements of a pe riod dating no longer ago than ten years are different from those of the present day, and it is the duty of an author, so far as it may be in his power and consistent with the scope of his labors, to fulfil these requirements by an entire remodelling of his subject." The most elaborate work on the subject that our literature possesses. — British and Foreign Medico-Chi- rurgical Review. It contains a vast body of facts, which embrace all that is important in toxicology, all that is necessary to the guidance of the medical jurist, and all that can be desired by the lawyer. — Medico-Chirurg. Review. One of the most practical and trustworthy works on Poisons in our language. — Western Jaum. of Med. It is, so far as our knowledge extends, incomparably the best upon the subject; in the highest degree cre- ditable to the author, entirely trustworthy, and indis- pensable to the student and practitioner. — N. Y. An- nalist. MEDICAL AND SCIENTIFIC PUBLICATIONS. 75 A Series of Text-books on Physical Science. HANDBOOKS OF MTURAL PHILOSOPHY AND ASTRONOMY. BY DIONYSIUS LARDNER, D. C. L., Formerly Professor of Natural Philosophy and Astronomy in University College, London. This valuable series is now complete, consisting- of three Courses, as follows : — FIRST COURSE. 51ECHANICS, RYDROSTATICS, HYDRAULICS, PNEUMATICS, SOUND, AND OPTICS. In one large royal \2mo. volume of 750 pages, with 424 illustrations / $1 75. SECOND COURSE. HEAT, MAGNETISM, COMMON ELECTRICITY, AND VOLTAIC ELECTRICITY. In one royal 12mo, volume of AbO 2:)ages, with 244 illustrations j $1 25. THIRD COURSE. ASTRONOMY AND METEOROLOGY. In one very large royal 12»io. volume of nearly 800 pages, with 2>1 pilates and over 200 illus- trations. $2. These volumes can be had either separately or in uniform sets, containing About two thousand pages, and nearly one thousand Illustrations on Steel and Wood. To accommodate those who desire separate treatises on the leadmg departments of Natural Philosophy, the First Course may also be had divided in three portions, as follows : — Part I.— MECHANICS, price 70 cents. «' II.— HYDROSTATICS, HYDRAULICS, PNEUMATICS, and SOUND, price 60 cts. " III.— OPTICS, price 70 cents. It will thus be seen that this work furnishes either a complete course of instruction on these subjects, or separate treatises on all the different branches of Physical Science. The object of the author has been to prepare a work suited equally for the collegiate, acade- mical, and private st udent, who may desire to acquaint himself with the present state of science, in its most advanced condition, without pursuing it through its mathematical consequences and details. Great attention has been paid throughout the w^ork to elucidating the principles ad- vanced by their practical applications to the wants and purposes of civilized life, a task to which Dr. Lardner's immense and varied knowledge, and his singular felicity and clearness of illustra- tion render him admirably fitted. This peculiarity of the work recommends it especially as the text-book for a practical age and country such as ours, as it interests the student's mind by show- ing him the utility of his studies, while it directs his attention to the further extension of that utility by the fulness of its examples. Its extensive adoption in many of our most distinguished colleges and seminaries is sufficient proof of the skill with which the author's intentions have been carried out. From Prof. W. L. Brown, Oakland College, Miss. I consider them most admirably suited for the pur- poses designed by the author — indeed, as the very best popular works on physical science with which I am acquainted The '-Third Course," on Astronomy, is especially valuable; its magnificent engravings and lucid explanations make it a most desirable text- book. From Sev. J. 6. Ralston, JVorristown, Pa. Lardner's Meteorology and Astronomy is a fit com- panion for his First and Second Course. It is won- derfully minute, and yet not prolix. The principles of Astronomy are probably as clearly defined and judiciously arranged in this book as they can be. I expect to introduce it in my school. From S. Schooler, Esq., Hanover Academy, Va. The three volumes constitute a body of information and detail on nearly the whole range of physical sci- ence which is not to be found together in any other publication with which I am acquainted. I hope that these works may be the means of inducing many of our youth to devote themselves to the development of the Laws of Nature, and the application of them to industry, and that tliey may be the vehicle for con- veying sound information and food for thought to every man who aspires to be well educated. From M. Conant, State Normal School, Mass. This is a treatise admirably adapted to its purpose. For the accurate knowledge it unfolds, and the very popular dress it appears in, I think I have met with nothing like it. I shall advise the students of the Normal School to add this to your edition of Lardner's Mechanics, &c. From Prof. E. Everett, New Orleans. I am already acquainted with the merits of this book, having had occasion to consult it in teaching the branches of which it treats, and I cannot give you a stronger assurance of my high opinion of it than the simple fact that I have selected it as the text-book of Physics in the course of study which I have just fixed upon for a new college to be established here. 76 BLANCHARD AND LEA'S In one very handsome oclavo volume, of nearly six hundred and fifty pages, with over five hundred beautiful Illustrations on wood, and two colored Plates, extra cloth, $3 50. Hart's Calorimotor Structure of the Larynx. PRKiCIPLES OF PHYSICS AND METEOPiOLOGY. BY J. MULLER, Professor of Physics at the University of Freiburg, &c. Revised, with Additions, by R. E. GRIFFITH, M. D. In the work before us, the first thing which strikes us is the profusion of wood-cuts beyond what we should have imagined any publisher could furnish at the price— and very good cuts. The matter of the work is sound, and ranges over the subjects well. The manner is popular, and the author, though a mathe- matician, introduces only formulae to look at now and then, without algebraic processes. We strongly re- commend this work, particularly for the libraries of schools, mechanics' institutes, and the like. The pre- sent volume contains excellent reading and reference for those who are to have but one book on the subject. — AtkencBum. The plan adopted by Muller is simple ; it reminds us of the excellent and popular treatise published many years since by Dr. Arnott, but it takes a much wider range of subjects. Like it, all the necessary explanations are given in clear and concise language, without more than an occasional reference to mathe- matics; and the treatise is most abundantly illus- trated with well-executed wood engravings. The author has actually contrived to comprise in about five hundred pages, including the space occupied by illustrations, Mechanics, the laws of Motion. Acous- tics, Light, Magnetism, Electricity, Galvanism, Electro- Magnetism, Heat, and Meteorology. Medical practitioners and students, even if they have the means to procure, have certainly not the time to study an elaborate treatise in every branch of science; and the question therefore is, simply, whe- ther they are to remain wholly ignorant of such sub- jects, or to make a profitable use of the labors of those who have the happy art of saying or suggesting much in a small space. ELEMENTS OF NATURAL PHILOSOPHY; BEING AN EXPERIMENTAL INTRODUCTION TO THE PHYSICAL SCIENCES. Illustrated with over three hundred wood-cuts. By Golding Bird, M. D., Assistant Physician to Guy's Hospital. From the Third and Improved London edition. In one handsome royal 12mo. volume, of over 400 very large pages, on small type, leather, f 1 50 ; extra cloth, $1 '25. ELEMENTS OF PHYSICS ; OR, NATURAL PHILOSOPHY, GENERAL AND MEDICAL. Written for universal use in plain, or non-technical language. By Neill Arnott, M. D. A new edition, revised and corrected from the last English edition, with Additions, by Isaac Hays, M. D. With about two hundred illustrations. In one octavo volume, of nearly five hundred pages, leather, $2 50. ZOOPHYTES AND CORALS. By James D. Dana. In one large imperial quarto volume, extra cloth, with wood-cuts. (U. S. Exploring Expedition.) f 15 00. Also, An Atlas to the above, in one imperial folio volume, half bound in morocco, with sixty- one large plates, exquisitely colored after nature, f 30 00. BY THE SAME ATJTHOE. THE STRUCTURE AND CLASSIFICATION OF ZOOPHYTES, volume, extra cloth. $4 50. In one imperial quarto ASPECTS OF NATURE IN DIFFERENT LANDS AND DIFFERENT CLIMATES. With Scientific Elucidations. By Alexander Von Humboldt. Translated by Mrs. Sabine. In one large and handsome royal 12mo. volume, extra cloth. $1 00. MEDICAL AND SCIENTIFIC PUBLICATIONS, 77 HBRSCHEL'S ASTRONOMY. OUTLINES OF ASTRONOMY. BY SIR JOHN F. W. HERSCHEL, Bart., F. R. S., &c. g. ncfa American, from tljc fourtlj anir nhm)i IToivbon tintioix. In one handsome crown octavo volume, with numerous plates and loood-euts, extra cloth, $1 60; or, half bound, leather backs and cloth sides, $1 75. The present work is reprinted from the last London edition, which was carefully revised by the author, and in which he embodies the latest investigations and discoveries. It may therefore be re°-arded as fully on a level with the most advanced state of the science, and even better adapted than its predecessors, as a full and reliable manual. From Prof. A. Casivdl, Brown University, R. J. As a work of reference and study for the more ad- vanced pupils, who yet are not prepared to avail them- selves of the higher mathematics, 1 know of no work to be compared with it. From Professor D. Olmsiead, Tale College. A rich mine of all that is most valuable in modern Astronomy. THE PRINCIPLES OF ANIMAL AND VEGETABLE PHYSIOLOGY. A POPULAR TREATISE ON THE FUNCTIONS AND PHENOMENA OF ORGANIC LIFE. BY J. STEVENSON BUSHNAN, M. D. "With 102 Illustrations. J||)j| ii^...^ ... Hill ' i^At'''' 4v1 I In one very neat royal 12mo. vohime, of 234 s|| \vi| ^ ^i^^ pages, extra cloth, 90 cents. % % il\jijlW^' Spiracle of Fly. The Illustrated Geographical Encyclopaedia. THE ENCYCLOPJlDll OF GEOGRAPHY. COMPRISING A COMPLETE DESCRIPTION OF THE EARTH, PHYSICAL, STATISTICAL, CIVIL, AND POLITICAL. EXHIBITING ITS RELATION TO THE HEAVENLY BODIES, ITS PHYSICAL STRUC- TtJRE, THE NATURAL HISTORY OF EACH COUNTRY, AND THE INDUSTRY, COMMERCE, POLITICAL INSTITUTIONS, AND CIVIL AND SOCIAL STATE OP ALL NATIONS. BY HUGH MURRAY, F. R. S. E., &c. Assisted in Botany, by Professor HOOKER— Zoology, &c., by W. W. SWAINSON— Astrono- my, &c., by Professor WALLACE— Geology, &c., by Professor JAMESON. REVISED, WITH ADDITIONS, BY THOMAS G. BRADFORD. In three large octavo volumes, containing about nineteen hundred large imperial pages, and ILLUSTRATED BY EIGHTY-TWO SMALL MAPS. And a colored Map of the United States, after Tanner's ; together vf'Ah. about Eleven Hundred Wood-cuts, executed in the best style. In leather, gilt, $5 00; library style, $4 50; extra cloth, $4. Skinner's Complete Edition of Youatt on the Horse. A BOOK FOR EVERY FARME R A ND COUNTRY GENTLEMAN. THE HORSE. By William Youatt. A new edition, with numerous Illui=trations. Toge- ther with a General History of the Horse; a Dissertation on the American Trotting Horse, how Trained and Jockeyed; an account of his remarkable performances; and an Essay on the Ass and the Mule. By J. S. Skinner, Assistant Postmaster-genera), and Editor of the Turf Register. In one octavo volume, of neai'ly 450 large pages, with numerous Illustra- tions; price, in extra cloth, f 1 50; leather, fl 7-S. 78 BLANCHARD AND LEA'S THE BOOK OF NATUKE AN ELEMENTARY INTRODUCTION TO THE SCIENCES OF PHYSICS, ASTRONOMY, CHEMISTRY, MINERALOGY, GEOLOGY, BOTANY, ZOOLOGY, AND PHYSIOLOGY. . BY FRIEDRICH SCHOEDLER, Ph. D., Professor of the Natural Sciences at Worms, &o. &c. First American Edition, with a Glossary and other Additions and Improvements, from the Second English Edition TRANSLATED FROM THE SIXTH GERMAN EDITION, BY HENRY ME BLOCK, F. C. S., &c. Profu sel y illustrated with nearly seven hundred beautiful engravings on wood. In one large and handsome crown octavo volume, of 692 pages, extra cloth ; price fl 80. The Geysers of Iceland (Geology). To accommodate those who desire to use the separate portions of this work, the publishers have prepared an edition in parts, as follows, which may be had singly, neatly done up in flexi- ble cloth, at 50 ceats each. Natural Phllosopliy, 114 pages, with 149 illustrations. Astrouomy, 64 " " 51 '• Chemistry, 110 « " 48 " Mineralogy and Geology, 104 pages, with 167 " This volume, as its title shows, covers nearly all the sciences, and embodies a vast amount of information for instruction. No other work that we have seen presents the reader with so wide a range of element- ary knowledge, with so full illustrations, at so cheap a rate. — Silliman's Journal. Written with remarkable clearness, and scrupu- lously correct in its details. — Mining Journal. His expositions are most lucid. There are few who will not follow him with pleasure as well as with pro- fit through his masterly exposition of the principles and primary laws of science. It should certainly be made a class-book in schools. — Critic. Botany, 98 pages, with 176 illustrations. Zoology and Pbyslology, 106 pages, with 84 " Introduction, Glossary, Index, ^c., Composed by the same distinguished author, all the departments have a uniformity of style and illustra- tion which harmoniously link the entire circle to- gether. The utility of such a connected view of the physical sciences, and on such an approved basis, is beyond price ; and places their acquisition within the reach of a vastly increased number of inquirers. Not only to such is it valuable, but to those who wish to have at hand the means of refreshing their memories and enlarging their views upou their favorite studies. Of such a book we speak cordially, and would speak more at length, if space permitted. — Southern Metlio- dist Quarterly Review. THE DOa. By WILLIAM YOUATT. Edited, with Additions, by E. J. LEWIS, M. D. WITH NUMEROUS PLATES AND WOOD-GUTS. In one very handsome volume, crown octavo, done up in beautiful crimson cloth, gilt stamps, $1 25. MEDICAL AND SCIENTIFIC PUBLICATIONS. 79 PRINCIPLES OF THE MECHANICS OF MACHINERY AND ENGINEERING. BY PROFESSOR JULIUS WEISBACH. Translated and Edited by PROFESSOR GORDON, or Glasgow. Jtrst American ^bition, iaitlj ^bititions, BY PROFESSOR WALTER R. JOHNSON. In tivo very handsome octavo volumes, of nearly 900 pages, with about 900 superb Jlhistraiions, handsomely and strongly bound in extra cloth, $6 50. Overshot Water- Wheel. These volumes contain an immense amount of practical information necessary to the ma- chinist, architect, and civil engineer ; the whole thoroughly brought up to the most advanced state of scientific investigation. The following rapid summary of the contents will show the manner in which the subjects are successively brought forward. Volume H. Section I. Application of Mechanics in Build- Volume I. Sect. I. Mathematical Science of Motion. Sect. II. Mechanics in the Physical Science of Motion in general. Sect. III. Statics of Rigid Bodies. Sect. IV. Dynamics of Rigid Bodies. Sect. V. Statics of Fluid Bodies. Sect. VI. Dynamics of Fluid Bodies. Division II. Application of Mechanics to Ma- chinery. Section II. Of Moving Powers and their Ef- fects. These subjects, treated in all their ramifications and practical applications to Machines, Water-powers, Dams, Bridges, Water-wheels, Turbines, Roofs, Walls, Arches, Windmills, Scales, Steam-boilers, Water-engines, &c. &c., present points of primary importance to every one engaged in machine-work of every description, building, or engineering ; while the compre- hension of the whole is greatly facilitated by the profusion of clear and beautifully executed illustrations which are interspersed throughout the work. THE YOUNG MILLWRIGHT, AND MILLER'S GUIDE. Illustrated by twenty-eight Descriptive Plates. By Oliver Evans, with Additions and Corrections by Thomas P. Jones. With a Description of an Improved Merchant Flour-mill, by C. and O. Evans. Fourteenth edition. In one handsome octavo volume, strongly bound in leather, $2 50. BLANCHARD AND LEA'S PUBLICATIONS. New and much Improved Edition — (Lately Issued.) PHYSICAL GEOGRAPHY. BY MARY SOMERVILLE. A NEW AMERICAN, FROM THE THIRD AND REVISED LONDON EDITION. WITH NOTES AND A GLOSSARY, BY W. S. W. RUSCHENBERGER, M. D., U. S. Navy. In one large royal llmo. volume, of nearly 600 pages, extra cloth, $1 2b; half hound in leather, $1 36. Eulog-y is unnecessary with regard to a work like the present, which has passed through three editions on each side of the Atlantic within the space of a few years. The publishers therefore only consider it necessary to stale that the last London edition received a thorough revision at the hands of the author, who introduced whatever improvements and corrections the advance of science rendered desirable, and that the present issue, in addition to this, has had a careful examination on the part of the editor, to adapt it more specially to this country. Great care has been exercised in both the text and the glossary to obtain the accuracy so essential to a work of this nature, and in its present improved and enlarged state, with no corresponding in- crease of price, it is confidently presented as in every way worthy a continuance of the striking favor with which it has been everywhere received. From Lieutenatjt Maury, U. S. iV. National Observatory, Washington. I thank you for the " Physical Geography ;" it is capital. I have been reading it, and like it so much that I have made it a school-book for my children, whom I am teaching. There is, in my opinion, no ■work upon that interesting subject on which it treats — Physical Geography — that would make a better text- book in our schools and colleges. I hope it will be adopted as such generally, for you have Americanized it and improved it in other respects. From Samuel H. Taylor, Esq., Philips Academy, Andcnxr, Mass., Feb. 15, 1854. We have introduced your edition of Mrs. Somer- ville's '■ Physical Geography" into our school, and tind it an admirable work. From Thomas Shermin, High School, Boston. T hold it in the highest estimation, and am confident that it will prove a very efiScient aid in the education of the young, and a source of much interest and in- struction to the adult reader. THE ENCYCLOPEDIA AMERICANA. A POPULAR DICTIONARY OF Arts, Sciences, Literature, History, Politics, Biography; INCLUDING A COPIOUS COLLECTION OF ORIGINAL ARTICLES IN AMERICAN BIOGRAPHY. ON THE BASIS OF THE SEVENTH EDH ION OF THE GERMAN CONVERSATIONS-LEXICON. Edited by FRANCIS LIEBER, Assisted by E. WIGGLESWORTH and T. G. BRADFORD. With Additions by Pkofessor HENRY VETHAKE, Of the University of Pennsylvania. In fourteen large octavo volumes, containing in all nearly nine thousand large double-columned pages, furnished in various styles of binding, at very low prices. THE GEOLOGICAL OBSERVER. BY SIR HENRY T. DE LA BECHE, C. B., l\ R. S., Director-General of the Geological Survey of the United Kingdom. WITH OVER THREE HUNDRED In one very large and handsome octavo volume, of 700 pages, extra cloth, $4 00. If '•^ '% \< -K .,r«.- ■;»"'■;);, r.'.'WJi, '?fjW-» ^^t^ Si