THIS BOOK IS THE GIFT OF The Van Cleef Library Cornell University Library QP 34.D15 1864 A treatise on human physiology. 3 1924 001 037 591 JAN 5 1965 Date Due Library Burea Cat. No. 1137 The original of tiiis book is in tine Cornell University Library. There are no known copyright restrictions in the United States on the use of the text. http://www.archive.org/details/cu31924001037591 A TREATISE HUMAN PHYSIOLOGY DESIGNED FOB TUB USE OP STUDENTS AND PMCTITIONERS OF MEDICINE. BY JOHN C. DALTON, Jr., M. D., PhOFESSOR OF PHYBIOLOaT AND MICROSCOPIC AWATOMT IN THE COLLEGE OF PHYSICIANS AND BURGEONS, NEW TOKK ; MEMBER OF THE NEW YORK ACADEMY OF MEDICINE j OF THE NEW YORK PATHOLOGICAL SOCIETY ; OP THE AMERICAN ACADEMY OP ARTS AND SCIENCES, BOSTON, MASS. ; AND OF THE BIOLOGICAL DEPARTMENT OF THE ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA. SC^irJj ffibittflti, '^zbisz'a aitb dnlargcb. TWO HUNDRED AND SEYENTY-THREE ILLUSTKATIONS. PHILADELPHIA: BLANCHAED AND LEA. 1864. KsiERED according to the Act of Congress, in the year 1864, by BLANCHARD AND LEA, in the Oifice of the Clerk of the District Court of the United States in and for the Eastern District of the State of Pennsylvania. 54- PlS PHILADELPHIA: COLLIKS, PKISTBH, 705 JAYNB STKEET. TO MY FATHER, JOHN C. DALTON, M.D., IH HOMAGE OP HIS LONG AND StJOCESSPUL DEVOTION TO THE • SCIENCE AND ART OF MEDICINE, AND I 17 • GRATEFUL RECOLLECTION OP HIS PROFESSIONAL PRECEPTS AND EXAMPLE, f lis ifllumj IS RESPECTFULLY AND AFFECTIONATELY INSCEIBED. ( iii ) PREFACE TO THE THIRD EDITION. In the present edition of this work, the general plan and arrange- ment of the two former ones are retained. The improvements and additions which have been introduced consist in the incorporation into the text of certain new facts and discoveries, relating mainly to details, which have made their appearance within the last three years. Such are the experiments of the author with regard to the secretion and properties of the parotid saliva in the human subject, and the quantitative analysis of this fluid by Mr. Perkins ; the valuable observations of Prof. Austin Flint, Jr., on Stercorine, Oholesterin, and the effects of permanent biliary fistula, and those of Prof. Jeffries Wyman on Fissure of Hare-lip in the median line, from arrest of development. Three new illustrations have been introduced, one of which (Fig. 183) repkces a previous one. The author ■ is much indebted to his friend. Dr. Foster Swift, for aid in carrying the work through the press. New Yoek, January, 1864. (V) PREFACE TO THE SECOND EDITION. In presenting a new edition of this work, tlie author desires to express his sincere acknowledgments to hia professional brethren for the very favorable manner in which it was received at the time of its first appearance, two years ago. In the present edition, the author has endeavored to supply, as fully as possible, the deficiencies which, he is well aware, existed in the former volume. Some of these deficiencies were evident to his own mind, while others were indicated by the suggestions of judicious criticism. These suggestions, accordingly, have been adopted in all cases in which they appear to be well founded, and not inconsistent with the general plan of the work. In those instances, on the other hand, in which the views of the author on physiological questions seemed to him to be positively sustained by the results of observa- tion, he has retained these views unchanged in the present edition. At the same time, he has abstained^ as before, from the lengthened discussion of theoretical points, and has purposely avoided even the enumeration of new experiments and observations, wherever they have not materially affected the position of physiological doctrines; for in a work like the present, it is not the object of the writer to give a detailed history of physiological science, but only such prominent and essential points in its development as will enable the reader fully to comprehend its actual condition at the present time. The principal additions and alterations which have thus been found advisable are: — First, the. introduction of an entire chapter devoted to the con- sideration of the Special Senses, which were only incidentally treated of in the former edition. ( vii ) Vm PKEFACE TO THE SECOND EDITION. Second, the re-arrangement of the chapter on the Oranial Nerves, and the introduction of some new views and facts in regard to their physiology. Third, an account of some new experiments, original with the ■ author, relating to the function of the Cerebellum, and the conclu- sions to which they lead. Fourth, certain considerations respecting the general properties of Sensation and Motion, as resident in the nervous system, which are important as an introduction to the more detailed study of these functions. Fifth, the introduction of a chapter on Imbibition and Exhalation and the functions of the Lymphatic System; including the study of endosmosis and exosmqsis, and their mode of action in the animal frame, the experiments of Dutrochet, Chevreuil, Gosselin, Matteucci) and others, on this subject, the constitution and circulation of the lymph and chyle, and, finally, a quantitative estimate of the entire processes of exudation and reabsorption, as taking place in the living body. Additions have also been made, in various parts, to the chapter.s on Secretion, Excretion, the Circulation, and the functions of the Digestive Apparatus. In every instance, these alterations have been incorporated with the text in such a manner as to avoid, so far as possible, increasing unnecessarily the size of the book. Twenty-two new and original illustrations have been introduced into the present volume, of which number five replace others in the former edition, which were regarded as imperfect, either in design or execution. The remaining seventeen are additional. It is hoped that the above alterations and additions will be found to be improvements, and that they will enable the work, in its pre- sent form, to accomplish more fully the object for which it was designed. New Yoek, February, 1861. PREFACE TO THE FIRST EDITION. This volume is offered to the medical profession of the United States as a text-book for students, and also as a means of commu nicating, in a condensed form, such new facts and ideas in physio- logy as have marked the progress of the science within a recent period. Many of these topics are of great practical importance to the medical man, as influencing, in various ways, his views on pathology and therapeutics; and they are all of interest for the physician who desires to keep pace with the annual advance of his profession, as indicating the present position and extent of one of the most progressive of the departments of medicine. It has been the object of the author, more particularly, to pre- sent, at the same time with the conclusions which physiologists have been led to adopt on any particular subject, the experimental basis upon which those conclusions are founded; and he has en- deavored, so far as possible, to establish or corroborate them by original investigation, or by a repetition of the labors of others. This is more especially the case in that part of the book (Section I.) devoted to the function of Nutrition; and as a general thing, throughout the work, any statement of experimental facts, not expressly referred to the authority of some other writer, is given by the author as the result of direct personal observation. ' The illustrations for the work have been prepared with special reference to the subject-matter ; and it is hoped that they will be found of such a character as materially to assist the student in comprehending the most important and intricate parts of the sub- ject. It is more particularly in the departments of the Nervous System and Embryonic Development that simple, clear, and faithful (ix) X PREFACE TO THE FIRST EDITTOK. illustrations are indispensable for the proper understanding of the printed descriptions ; the latter being often necessarily somewhat intricate, and requiring absolutely the assistance of properly arranged figures and diagrams. Of the two hundred and fifty- four illustrations in the present volume, only eleven have been • borrowed from other writers, to whom they will bf found duly credited in the list of woodcuts. Of the remaining illustrations, prepared expressly for the pre- sent work, the drawings of anatomical structures, crystals, and microscopic views generally were all taken from nature. The diagrams were arranged, for purposes of convenience, in such a manner as to illustrate known anatomical or physiological ap- pearances, in Uhe most compact and intelligible form. Physiological questions which are in an altogether unsettled state, as well as purely hypothetical topics have been purposely avoided, as not coming within the plan of this work, nor as calcu- lated to increase its usefulness. New York, January 1, 1859. CONTENTS. INTRODUCTIO-N. s PAGE Definition of Physiology — Its mode of study — Nature of Vital Phenomena — Division of the subject 49-59 SECTION I. NUTEITION. CHAPTER I. PROXIMATE PRINCIPLES IN GENERAL. Definition of Proximate Principles — Mode of their extraotion^Manner in which they are associated with each other — Natural variation in their relative quantities — Three distinct classes of proximate principles . . . 61-68 CHAPTER II. PROXIMATE PRINCIPLES OF THE EIRST CLASS. Inorganic substances — Water — Chloride of Sodium — Chloride of Potassium — Phosphate of Lime — Carbonate of Lime — Carbonate of Soda — Phosphates of Magnesia, Soda, and Potassa — Inorganic proximate principles not altered in the body — Theii: discharge — Nature of their function .... 69-78 CHAPTER III. PROXIMATE PRINCIPLES OF THE SECOND CLASS. Staboh — Percentage of starch in different kinds of food — Varieties of this substance — Properties and reactions of starch — Its conversion into sugar — SoGAB — Varieties of sugar — Physical and chemical properties — Proportion in different kinds of food — Fats — Varieties — Properties and reactions of fat — Its crystallization — Proportion in different kinds of food — Its condition in the body — Internal production of fat — Origin and destination of proximate principles cf this class 79-94 ( xi ) XU CONTENTS. CHAPTER IV. PROXIMATE PRINCIPLES OE THE THIRD CLASS. PACE General characters of organic substances — Their chemical constitution — Hygro- scopic properties — Coagulation — Catalysis — Fermentation — Putrefaction — Fibrin — Albumen — Casein — Globuline — Pepsine — Pancreatine — Mucosine — Ostelne' — Cartllaglne — Musculine — Hsematine — Melanine — Biliverdine — tJrosaclne — Origin and destruction of proximate principles of this class 95-104 CHAPTER V. or FOOD. Importance of inorganic substances as ingredients of food — Of saccharine and starchy substances — Of fatty matters — Insufficiency of these substances when used alone — Effects of an exclusive non-nitrogenous diet — Organic substances also Insufficient by themselves — Experiments of Magevdle on exclusive diet of gelatine or fibrin — Food requires to contain all classes of proximate principles — Composition of various kinds of food — Dally quantity of food required by man — Digestibility of food — Effect of cooking . 105-114 CHAPTER YI. DIGESTION. Kature of digestion — Digestive apparatus of fowl — Of ox — Of man — Mastica- tion — Varieties of teeth — Effect of mastication — Saliva — Its composition — Daily quantity produced — Its action on starch — Effect of its suppression — Function of the saliva — Gastric Juice, and Stomach Digestion — Structure of gastric mucous membrane — Dr. Beaumont's experiments on St. Martin — Artificial gastric fistulse — Composition and properties of gastric juice — Its action on albuminoid substances — Peristaltic action of stomach — Time re- quired for digestion — Daily quantity of gastric juice — Influences modifying its secretion^-lNTESTiNAL Juices, and the Digestion op Sugae and Starch Follicles of Intestine — Properties of intestinal juice — Pancreatic Juice, and the Digestion of pat — Composition and properties of pancreatic juice Its action on oily matters — Successive changes in intestinal digestion The large intestine and its contents 115-161 CHAPTER VII. ABSORPTION. Closed follicles and villi of small Intestine — Peristaltic motion Absorption by bloodvessels and lymphatics — Chyle — Lymph — Absorbent system— Jl,ac ■ teals and lymphatics — Absorption of fat — Its accumulation in the blood during digestion — Its final decomposition and disappearance . . 162-174 CONTENTS. Xm CHAPTER VIII. THE BILE. PAGE Physical properties of tlie bile — Its composition^— Blliverdine — Cholesterin — Biliary salts — Their mode of extraction — Crystallization — Grlyko-cholate of soda — Tanro-eholate of soda — Biliary salts in different species of animals and in man — Tests for bile— Variations and functions of bile — Daily quan- tity — Time of its discharge into intestine — Its disappearance from the ali- mentary canal-^Its reabsorption — Its ultimate decomposition . . 175-199 CHAPTER IX. FORMATION OF SUGAR IN THE LIVER. Existence of sugar in liver of all animals — Its percentage — Internal origin of liver-sugar — Its production after death — Glycogenic matter of the liver — Its properties and composition — Absorption of liver-sugar by hepatic veins — Its accumulation in the blood during digestion — Its final decomposition and disappearance ........ 200-207 CHAPTER X. THE SPLEEN. Capsule of Spleen — Variations in size of the organ — Its internal structure — Malpighian bodies of the spleen — Action of spleen on the blood — Effect of its extirpation ....... '208-212 CHAPTER XI. "^ THE BLOOD. Red Globules of the blood — Their microscopic characters — Structure and com- position — Variations in size in different animals — White Globdlbs of the blood — Independence of the two kinds of blood-globules — Plasma — Its com- position — Fibrin — Albumen — Fatty matters — Saline ingredients — Extractive matters — Coagulation of the Blood — Separation of clot and semm — Influ- ences hastening or retarding coagulation — Coagulation not a commencement of organization — Formation of bnffy coat — Entire quantity of blood in body 213-231 CHAPTER XII. RESPIRATION. Respiratory apparatus of aquatic and air-breathing animals — Structure of lungs in human subjects — Respiratory movements of chest — Of glottis — Changes in the air during respiration — Changes in the blood — Proportions of oxygen and carbonic acid, in venous and arterial blood — Solution of gases by the blood-globules — Origin of carbonic acid in the blood — Its mode of production — Quantity of carbonic acid exhaled from the body — Variations according to age, sex, temperature, &c. — Respiration by the skin . 232-252 S;iV CONTENTS. CHAPTER XIII. ANIMAL HEAT. PAGE Standard temperature of animals — How maintained — Production of heat by Vegetables — Mode of generation of animal heat — Theory of combustion — Objections to this theory — No oxidation in vegetables during production of heat — Quantities of oxygen and carbonic acid in animals do not correspond ■with each other — Production of animal heat a local process — Depends on the chemical phenomena of nutrition . . . . • 253-263 CHAPTER XIV. THE CIRCULATION. Giroulatory apparatus of fish — Of reptiles — Of mammalians — Course of blood through the heart— Action of valves — Sounds of heart — Movements — Im- pulse — Successive pulsations — Arterial system — Movement of blood through the arteries — Arterial pulse — Arterial pressure — Rapidity of arterial circula- tion — The veins — Causes of movement of blood in the veins — Rapidity of venous current — Capillary circulation — Phenomena and causes of capillary circulation — Rapidity of entire circulation — Local variations in different parts ......... 264-306 CHAPTER XV. IMBIBITION AND EXHALATION. — THE LYMPHATIC SYSTEM. Endosmosis and exosmosis — Mode of exhibiting them — Conditions which regu- late their activity — Nature of the membrane — Extent of contact — Constitu- tion of the liquidS-^Temperature — Pressure — Nature of endosmosis — Its conditions in the living body— Its rapidity— Phenomena of endosmosis in the circulation — The lymphatics — Their origin — Constitution of the lymph and ehyle— Their quantity— Liquids secreted and jreabsorbed in twenty- four hours ........ 307-323 CHAPTER XVI. SECRETION. Nature of secretion — Variations in activity — Mucus — Sebaceous matter Its varieties — Perspiration — Structure of perspiratory glands — Composition and quantity of the perspiration— Its use in regulating the animal temperature- Tears — Milk— Its acidification— Secretion of bile — Anatomical peculiarities 324-340 COXTENTS. XT CHAPTER XYII. EXCRETION. PAGE Nature of excretion — Exorementitious substances — Effect of their retention — Urea — Its source — Couversiou into carbonate of ammonia — Dally quantity of urea — Creatine — Creatinine — Urate of soda — Urates of potassa and ammo- nia — General characters of the urine — Its composition — Variations — Acci- dental Ingredients of the urine — Acid and alkaline fermentations — Final decomposition of the urine . . . ' . . • 341-364 SECTION II. NERVOUS SYSTEM. CHAPTER I. GENERAL CHARACTER AND FUNCTIONS OF THE NERVOUS SYSTEM. Nature of the function performed by nervous system — Two kinds of neryous tissue — Fibres of white substance — Their minute structure — Division and inosculation of nerves — Gray substance — Nervous system of radiata — Of moUusoa — Of artlculata — Of mammalia and human subject — Structure of encephalon — Connections of its different parts . . . 365-?87 CHAPTER II. OF NERVOUS IRRITABILITY, AND ITS MODE OF 'ACTION. Irritability of muscles — How exhibited — Influences which exhausj; 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 — Differences between the two 388-397 CHAPTER III. THE SPINAL CORD. Power of sensation — Power of motion — 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 during disease — Influence in health on sphincters, voluntary muscles, urinary bladder, &c. . . . 398-416 XVI contents'. CHAPTER IV. THE BRAIN. PAGE Seat of sensibility and excitability in different parts of the encephalon — Olfac- tory ganglia — Optic tlialami— Corpora striata — Hemispheres — Remarkable cases of injury of hemispheres — Effect of their removal — Imperfect develop- ment in idiots — Aztec children — Theory of phrenology — Cerebellum — Effect of its injury or removal — Comparative development in different classes — Tubercula quadrigemina — Tuber annulare— MeduUaoblongata — Three kinds of reiiex action in nervous system ..... 417-445 CHAPTER V. THE CRANIAL NERVES. Olfactory nerves — Optic nerves — Auditory .nerves — Classification of cranial nerves — Motor nerves — Sensitive nerves — Motor oeuli communis — Patheti- cus — Motor externus — Fifth pair — Its sensibility — Effect of division — Influ- ence on mastication — Influence on the organ of sight — Facial nerve— Effect of its paralysis — Glosso-pharyngeal nerve — Pneumogastric — Its distribution — Influence on phj^rynx and CBSophagus — On larynx — On lungs — On stomach and digestion — Spinal accessory nerve — Hypoglossal . . . 446-477 CHAPTER VI. THE SPECIAL SENSES. General and special sensibility — Sense of touch in the skin and mucous mem- branes — Nature of the special senses — Tastk — Apparatus of this sense — Its conditions — Its resemblance to ordinary sensation — Injury to the taste in paralysis of the facial nerve — Smell — Arrangement of nerves in nasal pas- sages — Conditions of this sense — Distinction between odors and irritating vaporsr-SiGHT— Structure of the eyeball — Special sensibility of the retina — Action of the lens — Of the iris — Combined action of two eyes — Vivid nature of the visual impressions — Hearino — Auditory apparatus — Action of mem- brana tympani — Of chain of bones — Of their muscles — Appreciation of the direction of sound — Analogies of hearing with ordinary sensation . 478-513 CHAPTER VII. 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 .... 514-524 CONTENTS. 2V11 SECTION III. REPRODUCTION. CHAPTER I. ON THE NATURE OP EEPRODUOTION, AND THE ORIGIN OF PLANTS AND ANIMALS. PAGE Nature and objectg 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 inffisoria — Conditions of their development — Schultze's experiment on generation of infusoria — Pro- duction of animal and vegetable parasites — Encysted entozoa — Trichina spiralis' — Taenia, — Cysticerous — Production of taenia from oysticercus — Of cysticercus from eggs of taenia — Plants and animals always produced by generation from parents ...... 525-539 CHAPTER II. ON SEXUAL (^ENERATION AND THE MODE OP 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 640-543 CHAPTER III. ON THE EGG, AND THE PEMALE ORGANS OP GENERATION. 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 . . 644-555 • CHAPTER IV. ON THE SPERMATIC PLUID, AND THE MALE ORGANS OP GENERATION. The spermatozoa — Their varieties in different species — Their movement — For- mation of spermatozoa in the testicles — Accessory male organs ol^eneration — Epididymis — Vas deferens — Vesiculae seminales — Prostate — Cowper's glands — Function of spermatozoa — Physical conditions of fecundation 556-562 2 • XVIU CONTENTS. CHAPTER V. t I ON PEKIODICAL OVULATION, AND THE FUNCTION OF MENSTRUATION. . PAGE Periodioai, Ovulatioit — 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 oestruation — Menstkuation — 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 663-575 CHAPTER VI. ON THE CORPUS LUTEUM OF MENSTKUATION AND PREGNANCY. CoEPtJS LvTEum. OP Menstritation — Discharge of blood into the ruptured Graafian follicle — Deoolorization of the clot, and hypertrophy of the membrane of the vesicle — Corpus luteum of menstruation, at the end of three weeks^Tellow coloration of convoluted wall — Corpus luteum of menstruation at the end of four weeks — Shrivelling and condensation of Its tissues; — Its condition p,i the end of nine weeks — Its final atrophy and disappearance^CoEpus Lcteum OF Presnakcy — 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 . f . 576-585 CHAPTER VII. ON THE DEVELOPMENT OF THE IMPREGNATED EGGi. Segmentation of the vitellus — Formation of blastodermic membrane — Two layers of blastodermic membralie — 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 ........ 586-595 CHAPTER VIII. 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 . . 696-598 CONTENTS. XIX CHAPTER IX. AMNION AND ALLANTOIS — DEVELOPMENT OP THE CHICK. • PAGE Necessity for accessory organs iu the development of birds and quadrupeds — Formation of amniotic folds — TJieir union and adhesion — Growth of allantois from lower part of intestine — Its vascularity — Allantois iu 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 ........ 599-607 CHAPTER 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 vUlosities — 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 ...... 608-613 CHAPTER XI. DEVELOPMENT OF UTERINE MUCOUS MEMBRANE — FORMATION OF THE DECIDUA. Structure of uterine mucous membrane — Uterine tubules — Thickening of ute- rine mucous membrane after impregnation — Decidua vera — Entrance of egg into uterus — Deoidua reflexa — Inclosure of egg by decidua reflexa — Union of chorion with decidua — Changes in the relative development of different portions of chorion and decidua ..... 614-620 CHAPTER XII. THE PLACENTA. Nourishment of foetus by maternal and foetal vessels — Arrangementof the vascular membranes in different species of animals — Membranes of foetal pig — Cotyledon of cow's uterus — Development of foetal tufts in human pla- centa — Development of uterine sinuses — Relation of foetal and maternal bloodvessels in the' placenta — Proofs that the maternal sinuses extend through the whole thickness of the placenta — Absorption and exhalation by the placental vessels ...... 621-629 XX CONTENTS. CHAPTER XIII DISCHARGE OF THE OVUM, AND INVOLUTION OF THE UTERUS. ' PAG» Enlargement of amniotic cavity— Contact of amnion and chorion— Amniotic fluid— Movements of fcetus- Union of deoidua 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 deoidua Fatty degeneration and reconstruction of muscular -walls of uterus 630-636 CHAPTER XIV. DEVELOPMENT OP THE EMBRYO — NERVOUS SYSTEM, ORGANS OP SENSE, SKELETON AND LIMlBS. 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 vertebrse— Laminae and ribs— Spina bifida — Anterior and pos- terior extremities — Tail — Integument — Hair — Vernix caseosa — Exfoliation of epidermis ........ 637-643 CHAPTER XT. DEVELOPMENT OF 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 — Uraohus — Vesico-rectal septum — Perineum — Liver — Secretion of bile — Gastric juice — Meconium — Glyco- genic function of liver — Diabetes of foetus — Bharynx and oesophagus — Dia- phragm — Diaphragmatic hernia — Heart and pericardium — Ectopia cordis — Development of the face ...... 644-654 CHAPTER XYI, DEVELOPMENT OF THE KIDNEYS, WOLFFIAN BODIES, AND INTERNAL ORGANS * OF GENERATION. Wolflan 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 .... 655-664 CONTENTS. XXI CHAPTER XVII. DEVELOPMENT OF THE CIRCULATORY APPARATUS. PAGE 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-mesenterio arteries and vein-^- Ciroulation 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-r-Aorta and pulmonary artery — Ductus arteriosus — Foramen ovale and Eustachian valve — Changes in circulation at the period of birth 665-686 CHAPTER 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 — Difference in relative size of organs, in infant and adult — Withering and separation of umbilical cord — Exfoliation of epidermis — First and second sets of teeth — Subsequent changes in osseous, muscular and tegumentary systems, and general devel- opment of the body . 687-690 LIST OF ILLUSTRATIONS, ALL OF WHICH HAVE BEEN PEEPAEED PBOM OEIMNAL DBAWINOS, WITH THE EXCEPTION OF TEN, CREDITED TO THEIR AUTHOEITIES. FIO. 1. Fibula tied in a knot, after maceration in a dilate acid 2. Grains of potato starcK . 3. Starch grains of Bermuda arrowroot 4. Starch grains of wheat flour 5. Starch grains of Indian com ^. Starch grains from wall of lj,teral ventricle 7. Stearine .... 8. Oleaginous principles of human fat 9. Human adipose tissue 10. Chyle 11. Gclohulea 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'a 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 Aohille Richard PAGE 75 .80 80 81 81 82 87 88 90 90 91 91 92 92 93 117 118 119 121 122 122 122 123 124 133 133 134 134 134 140 151 152 158 158 159 XXUl XXIT LIST OF ILLUSTKATIONS. 36. From last quarter of small intestine 37. One of the closed follicles of Peyer's patches 38. Grlandulse agminatse 39. Extremity of intestinal villus 40. Panizza's experiment on absorption by bloodvessels 41. Chyle, from commencement of thoracic duet 42. Laoteals, thoracic duct, &c. 43. Lacteals and lymphatics . 44. Intestinal epithelium, in intervals of digestion 45. Intestinal epithelium, during digestion . 46. Cholesterin .... 47. Ox-bile, crystallized 48. Glyko-oholate of soda from ox-bile 49. Glyko-oholate 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 fistula .... 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 menobranchus . 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. XXV PIG. 86. 87. 88. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 136. Heart of frog, in contraction Simple looped fibres .... Bullock's heart, showing superficial muscular fibres Left ventricle of bullo'ek's>Tieart, showing deep fibres Diagram of circular fibres of the heart Converging fibres of the apex of the heart Arterj in pnlsation Curves of the arterial pulsation Volkmann's apparatus . The same .... Vein, with valves open . Vein, with valves closed Small artery, with capillary branches . Capillary network Capillary circulation Diagram of the circulation Follicles of a compound mucous glandule Meibomian glands Perspiratory gland . .n Glandular structure of mamma Colostrum corpuscles Milk-globules .... Division of portal vein in liver Lobule of liver Hepatic cells Urea .... Creatine Creatinine Urate of soda Uric acid Oxalate of lime . Phosphate of magnesia and ammonia Nervous filaments, from brain . Nervous filaments from sciatic nerve Division of a nerve Inosculation of nerves . Nerve cells . . • . Nervous system of starfish Nervous systemuaf aplysia Nervous system of centipede Cerebro-spinal system of man . Spinal cord Brain of alligator Brain of rabbit Medulla oblongata of human brain Diagram of human b>ain Experiment showing irritability of muscles Experiment showing irritability of nerve Action of direct and inverse currents Diagram of spinal cord and nerves From KoUiker From Ludovic From Todd and Bowman From Lehmann (Funke's Atlas) From Lehmann (Funke's Atlas) From Lehmann (Fnnke's Atlas) XXVI LIST OF ILLUSTEATIONS. PIG. 136. Spinal oord in vertical section 137. Experiment, showing effect of poisons upon 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. Brain of healthy pigeon in profile 144. Brain of operated pigeon in profile 145. Brain of healthy pigeon, posterior view 146. Brain of operated pigeon, posterior view 147. Inferior surface of brain of cod 148. Inferior surface of brain of fowl . 149. Course of optic nerves in man . 150. Distribution of fifth nerve upon the face 151. Facial nerve .... 152. Pneumogastrie nerve 153. Diagram of tongue 154. Distribution of nerves in the nasal passages 155. Vertical section of eyeball 156. Dispersion of rays of light 157. Action of crystalline lens 158. Myopia ...... 159. Presbyopia .... 160. Vision at short distance 161. Vision at long distance . 162. Eefraotion of lateral rays 163. Skull, as seen by left eye 164. Skull, as seen by right eye 165. Human auditory apparatus 166. Great sympathetic 167. _Cat, after division of sympathetic in the neck 168. Different kinds of infusoria 169. Experiment on spontaneous generation 170. Trichina spiralis 171. Taenia ..... 172. Cystieercus, retracted . . . • 173. Cystieercus, unfolded 174. Blossom of Convolvulus purpureus 175. Single articulation of Taenia crassicollis 176. Human ovum ... 177. Human ovum, ruptured by pressure 178. Female generative organs of frog 179. Mature frogs' eggs ... 180. Female generative organs of fowl 181. Fowl's egg ... . 182. Uterus and ovaries of the sow . 183. Generative organs of human female 184. Spermatozoa 185. Graafian follicle .... From Sehultze LIST OF ILLUSTRATIONS. XXTll pie. 186. Ovary with Graafian follicle ruptured . 187. Graafian follicle, ruptured and filled with blood 188. Corpus luteum, three weeks after menstruation 189. Corpus luteum, four weeks after menstruation 190. Corpus luteum, nine weeks after menstruation 191. Corpus luteum of pregnancy, at end of second month 192. Corpus luteum of pregnancy, at end of fourth month 193. Corpus luteum of pregnancy, at term . 194. Segmentation of the vitellus 195. Impregnated egg, showing embryonic spot 196. Impregnated egg, showing two layers of blastodermic membrane 197. Impregnated egg, farther advanced 198. Prog's egg, at an early period . 199. Egg of frog, in process of development 200. Egg of frog, farther advanced 201. Tadpole, fully developed 202. Tadpole, changing into frog 203. Perfect frog 204. Egg of fish 205. Young fish, with umbilical vesicle 206. Human embryo, with umbilical vesicle 207. Fecundated egg, showing formation of amnion . 208. Fecundated egg, showing commencement of allantois 209. Fecundated egg, with allantois nearly complete 210. Fecundated egg, with allantois fully formed 211. Egg of fowl, showing area vasoulosa 212. Egg of fowl, showing allantois, amnion, &c. 213. Human ovum, showing formation of chorion 214. Compound villosity of human chorion 215. Extremity of villosity of chorion 216. Human ovum, at end of third month . 217. Uterine mucous membrane 218. Uterine tubules .... 219. Impregnated uterus, showing formation of decidua 220. Impregnated uterus, showing formation of decidua reflexa 221. Impregnated uterus, with decidua reflexa complete 222. Impregnated uterus, showing union of chorion and decidua 223. Pregnant uterus, showing formation of placenta 224. Foetal pig, with membranes 225. Cotyledon of cow's uterus 226. Extremity of foetal tuft, human placenta 227. Foetal tuft of human placenta injected 228. Vertical section of placenta 229. Human ovum, at end of first month 230. Human ovum, at end of third month . 231. Gravid human uterus and contents 232. Muscular fibres of unimpregnated uterus 233. Muscular fibres of human uterus, ten days after parturition 234. Muscular fibres of human uterus, three weeks after parturition 236. Formation of cerebro-spinal axis .... XXVIU LIST OF ILLUSTKATIONS. pia. 236. Formation of cerebro-spinal axis 237. Foetal pig, showing brain and spinal cord 238. FcEtal pig, showing brain and spinal oord 239. Head of foetal pig ... . 240. Brain of adult pig .... 241. Formation of alimentary canal . 242. Foetal pig, showing Umbilical hernia 243. Head of human embryo, at twenty days 244. Head of human embryo, at end of sixth week . 245. Head of human embryo, at end of second month 246. Foetal pig, showing Wolffian bodies 247. Foetal pig, showing first appearance of kidneys 248. Internal organs of generation 249. Internal organs of generation . 250. Formation of tunica vaginalis testis 251. Congenital Inguinal hernia . . 252. Egg of fowl, showing area vasculosa 253. Egg of fifih, showing vitelline circulation 254. Young embryo and its vessels . 255. Embryo and its vessels, farther advanced 256. Arterial system, embryonic form . ' . 257. Arterial system, adult form 258. Early condition of venous system 259. Venous system, farther advanced 260. Continued development of venous system 261. Adult condition of venous system 262. Early form of hepatic circulation 263. Hepatic circulation farther advanced 264. Hepatic circulation, during latter part of foetal life 265. Adult form of hepatic circulation 266. Foetal heart . 267. Foetal heart 268. Foetal heart 269. Foetal heart 270. Heart of infant 271. Heart ef human foetus, showing Eustachian valve 272. Circulation through the foetal heart 273. Adult circulation through the heart From Longet PASE 638 638 639 639 639 645 646 652 652 653 655 657 657 659 660 661 666 666 667 668 670 670 672 673 673 674 675 676 676 677 678 678 678 679 679 681 682 685 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 pheno- 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 resemble 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, then, 4 ( 49 ) 50 INTRODUCTION. 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 differences, both of structure and function, which require a special study. Thus, the physiology of fishes is not exactly/* the same with that of reptiles, nor the physiology of birds with that of quadrupeds. 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; beside which, there are many physiological questions that require for their solution experiments and observations, which can only 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 living 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, when plaped under the requisite , external conditions, present certain active phenomena, which are characteristic of them, and by which they may be recognized. INTHODUCTIOX. 51 Thus lime, dissolved in water, if brought into contact with car- bonic acid, alters its condition, and takes part in the formation 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 tp 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 body, 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, &o., 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, from4he 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 52 INTRODUCTIOlSr. interior of the body, which 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 body, 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 b§ determined by the means of investigation which are at his disposal. Consequently, all efforts to- solve them will 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 occur 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 profitably 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, 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. 53 As, for example, wlieii he 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 prac- tical 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, form any original principlefe, 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 54 INTRODUCTION. the latter can never he in the least degree inferred from those of the former, but must be studied by 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, infer its presence in the body or otherwise, nor in what quantities nor in what situa-r 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 anatomy, 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 ascertain 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 ordi- nary 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 character- istic 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- Fig. 24. scppe (Fig. 24), is seen to consist of abutidant fepithe- 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, and becomes slightly opalescent by boil- B«CCAL AND OLANDUI.AEEpiTH£r.HJM, with i^g- ^^^ ^J t^S addltlOn Of Granular Matter and Oil-globules; depoBited as sedi- nitric acid. ^Icohol in ex- meat from humaa saliva. . .... cess causes the precipitation of abundant whitish flocculi. According to Bidder and Schmidt,' the composition of saliva is as follows : — Composition op Saliv*. Water 995.16 Organic matter 1.34 Sulpho-oyanide of potassium 0.06 Phosphates of soda, lime, and magnesia .98 Chlorides of sodium and potassium .84 Mixture of epithelium 1.62 1000.00 The organic substance present in the saliva has been occasionally known by the name of jptyaline. It is coagulable by alcohel, 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 '• Verdauungssaefte und StoflFwechsel; Leipzig, 1852. SALIVA. 125 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, ■whicli 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 degitee of viscidity than that from the submaxillary. The mucous secre- tion of the follicles of the mouth, whicli 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 wben received in a glass vessel, but adheres strongly to the surface of the glass. We bave obtained the parotid saliva of the human subject in a state of purity by introducing directly into the orifice of Steno's duct a silver canula ■ j'j to jV oi an inch in diameter. The other extremity of the canula projects from the mouth, between the lips, and the saliva is collected 'as it runs from the open orifice. This method gives results much more valuable than observations made on salivary fistulse and the like, since the secretion is obtained under perfectly healthy conditions, and unmixed with other animal fluids. 126 DIGESTION. The result of many different observations, conducted in this way, is that the human parotid saliva, like that of the dog, is colorless, watery, and distinctly alkaline in reaction. It difeers from the mixed saliva of the mouth, in being perfectly clear, without any turbidity or opalescence. Its flow is scanty while the cheeks and jaws remain at rest • but as soon as the movements of mastication are excited by the introduction of food, it runs in much greater arbundance. We have collected, in this way, from the parotid duct of one side only, in a healthy man, 480 grains of saliva in the course of twenty minutes ; and in seven successive observations, made on dififerent days, comprising in all three hours and nine minutes, we have collected a little over 3000 grains. The parotid saliva obtained in this way has been analyzed by Mr. Maurice Perkins, Assistant to the Professor of Chemistry* in the Qollege of Physicians and Surgeons, with the following result : — • Composition op Hpman Parotid Saliva. Water 983.300 , Organic matter precipitable by alcohol . . . . . 7.352 Substance destructible by heat, but not precipitated by alcohol or acids 4.81 Sulpho-cyanide of sodium 0.33 = Phosphate of lime ......... 0,24 Chloride of potassium 0.90 Chloride of sodium and carbonate of soda . . . . 3.06 1000.00 Mr. Perkins found, in accordance with our own observations, . that the fresh parotid saliva, when treated with perchloride of iron, showed no evidences of sulpho-cyanogen ; but after the organic mat- ters had been precipitated by alcohol, the filtered fluid was found to contain an appreciable quantity of the sulpho-cyanide. The organic matter in the parotid saliva is in rather large quan- tity as compared with the mineral ingredients. It is precipitable by alcohol, by a boiljng temperature, by nitric acid, find by sulphate of soda in excess, but not by an acidulated solution of ferrocyanide of potassium. It bears some resemblance, accordingly, to albumen, but yet is not precisely identical with that substance. The parotid saliva also differs from the mixed saliva of the mouth in containing some substance which ma'sks the reaction of sulpho- cyanogen. For if the parotid saliva and that from the mouth be drawn from the same person within the same hour, the addition of perchloride of iron will produce a distinct red color in the latter, while no such change takes place in the former. And yet the parotid SALIVA. 127 saliva contains sulpho-cyanogen, -which may be detected, as we have already seen, after the organic matters have been precipita,ted by alcohol. 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 versd. In these animals mastication is said to be linilateral, 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 the jaw may be reversed many times during the course of a meal. By establishing a salivary fistula simultaneously on ieach 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. • We have observed a similar alternation in the flow of parotid saliva in the human subject, when the mastication is changed from side to side. In an experiment of this kind, the tube being inserted into the parotid duct of the left side, the quantity of saliva dis- charged during twenty minutes, while mastication was performed mainly on the opposite side of the mouth, was 127.5 grains; while the quantity during the same period, mastication being on the same side of the mouth, was 374.4 grains — being nearly three times as much in the latter case as in the former. 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 fhe presence of food, it becomes somewhat difficult 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 • Traits de Pliysiologie ComparSe, Paris, 1854, p. 468. ' Simon's Chemistry of Mau. Fhila. ed., 1846, p. 295. 128 DIGESTION. duct ia the human subject ; from which it was supposed that the total amount secreted bj 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 regarded 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 un- irritating substance, as glass beads, or the like; and during the mastication of food, the saliva is poured out in very much greater abundance. 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 ^otal quantity • of saliva, based on the amount secreted in this intervals of mastication, would be a very imperfect one. We may make a tolerably accurate calculation, however, by ascertaifting 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 purpose, 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 ^ ] 2232 " Total quantity in twenty-four hours = 20164 grains ; or rather less than 3 pounds avoirdupois. ' Op. cit., p. 14. SALIVA. 129 The most important question, connected 'witli this subject, relates to the function of the saliva in the digestive process. A very remark- able 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 kef)t 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 boMed 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 temperaturei of 100° P., will yield traces of sugar at the end of half a minute. The rapidity however, with which this action is mani- fested, varies very much, as was formerly noticed by Lehmann, 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 fiftpen 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. It was supposed, when this property of converting starch into sugar was first discovered in the saliva, that it constituted the true physiological action of this secretion, and that the function of the saliva was, id reallity, the digestion and liquefaction of starchy substances. It was very soon noticed, however, by the French ' Chimie appliquee i, la Physiologie et h, la Therapentique, Paris, 1856, p. 43. 9 130 DIGESTIOU'. 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 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. «[f a dog, with a gastric fistula, be fed with a mixture of meat and boiled starch, and portions of the fluid con- tents of the stomach 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 apy time. Somej;imes 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 Schmidt have also concluded, from subsequent investigations,' that the first experiments performed under their direction by Jacubowitsch were erroneous ; and it is now acknowledged by them, as well as by the French 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, starchy 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 both 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, ' Op oit., p. 26. SALIVA. 131 therefore, of saliva on starch, thougli a curious and interesting pro- perty. Las 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- 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 difl&cult and laborious, not only for dry food, like bread, but even for that of a tolerably moist consistency, like fresh meat. The animals also became very thirsty, and jfere 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 I ' Op. cit., p. 3. * Lesons de Physlologie Expfirimentale, Paris', 1856, p. 146., 132 DIGESTION. ■ retarded. The alimentary masses passed down the oesophagus at longer intervals, and their interior was no longer moist and pastj, 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 appropriate vessels and again weighed. The difi'erence 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 Saliva absorbed. For 100 parts of hay 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 ijiuoh mastipation, 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 preliqiinary preparation ^ ' Comptes Eendus, vol. xxi. p. 362. . GASTRIC JUICE, AND STOMACH DIGESTION. 133 is accomplisljed. 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 stoinach. Gastric Juice, and Stomach DiGBSTiON.-r-The mucous mem- branp 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 wha* is required for the 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 polygonal interspaces, each of which is icncircled by a capillajy network. In the pyloric portion (Fig. 26), the eminences are more ^ig. 25. Fig. 26. Fig. 25. Free surfiico of Gastric Mucous MemsrAke, viewed from above; from Pig's Sto- macli, Cardiac portion. Magnified 70 diameters. Fig. 26. Free surface of Gastric Mucous Membrane, viewed in %'erticaL section; from Pig's Stomach, Pyloric portion. Magnified 420 diameters. or less pointed and conical *in form, and generally flattened from side to side. They contain each a capillary bloodvessel, which re- 134 DIGESTION. turns 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 ^'S- ^''- parts of the stomach. In the pyloric portion (Fig. 27), they are nearly straight, simple- tubules, ^{jj of an inch in diameter," easily separated from each other, lined with glandular epithelium, and ter- minating in blind extremities at the under surface of the mucous membrane. They are sometimes slightly branched, > or provided with one or two rounded diverticula, a. short distance above their termina- tion. They open on the free surface of the mucous mem- brane, in the interspaces be- tween the projecting folds or villi. Among these tubular glandules there is also found, in the gastric mucous membrane, smother kind of glandular organ, consisting of closed follicles, similar to the soli- MncoDS Membrane OF Pig's Stomach, from Pyloric portion; vertical section; showing gastric tubules, and, at a, a closed follicle. Magnified 70 diapetei's. Fig. 28. Fig. 29, Fig, 28. Gastric Tubules from Pig's Stomach, Pyloric portion, showing their Csecal Extremities. At a, the torn extremity of a tubnle, sho^v^Dg its cavity. Pig. 29. Gastric Tubuj.es from Pig's Stomach; Cardiac portion. At a, a large tubule dividing into two small ones. b. Portion of a tubule, been endwise, c. Its central cavity. GASTRIC JUICE, AND STOMACH DIGESTION. 135 tary 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 caecal 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 membi^ane they become branched and conside- rably reduced in size. From the point where the branching takes place to their termination below, 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 membrane with an abundant supply of blood, for the purposes both 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 Eeaumur and Spallanzani, who showed by various methods that the reduction and liquefaction of the food in the stomach could not be owing to 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 afterward withdrawn, the fluids which they had absorbed being then 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, the result of an accidental 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 136 DIGESTION. 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 diarfteter, 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 extraclied foi* 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 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 efiect of various kinds of stimulus on the secretion of the stomach, the rapidity with whicb the process of digestion takes place, the com- parative digestibility of various kinds of food, &c. &o. Since Dr. Beaumont's time it has been ascertained that similar gastric fistulas 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 gresat 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 ' Experiments and Observations upon the Gastric Juice. Boston, 1834. GASTRIC JUICE, AND STOMACH DIGESTION. 137 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 away, 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 some respects, indeed, more satisfactory when made upon the lower animals, than upon the human subject; since animals are entirely under the control of the experimenter, and all sources of 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 fistul£e 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. 138 DIGESTIOK. 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 abundatit supply. In other instances, on the con- trary, the introduction of metallic catheters, &c., 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, or eight days. There is at no time, however, under these circumstances, any considerable amount of fluid present in the stomach ; but only just suificient 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 suf&cient 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 §ij to giiss may be collected in the course of fifteen minutes. Afterward 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 io 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 GASTRIC JUICE, AND STOMACH DIGESTION. 139 filtration from accidental impurities. Obtained in this way, the gastric juice is a clear, wa,tery fluid, without any appreciable vis- cidity, very distinctly acid to test paper, of a faint ^mber 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 : — Composition of Gastbio Jpick. Water 975.00 Organic matter 15.00 Lactic acid 4.78 Chloride of sodium 1.70 " " potassium « 1.08 " " calcium 0.20 " " ammonium ........ 0.65 Phosphate of lime 1.48 " " magnesia 0.06 "■ " iron 0.06 1000.00 In place of lactic acid. Bidder and Schmidt 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. Eobin and Yerdeil 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 chlo- rides, 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 > Leijons de Fhjsiologie Exp^rimentale, Paris, 1856, p. 396. 140 DIGESTION. 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 egg. 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 which 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, on somewhat insufficient grounds, as a product of the alteration of the mucus ^f 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 no 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. ^'S- 30- It is precipitated also,, and coagulated, by a boiling tem- perature; and the gastric juice, accordingly, after being boUed, becomes turbid, and loses altogether its power of dissolving alimentary sub- stances. Grastric 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 CoNFERvoiD Vegetable, growing in the Gas- tric Juice of the Dog. Tlie fibres have an average diameter of 1-7000 of an incht GASTRIC JUICE, AND STOMACH UIGJiSTION. 141 common glass-stoppered bottle without developing any putrescent odor. A light deposit generally collects 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 composed of deli- cate 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 accompanied by any putrefactive changes, and the gastric juice retains it's acid reaction and its digestive properties for many months. By experimenting artificially with gastric juice on various ali- mentary substances, such as meat, boiled white of egg, &o., it is found, as Dr. Beaumont formerly. observed, to exert a solvent action on these sub"fetances 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 Dr. Beaumont, and his predeces- sors, the gastric juice is not a universal "solvent for all alimentary substances, but, on the contrary, afiects 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 influ- ence of the warmth and moisture. Solid and semi-solid albuminoid matters, however, are at once attacked and liquefied by the diges- tive fluid. Pieces of coagulated white of egg suspended in it, in a test-tube, are 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 de- posit, which collects at the bottom of the test-tube. "While the disintegrating process is going on, it may almost always be noticed that minute, opa'que 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 bot- tom is probably also composed of some ingredient, not soluble in the gastric juice. If pieces of fresh meat be treated in the same manner, the areolar tissue entering into its composition is first 142 DIGESTION. dissolved, so that the muscular bundles become more distinct, and separate from each otber^ 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 subjected to the same process, the walls of the fat vesicles and the inter- vening areolar tissue, together with the capillary bloodvessels, &c., are dissolved ; while the oily matters are set free from their en- velops, and collect in a white, opaque layer on the surface. In cheese, the casein is dissolved, and the oil which it contains set free. In bread the gluten is digested, and the starch left un- changed. In milk, the casein is first coagulated by contact with the acid gastric fluids, and afterward slowly liquefied, like other , albuminoid substances. The time required for the complete liquefaction ef 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 diges- tible substance. The liquefying process which the food undergoes in the gastric juice is not a simple solution. It is a catalytic transformation, produced in the albuminoid substances by contact with the organic 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 requires to be dis- solved 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, a«id a very high temperature. By its opera- tion the albuminoid matters of the food, whatever may have been their original character, are all, without distinction, converted into a new substance, viz., albuminose. This substance has the general characters belonging to the class of organic matters. It is uncrys- tallizable, and contains nitrogen as an ultimate element. It is pre- cipitated, like albumen, by an excess of alcohol, and by the metallic salts ; but unlike albumen, is not affected by nitric acid or a boil- ing 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 GASTRIC JUICE, AND STOMACH DIQESTION. 143 cheese is entirely converted into albuminose, and dissolved nnder that form. A very considerable portion of raw white of egg, how- ever, dissolves in the gastric juice directly as albumen, ^.nd retains its property of coagulating by heat. Soft-boiled white of egg and raw meat are principally converted 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, 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 timfc 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 84). 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 ffebdomadaire 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 ' American Jonm. Med. Sci., Oct. 1854, p. 319. 2 Nouvelles recherches relatives h Taction du sue gastrique sur lea substances albuminoidea. — Gaz. Hehd. 9 Ftvrier, lS55, p. 103. 144 DIGESTION. of food, already contains some albuminose in solutjon, 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 potassa, 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 quantity, an incomplete reduction of the copper takes place, so that the mixture becomes opaline and cloudy, but still without any weU 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 mutual reaction of starch and iodine. If 3j 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 he 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 GASTBIC JUICE, AND STOMACH DIGESTION. 145 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 peristfiMia 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 stomach the bulb and stem of a thermometer. According to his observations, when the food first passes into the stomach, and the secretion of the gastric juice commences, the muscular coat, which was before quiescent, is excited and begins to contract act- ively. The contraction takes place in such a manner that the food, after entering the cardiac orifice of the stomach, is first carried to the left, into the great pouch of the organ, thence downward and a,long the great curvature to the pyloric portion. At a short distance from the pylorus. Dr. B. often found a circular constriction of the gastric 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 constric- tion was relaxed, and the bulb of the thermometer again released, and carried together with the food along the small curvatui'e of the organ to its cardiac extremity. This circuit was repeated so long as any food 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 ordi- nary 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 carefully in place, or it will fall out by its own weight. But immediately 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, 10 146 DIGESTION. in opening the abdomen, any such energetic or extensive contrac- tions of the stomach, even when full of food, as may he easily excited in thb 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 pro- duce a constant churning movement of the masticated 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 mucous membrane ; so l^at thjg digestive fluid is made to penetrate equally every part of the ali- mentary mass, and the digestion of all its albuminous ingredients goes on simultaneously. This gentle and continuous movement of the stomach is one which cannot be successfully imitated in experi- ments 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 reqjiires 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. Bv examinmg 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 : — KiSD OF Food. Hours. Mikutes. 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 OO Salt beef (boiled) 4 15 Roasted pork 5 15 GASTRIC JOICE, AND STOMACH DIGESTION. 147 The comparative digestibility of different substances varies more or less ia 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 quantity 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 34 pounds, they were able to obtain by separate experiments, con- suming in all 12 hours, one pound and three-quartfers of gastric juice. The total quantity, therefore, for 24 hours, in the same ani- mal, would be 3J pounds ; and, by applying the same calculation 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 deter- mine 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 desiccation, 78 per cent, of its weight. 300 grains of such meat, cut into small pieces, were then digested for ten hours, in Siss 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 quantity 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 3iss of gastric juice, or 33J 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 which 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 148 DIGESTION. 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 circulation of the digestive fluids from the bloodvessels to the ali- mentary canal, and from the alimentary canal back again to the bloodvessels. That this circulation really takes place is proved by the fol- Ipwing 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 sujB&cient to smear over and penetrate the half digested pieces of meat ; and, secondly, in the living animal, gastria (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 ab- sorption 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 albu- minous substances, should not destroy the walls of the 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 Eeauraur 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 thus destroying the organ by which it was itself 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 -pepsine of the gastric juice. "We know that all the or- ganic substances in the living tissues are constantly undergoing, in GASTRIC JUICE, AND STOMACH DIGESTION. 149 the process of mitrition, a series of catalytic changes, which are characteristic 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 putrefective changes are stopped or altogether 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 difiBculty. 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. 150 DIGESTION. 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 stomach, 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 op 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 sinall intestine. As already mentioned, it is only the albuminous matters which are digested in 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 gss 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 sugar cane 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 JUICES, DIGESTION OF SUGAR, ETC. 151 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 late. For it is remarkable how rapidly starqhy substances, if pre- viously disintegrated by boiling, are disposed of in the digestive 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 rafndity with which this passage of the starch into the duodenum takes place varies, to some extent, in different animals, '^'^" ^^* according to the general ac- tivity of the digestive appa- ratus; but it is always a comparatively rapid process, when th^ 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 ac^ coraplished 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 accomplished. 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. Follicles of Liebi^bkitqn, testiue of dog. from Small la- 152 DIGESTION. exert any digestive action on starch. The pancreatic juice, on the other hand, has the property of converting starch into sugar ; but . it is not known whether this fluid be always present in the duode- num. 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 Lieberkiihn and the glands of Brun- ner. The first of these, or Lieberkiihn's follicles (Fig. 31), are the most numerous. They are simple, nearly straight tubules, lined with a continuation of the intestinal epithelium, and somewhat 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 glandulss, as they are sometimes called, are confined to the upper part of the duO' denum, where they exist as a ^'^" ^^' closely set layer, in the deeper portion of the mucous mem- brane, extending, downward a short distance from the pylo- rus. They are composed of a great number of rounded follicles, clustered round' a central excretory duct. Each follicle consists of a delicate membranous wall, lined with glandular epithelium, and covered on its surface with small, distinctly marked nu- clei. The follicles collected around each duct are bound together by a thin layer of areolar tissue, and covered with a plexus of capillary bloodvessels. The intestinal juice, which is the secreted product of the above glandular organs, has been less successfully studied than the other digestive fluids, owing to the difiiculty 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 . Portioa of one of Buunner's Glands, from Humau Intetstine, DnoDESi PANCKEATIO JUICE, ^ND THE DIGESTION OF FAT. 153 into the abdomen, and the external wound closed by sutures. After six 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- lished an intestinal fistula below, from which they extracted the fluids which accumulated in the cavity of the gut. Frora 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 folli- cles 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 tempera- ture of the living body. Pancreatic Juice, and the Digestion of Fat. — The only re- maining ingredients of the food that require digestion are the oily matters. These are not affected, as we have already stated, by con- tact 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. Very soon, however, after its entrance into the intestine, the oily portion of the food loses its characteristic appearance, and is con- verted 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 valvular conniventes, and adhering to the surface of the villi. The digestion of the oil, however, and its conversion 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 154 DIGESTIOjr. these ducts almost invariably open into the intestine at or near the same point, it was for a long time difficult 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 ex- perimenting on some of the lower animals, in which the two ducts open separately. In the rabbit, for example, the biliary duct opens as usual just below the pylorus, while the pancreatic duct com- municates with the intestine some eight or ten inches lower down. Bernard fed. these animals with substances containing oil, or in- jected melted butter into the stomach ; and, on killing them after- ward, found that there was no chyle in the intestine between the opening of the biliary and pancreatic ducts, but that it was abun- dant immediately below the orifice of the latter. Above this point, also, he found the lacteals empty or transparent, while below it they were full of white and opaque chyle. The result of these ex- periments, which have since been confirmed by Prof. Samuel Jack- son, of Philadelphia,' led to the conclasion that the pancreatic fluid is the active agent in the digestion of oily substances ; and an ex- amination of the properties of this secretion, when obtained in a pure state from the living animal, fully confirms 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 eftects 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 re- commences, and continues to exhibit similar fluctuations during ' American Journ. Med. Soi., Oct. 1S54. ■ ■ PANCREATIC JUICE, AND THE DIGESTION OF FAT. 155 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 more than a little over two fluidounces_ and a half during a period of three hours, in a dog weighing not more than forty-five pounds. This is equivalent to about 864 grains per hour ; but as the pancreatic juice in the dog is secreted with freedom only during digestion, and as this J)rocess is in opera- tion not more than twelve hours out of the twenty-four, the entire amount of the secretion for the whole day, in the dog, may be esti- mated at 4,368 grains. This result, applied to a man weighing 140 pounds, would give, as the total daily quantity of the pancreatic juice, about 13,104 grains, or 1,872 pounds 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 PA^•CIlEATIO Juice. Water ... 900.76 Organic matter (pancreatine) 90.38 Chloride of sodium 7.36 Free soda 0.32 Pliosphate of soda 0.45 Sulphate of soda 0.10 Sulphate of potassa O.02 r Lime 54 Combinations of J Magnesia 0.05 I 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 flaid. If 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 re- mains 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 156 UIGESTION. 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 the 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 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 sufier 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, ' Lehmann's Physiological Chemistry. Philada. ed., vol. i, p. 507. PHENOMENA OF INTESTINAL DIGESTION. 157 by the influence of whicli 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 iiitervening 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 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 which 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 ingestion 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 Troramer'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 158 DIGESTION. Fig. 33. fibres, fat vesicles, and oil drops; substances whicli are easily recognizable under the microscope, and which produce a grayish turbidity in the fluid drawn from the fistula. This turbid admixture grows constantly thicker from the second to the tenth or twelfth hour; after which the intestinal ■ fluids become less abundant, and finally disappear almost entirely, as the process of di- gestion comes to an end. The passage of disintegrated, muscular tissue into the intes- tine 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 and adipose, tissue, the stomach contains masses of softened meat, smeared over with gastric juice, and also a moderate quantity of grayish, grumous fluid, with an acid reaction. This fluid contains, muscular fibres, isolated from each other, and more or less dis- integrated, by the action of the gastric juice. (Fig. 33.) The fat vesicles are but little or not at sd\ altered in the stomach, and there are only a few free oil globules to be Contents of Stomach during Digestion OF Meat, from the Dog.— n. Fat Vesicle, filled with opaque, solid, granular fat. &, b. Bits of pUrtially dieiiategrated muscular fibre, c. Oil globules. Fig. 34. From Duodknum of Dog, during Diges- tion OP Meat. — fi. Fat Vesicle, with its contents diminishing. The vesicle is beginning to shrivel and' the fat breaking up. h, b. Disintegrated muscular SCeU floating in the mixCd iibre. CO. Oil Globules. fluids. Contained in the cavity of the organ. In the duodenum the muscular fibres are further* disintegrated. (Fig. 34.) They become very much broken up, pale and transparent, but can still be recognized by the granular mark- . ings and striations which are characteristic of them. The fat vesi-- PHENOMENA OF INTESTINAL DIGESTION. 159 From Middlk op Small iNTKariHE.- Fat vesicles, nearly emptied of their contenta. Fig. 36. cles also begin to become altered in the duodenum. The solid granular fat of beef, and similat kinds of meat, becomes liquefied and emulsioned ; and appears under the form of free oil ^'^' ^^' drops and fatty molecules; while the fat vesicle itself is partially emptied, and becomes more or less collapsed and shrivelled. In the middle and lower parts of the intes- tine (Figs, 35 and 36) these changes continue. The mus- cular fibres become constantly more and more disintegrated, and a large quantify of granu- lar debris is produced, which is at last also dissolved. The fat also progressively disap- pears, and the vesicles may be seen in the lower part of the intestine, entirely collapsed and empty. In this way the digestion of the different ingredients of the food goes on in a continu- ous manner, from the stomach throughout the entire length of the small intestine. At the same time, it results in the production of three different substances, viz : 1st. Albumi- nose, produced by the action of the gastric juice on the albuminoid matters; 2d. An oily emulsion, produced 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 as the mingled ingredients of the intestinal contents pass successively downward, through the duodenum, jejunum, and ileum, the products of digestion, together with the digestive secretions themselves, are gradually absorbed, Fhom last quarter of Small Intestine. — a, a. Fat vesicles, quite empty and shrivelled. 160 DIGESTION. one after anotlier, by tlie vessels of tlie mucous membrane, and carried away by the current of the circulation. The Large Intestine and its Contents. — Throughout the small intestine, as we have just seen, the secretions are intended exclusively or mainly to act upon the food, to liquefy or disinte- grate it, and to prepare it for absorption. But below the situation of the ileo-caecal valve, and throughout the large intestine, the con- tents of the alimentary canal exhibit a diflferent appearance, and are distinct in their color, odor, and consistency. This portion of the intestinal contents, or the feces, are not composed, for the most part, of the undigested remains of the food, but consist principally of animal substances discharged into the intestine by excretion. These substances have not all been fully investigated ; for although they are undoubtedly of great importance in regard to the preser- vation of health, yet the peculiar manner in which they are dis- charged by the mucous membrane and united with each other in the feces has interfered, to a great extent, with a thorough investi- gation of their physiological characters. Those which have been most fully examined are the following : — Excretine. — This was discovered and described by Dr. W. Mar- cet,' as the most characteristic ingredient in the contents of the large intestine. It is a slightly alkaline, crystallizable substance, insoluble in water, but soluble in ether and hot alcohol. It crys- tallizes in radiated groups of four-sided prismatic needles. It fuses at 204° F., and burns at a higher temperature. It is non-nitrogen- ous, and consists of carbon, hydrogen, oxygen, and sulphur, in the following proportions : — C78 H78 O2 S. It is thought to be present mostly in a free state, but partly in union with certain organic acids, as a saline compound. Stercorine. — This substance was found to be an ingredient of the human feces by Prof. A. Flint, Jr.^ It is soluble in ether and boiling alcohol, and, like excretine, crystallizes in the form of radiating needles, but fuses at a much lower temperature. It is regarded by its discoverer as produced, by transformation, from cholesterine, one of the ingredients of the bile. Beside these substances, the feces contain a certain amount of ' American Journal of the Medical Sciences, January, 1855, and January, 1858. » Ibid., October, 1862. THE LARGE INTESTINE AND ITS CONTENTS. 161 fat, fatty acids, and the remnants of undigested food. Vegetable cells and fibres may be detected, and some debris of the disin- tegrated muscular fibres may almost always be found after a meal composed of animal and vegetable substances. But little absorp- tion, accordingly, takes place in the large intestine. Its of&ce is mainly confined to the separation and discharge of certain excre- mentitious substances. 11 162 AB30BPTI0K. CHAPTEE VII. ABSORPTION. . Beside the glands of Brunner and the follicles of LieberkHhn, already described, there are, in the inner part of the walls of the intestine, certain glandular- ^'^' ^^' looking bodies vrhich are termed "glandulae.solitariaB," and "glandulaa agminatse." The glandulae solitarisB are globular or ovoid bodies, . about one-thirtieth of an inch in diameter, situated partly in and partly beneath the in- testinal 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- Onr of the closed Follicles of Pkyeb's , . Patches, from Small Intestine of Pig. Miigniaod sistS of minutC Cellular bodicS, 50 diameter-. Fii. 38. Glakdoi.^ AilMivAT.B, fom Small Inteatine if Vii. Maii'uifled 20 dlamuters. imbedded in a homogeneous substance. The inclosed mass is. penetrated by capillary bloodvessels, which pass in 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 vesiicle, being pulpy and vascular, as already de- scribed, are not to be regarded as a secretion, but as consti- tuting a kind of solid gland- ABSOBPTIOX. 163 Fig. 39. tissue. The glandulse agminatse (Fig. 38), or " Peyer'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 mesenteric 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. 39), 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 epith^ial layer. They 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 blind extremity, or by an irregular plexus, passes, in a straight or J:IXTUBUlT]f UF iNTJJSTlNAr. Villus, from the Dog.— «. Liiyer o( epithelium, b. Blood veitBel. c. Lacteal vessel. 164 ABSORPTION. somewhat wavy line, toward the base of the villus, and then be- comes 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-flu-id mass cont^iiied in the intestinal cavity, as the roots of a tree penetrate the soil; and they imbibe the liquefied portions of the food, with a rapi(Jj.ty 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 ma,y 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 (|iameter, 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 relaxatipn, 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 snfall intestine, there remains only .the Undigestible portion of the food, together with the refuse of the intestinal "secretions. These ABSORPTION. 165 pass throTigh the ileo-csecal 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 the blood of the portal vein, but that absorption takes place more rapidly and abundantly by the bloodvessels than by the lacteals. The most decisive of these experiments were those performed by Panizza on the abdominal circulation.' This ob- server 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), which Fiff. 40., Panizza's Expekiment. — 'in. iDtestiud. ft. Point of ligature of mesenterie vein. c. Opening in intestiue for introduction of poison, d. , Opening i a meHBUteric vein behind the ligature. he incliided between two ligatures. A ligature was then placed (at b) upon the mesenteric vein receiving the blood from this portion of intestine; aud, in order that the circulation might not be inter- rupted, an opening was inade (at d) in the vein behind the ligature, so that the blood brought by the mesenteric artery, after circulating ' In Matteucci'd Lecjtures on the Pliysical Phenomena of Living Beings, Pereira'a edition, p. 83. . 166 ABSORPTION. « in the intestinal capillaries, passed out at the opening, and was collected in a vessel for examination. Hydrocyanic acid was then introduced into the" intestine by an opening at c, and almost imme- diately afterward its presence was detected in the venous blood flowing from the orifice at d. The animal, however, was not poi- . soned, since the acid was prevetited from gaining an entrance into the general circulation by the ligature at b. Panizza afterward varied this experiment in the following man- ner : Instead of tying the mesenteric vein, he simply compressed it. Then, hydrocyanic acid being introduced into the intestine, as above, no effect was produced on the animal, so long as compressioii was maintained upon the vein. But as soon as the blood was allowed to pass again through the v«ssels, 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. Hydro- cyanic acid now being introduced into the intestine, 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 aliment- ary matters indiscriminately. It is one of the most important of the facts which have been, established by modern researches on digestion that the 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 both 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, however, 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 J^he condition of ordinary albumen, and probably also partly into that of fibrin ; while the sugar rapidly .becomes decomposed, and loses its characteristic properties; so ABSORPTION. 167 Fiff. 41. 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, chti,e trom commf.ncemkxt op thoe^cio ^ • • ^ Dpct, from the Dog.— The molecales vary in size after a meal containing fatty from i-io,oooth of an inch downward, ingredients, the lacteals may be seen as white, opaque vessels, distended with milky chyle, pass- ing through the mesentery, and converging from its intestinal bor- der 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 discharged, 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 168 ABSORPTION. throughout tbe body. The walls of these vessels are thinner and more transparent than those of the arteries and veins, tad they are consequently less easily detected by ordinary dissection. They originate in the tissues of the above-mentioned parts by an irregular plexus. They pass from the 'extremities toward the trunk, converging and uniting with each other like the veins, their principal branches taking usually the same di- rection with the nerves and bloodvessels, and passing, at various points in their course> through certain glandular bo- dies, the "lymphatic" or "ab- sorbent" glands. The lym- phatic glands, among which are included the mesenteric glands, consist of an external layer of fibrous tissue and a contained pulp or parenchy- ma. The investing layer of fibrous tissue sends off thin septa or laminae from its in- ternal surface, which pene- trate 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 rounded spaces or alveoli. These alveoli are not completely isolated, but communicate 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 Kdlliker, like the solitary and agmiiiated glands of the intestine, by a fine network of capillary bloodvessels. The solitary and agminated glands of thfe intestine are, therefore, closely analo- gous 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 '■■■a. Lacteals, Thoracic Duct^ &c. — a.. Intes- tine, h. Vena cava inferior. c, c. Bight and left Bubclavian veins, e/. Point of opening of thoracic duct into left suhclavian. ABSOBPTIOX. 169 glands, is not precisely understood. Each lymphatic vessel, as it enters the gland, breaks up injo 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 farther side of the gland ; but the exact mode of communication between the two has not been definitely ascertained. 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. So far as regards its composition,, it is known to contain, beside water and saline matters, a small quantity of fibrin and albumen. Its ingredients are evidently derived from the metamorphosis 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, how- ever, with regard to the precise nature of their character 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 w'hich 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. 43.) 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 aftqr 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 170 ABSORPTION. digestion, as soon as they have been disintegrated and emulsione^ by the potion of the intestinal fluids. As digestion proceeds, they accumulate in larger quantity, and gradually fill the whole lacteal ■ Fig. 43. liACTEALS AND LyjIFHATICS. 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 oijy molecules and granules, between the orifice of the thoracic duct and the right side of the heart. While passing through the pulmonary circula- tion, however, they disappear. Precisely what becomes of them, or what particular chemical changes they undergo, is not certainly ABSOKPTION. 171 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 ife 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 dif&oulty has long been experienced in accounting for the ab- sorption of fat from the intestine, owing to its being considered as a non-endosmotic substance ; that is, as incapable, in its free or undis- solved 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 endos- mometer, and pure water on the other, the water will readily pene- trate the substance of the membrane, while the oily particles cannot be made to pass, even under a high pressure. Though this be true, however, fdr pure water, it is not true for slightly alkaline fluids, like the serum of the blood and the lymph. This has been de- monstrated by the experiments of . Matteucci, in which he made an emulsion with an alkaline fluid containing 43 parts per thou- sand of caustic potassa. Such a solution has no perceptible alkaline taete, and its action on reddened litmus paper is about equal to that of the lymph and chyle. If this emulsion were placed in an < Lefons dH Phjsiologie GxpSrimentale. Paris, 1856, p. 325. 172 ABSORPTION. Intestinal Epithelium fasting. from the Dog, while endosmometer, together with a watery alkaline solution of similar strength, it was found that the oily particles penetrated through the animal membrane without ^'S- ^^ 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 endosmosia 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 directly into and through the substance of the villi; ^" ' " The epithelial cells covering the external surface of the villus are the first active agents in tbis absorption. In the intervals of digestion (Fig. 44) these cells are but slightly granular and nearly trans- parent in appearance. But if examined during the diges;- tion and absorption of fat (Fig. 45), their substance ia seen to be crowded with oily particles, which they hav« taken up from the intestinal cavity by absorption. The oily matter then passes q/a.- ward, penetrating deeper and d;eeper into the substance of the villus, until it is at last received by the capillary vessels and lacteals in its centre. I STK3TISAL Epithelium; the digestioa of fat. from tlie Dog, dnring ABSOBPTION. 173 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 peculiar appearance known as that of " chylous" or " milky" blood. After the separation of the clot, the serum presents a turbid appearance ; 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 occasionally observed in the human subject, particularly in bleeding for apoplectic attacks occurring after a full meal, and has been mistaken, in some instances, for a morbid phenomenon. It is, however, a perfectly natural one, and depends simply on the rapid absorption, 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, ac- cording to the amount of oleaginous matters contained in the food. • When digestion is terminated, and the fat ceases to be introduced in unusual quantity into the circulation, its transformation and decomposition continuing to take place in the blood, it disappears gradually from the veins, arteries, and capillaries of the general 174 ABSOBPTION. 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. Thus they 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. TUB BILE. 175 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 : — 176 THE BILE. CoMPiisiTios or Ox Bile. Water 888.00 Glyko-oholate of soda" 1 90 00 Tauro-cholate " " ' Biliverdine j Fats ! 13.42 Oleates, margarates, and stearates ofsoda and potassa . i Cholesterin , ^ J Chloride of sodium 1 Phosphate of soda ■ | " " lime } 15.24 " " magnefiia Carbonates of soda and potassa J Mucus of the gall-bladder .1.34 1000.00 BiLiVEBDiNE. — Of the above mentioned ingredients, biliverdine is peculiar to the bile, and therefore importafnt; 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. Cholestebin;(C25H220). — 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 watel". It is not saponifiable, however, by the action of 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 ingredientiin most biliary calculi ; and is found also in different re^ons 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. 177 Fig. 46. the subjacent crystals show; very distinctly through. the substance of those which are placed above. Cholesterin is not formed in the 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. It has> also been found, by Dr. W. Marcet,' to exist, in considerable abund- ance, in the tissue of the spleen. From all these tis- sues it is absorbed by the blood, then conveyed to the liver, and discharged with the bile. Cholestebik, from an EaCyHted Tumor. This fact has been fully confirmed by the researches of Prof. A. Flint, Jr.,' who has found that there is nearly one-quarter part more cholesterin in the blood of the jugular vein, returning from the brain, tha,n in that of the carotid artery, before its passage through that organ ; and that, on the other hand, the blood of the hepatic artery, as well as that of the portal vein, loses cholesterin in passing through the liver, so that but a small quantity can be found in the blood t>t the hepatic vein. The cholesterin, however, after #being poured into the iiitestine with the bile, is decomposed or transformed into some other sub- stance, since it is nat discharged with the feces.^ Its decomposition is probably effected by the contact of the intestinal fluids. - BiLiABT Salts.— rBy far the most important and charaeteristic ingredients of this secretion are the two saline substances mentioned above as the ghfho-cholate and tauro-cholate of. soda. These sub- stances were first discovered by Strecker, in 1848, in the bile of the ox. They are both freely soluble in water and in alcohol, but in- soluble in ether. One of theni, the tauro-cholate, has the property, I In American Joum. Med. Sci., January, 185S. ' American Joum. Med. Sci., October, 1862. » Trof. A. 'Flint, Jr., iu Ala. Journ. Med. Sci., Oct. 1862. 12 178 TEB BILE. •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 following manner : — The bile is first evaporated to dryness by the water-bath. The dnr residue is then pulverized and treated with. absolute alcohol, in the proportion of at least 3j 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-oholate of soda, the coloring matter and more or less of the fats originally, present in the bile. On the addition of a small qua,ntity 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 the fflyko-cholate and tauro-cholate of soda, thrown down under the form of heavy flakes and granules, part of which subside to^ Fig. 47. Fig. 49. Ox-bile, extracted with abnolute alcohol and precipitated with ether. Gi.TKO-rnoLATE OF Soda prom Ox-bile, after two dajm' crystalUialion. At the lower part of the figure the crystalB are melting into drops, from the eraporatton of the ether and absorption of moisture. .the bottom of the test-tube, while part remain fo'r a time in suspen- sion. Gradually these flakes and granules., unite 'wjth each other and fuse together into clear, brownish-yellow> oily,, or resinous^ TH£ BILE. 179 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 di;ops. The deposit is semi-fluid in consistency, and sticky, like Canada balsam or half- melted 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 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 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 crystallization in the test-tube goes on after the first day, and the crystals in- crease 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-oholate, however, is uncrystallizable, and re- mains in an amorphous con- dition. If a portion of the deposit be now removed and Jig. 49. Gltko-ch TAUEO-CHOL ATE OP examined by the microscope, soda, prom ox-bub, after hx days* crystamzii- ... ..1 J j1 i. ^ e tion. The glyko-cholate is crystallized: the tanro- It is seen that the crystals of ^^„,,^ j. ,/j^i, ^^p,. glyko-cholate of soda have increased considerably in thickness (Fig. 49), so that their trans- verse diameter may be readily estimated. The uncrystallizable taui-o-cholate appears under the form of circular drops, varying 180 ' THE BILE. considerably in size, clear, transparent, strongly refractive, and bounded by a dark, well-defined outline. These drops are not to be distinguished, by any of their optical properties, from oil-globuhs, as they usually appear under tbe 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 off and distilled water added, the deposit dissolves, rapidly and completely, with a more or less distinct yellowish color, according 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 the addi- tion oi acetate of lead, the glyko-cholate of soda is decomposed, and precipitates as a glyko-cholate of lead. The precipitate, sepa- rated by 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 subacetate of lead, which precipitates a tauro-cholate of lead. This is separated by ■filtranon, 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 siibsequent 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 : — Glyko-cholate of soda (NaOjOjjH^jNO,,) crystallizes, when precipi- tated by ether frSm 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-eholic THE BILE. 181 acid (C^jjHjj!N"0„,HO). This acid is crystallizable and contains nitro- gen, as shown by the above formula, which is that given by Leh- mann. If boiled for a long time with a dilute solution of potassa, glyko-cholic acid is decomposed with the production of two new substances; the first a non-nitrogenous acid body, cholic acid (O^gHjgOgjHO) ; the second a nitrogenous neutral body, glycine (O^HjNOJ. 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 n(Jt exist origfnally in the glyko-cholic acid, but are rather new combinations of its elements, produced by long boiling, in contact with potassa 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 gfyko-cholic acid. Tauro-cholate of soda (NaO,CjjH^S20„) is also a very abundant ingredient of the bile. It is said by Eobin and Verdeil' that it is 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-choUc 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 potassa, 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 add; and the second a new nitrogenous neutral body, viz., taurine (C^HjNSjOj). The same remark holds good with regard t© 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 ' Chimie Anatomiqne et Physiologiqne, vol. ii. p. 473. * Pliysiologicai Chemistry, Phil, ed., vol. i. p. 209. » Lehmann's Physiological Chemistry, Phil, ed., vol. i. p. 47G. 182 THE BILK. Fig. 50. 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-oholate 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 sepretions of difierent species of animals, Strecker found so great a resemblance between them, that he was disposed to rega,rd 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 qtfentity in which the two were present. The only exception to this was supposed to be pig's bile, in which Strecker found a peculiar organic acid, the "hyo-cholic" or " hyo-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 alco- hol, 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 present 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 crystallizable, 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 crystallizatioii is fully estab- D * s B T I. B, extract- ed withaljsoluteaylcohol and precipitated with ether. * THE BILE. 183 llshed. But it is more particularly in their reaction with the salts of lead that the difference between these substances becomes mani- fest. For while the crystallizable substance of ox-bile is precipi- tated 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 slighest turbidity. If subacetate of lead be then added in excess, a copious precipitate falls, composed of both the crystallizable and uncrystallizable substances. If tlfe lead pre- cipitate be then separated by filtration, washed, and decomposed, as above described, by carbonate of soda, the watery solution 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 precipitated by ether ; when the ether precipitate crystallizes partially after a time as in fresh bile. Both the biliary matters of dog's bile are therefore precipitable by subacetate of lead, but neither of them by the acetate. Instead of calling them, consequently, glyko-cholate and tauro chelate of soda, we shall speak of them simply as the " crys- talline" and " resinous" biliary substances. In cat's bile, the biliary substances act very much as in dog's bile. The ether precipitate of the alcolfclic solution contains here also a crys- talline 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 substance, but the ether precipitate is altogether resinous in appearance. Notwith- standing 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 precipitate Human Bi>LE, ex- tracted with ahfielnte alcohol and precipitated bj ether. 18i THK BILE. 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 presence 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 subacetate it gives no precipitate after filtration by the acetate. The entire biliary in^edients, therefore, of human bile are precipitated by both or either of the salts of lead. Different kinds of bile vary also in other respects ; as, for ex- ample, 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 alco- holic solution of dried ox-bile, furthermore, does not precipitate at all on the addition of water ; while that of human 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 fob 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 by 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 preci- pitate which takes a bright grass-green hue. Tincture of iodine produces the same change of color, when added in small quantity ; and probably there are various other substances which would have the same effect. It is by this test that the bile has so often been recognized in the urine, serous effusions, the solid tissues, &c., in TESTS FOR BILE. 185 cases o^ jaundice. But it is very insufficient for anything . like accurate investigation, since t^e 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 biliverdine. On the other hand, if the coloring matter be ab- sent, the biliary substances themselves cannot be detected by it. For if the biliary substances of dog'« bile be precipitated by ether from an alcoholic solution, dissolved in water 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. Pettenkofer'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 th^ 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 prop'ortion 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 passes 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 186 THE BILE. or twenty-five minutes. If foreign matters, again, not of a biliary nature, be also present, tbey 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 all chance of deception from this source. "When sulphuric acid is mixed with a watery solution containing cane sugar, after it has been added i«i 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 *ngy 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 TESTS FOB BILE. 187 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. Yarious 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 bile, and may therefore give rise to mistakes." Some fatty sub^ stances and volatile oils (oleine, 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 substances of organic origin. No other substance is, in point .of fact, liable to be met with in the intestinal fluids or fhe blood, which would simulate the reactions of the biliary matters. We have found that the fatty matters of the chyle, taken from the tho- racic 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, empyren- matic odor is developed- at the same time ; but we do not get the l^te 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 preseace 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. ' Op. cit., vol. ii.^ p. 46S. 188 THE BILE. Yabiatioits and Functions of Bile. — ^Witk 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 performed 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 open- ing 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, and its quantity accurately determined. Bach observation usually occupied about two hoUrs, during which period the temporary fluctuations occasionally observ- able in the quantity of bile discharged were mutually cco-rected, so far as the entire result was concerned. The animal was then killed, weighed, and carefullf examined, in order to make sure that the biliary duct had been securely tied, and that no inflammatory alter- ation had taken place in the p,bdominal 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 calculated from a comparison of the above results ; and the quantity of its solid ingi-edients was also ascer- tained in^each instance by evaporating 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 twenty -four hours • Fresh Bile. Dbt Residue. In the cat 102 grains. 5.712 grains. •'dog 140 " 6.916 " " . sheep .• 178 " 9.408 " - " rabbit 9.')8 " 17.290 " Since, in\he 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. ' Verdau:igssaefte und StoiTwechBel. Leipzig, 1852. VARIATIONS AND FUNCTIONS OF BILE. 189 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 jiiic% and discharged only during the digestive process. In order to determine this point, we have performed the following series of experiments on dogs. The aniipala 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 con- tents 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 intestinal con- tents always presented some peculiarities of appearance when treated with alcohol and ether, owing probably to the presence of other substances than the bile; but they ailways gave evidence of the presence of biliary matters as well. The biliary sub- stances could almost always be recognized by the mi- croscope in the eth«r preci- pitate of the alcoholic solu- tion ; 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 bun- dles of slender, radiating, slightly curved or wavy, needle - shaped crystals. These substances, dissolved in water, gave a purple color with sugar and sulphuric acid. These experiments 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. Cbtstait.ike akj) EESisniTs BiiiART Srn- BTAXCES; fromSmall Intestine of Dog, after two days ftstlng. 190 THE BILE. ia all these instances, bile was present in the small intestine. It is, therefore, plainly not an intermittent secretion, nor one which is concerned exclusively in the digestive process ; but its secretion is constant, and it continues to be discharged into the intestine for many days after the animal has been deprived of food. The next point of importance to be examined relates to the time, after feeding at which the bile 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, by tying the common bile-duct, and then opening the fundus of the gall-bladfler, 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 diflferent periods, they came to the conclusion that the flow *^8' 53. Qf l^jlg 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 fifteea , hours. Other observers, how- ever, have obtained different results. Arnold;' for example, found the quantity to be largest soon after meals, decreasing • again after the fourth hour. Kolliker and Muller,' again, found it largest between the sixth and eighth hours. Bidder and Schmidt's experiments, in- deed, 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, bear- ing on this point, were performed on dogs, by making a permanent duodenal fistula, on the same plan that gastric fistulse have so often DnoDBNAi. Fistula.— «. Stomach. 6. Dao- deaum. c^ c^ c. Pancreas; its two dacts are seen opening into tlie duodenum, one near the orifice of the biliary duct, <2, the other a Khort distance lower down. e. Silver tube paRsin^ through the abdomiual walls and openiu(^ into the duodenum. > lu Am, Joarn. Med. Sci., April, 1856. « Ibid., April, 1857. VARIATIONS AND PDNCTIONS OF BILK. 191 been established for the examination of the gastric juice. (Fig. 53.) An ineisioa was made through the abdqminal 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 in- cision, and a silver tube, armed at each end with a narrow projecting collar or flange, inserted 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 parietes, the parts secured by sutures, and the wound allowed to heal. After cicatrization was complete, and the animal had entirely recovered his healthy condition and appetite, the intestinal fluids were drawn off 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 wheii 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. 192 THE BILE, • Time after Quantity of fluid Dry residue Quantity of Proportion of feetfittg. is 15 lainutes. of same. biliary matters. VlllaryMnatters io dry residue. Iintuediatel^ 640 grains 33 grains 10 grains .30 1 hoar 1,990 " 105 " 4 '• .03 3 hours 780 " . 60 " 4 " .07 1 6 " 750 " 73 " 3 " .05 9 " 860 " 78 " 4. " .06 12 " 325 . " 23 " 3| " .16 15 " 347 " 18 " 4 u .22 18 " — 21 " 384 " 11 " 1 " .09 24 " 163 " 9.i " 3J " .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. The next point to be ascertained with regard to this question is the following, viz : What becomes of the bile in its passage through the intestine P 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 npon the intestine at varions 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- csecal valve; and at the same time their color changes from flight 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 approximately that of the biliary substanc^ 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 VARIATIONS AND FUNCTIONS OF BILE. 193 large intestine as it is in the small. But even this inconsiderable quantity, found in 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 function of the bile in the intestine, our first object must be to determine what part, if any, it takes in the digestive process. As the liver is situated, like the salivary glands and the pancreas, in the immediate vicinity of the alimentary canal, and lik-e 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. It is a very remarkable fact, in this connection, that the bile pre- cipitates by contact with the gastric juice. If four drops of dog's bile be added to 3j of gastric juice from the same animal, a copious yellowish-white precipitate falls down, which contains the whole of the coloring matter of the bile which has been added ; and if the mixture be then filtered, the filtered fluid passes through quite colorless. The gastric juice, however, still retains its acid reaction. This precipitation depends upon the presence of the biliary sub- stances proper, viz., the glyko-cholate and tauro-cholate of soda, and not upon that of the incidental ingredients of the bile. For if the bile be evaporated to dryness and the biliary substances extracted 13 194 THE BILE. by alcohol and precipitated by ether, as above described, their watery solution will precipitate with gastric juice, in the same manner as fresh bile would do. Although the biliary matters, however, precipitate by contact with fresh gastric juice, they do not do so with gastric juice which holds alhuminose in solution. We have invariably found that if the gas- tric juice be digested for several hours at the temperature of 100° F., with boiled white of egg, the filtered fluid, which contains an abundance of albuminose, will no longer give the slightest precipi- tate on the addition of bile, or of a watery solution of the biliary substances, even in very large amount. The gastric juice and the bile, therefore, are not finally antagonistic to each other in the digestive process, though at first they produce a precipitate on being mingled together. It appears, however, 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 constantly, and continue even after the animal has been kept for many days with- out food. These facts would lead us to jegard 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 discharge'Il 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 efiete in- gredients; 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 VARIATIONS AND FUNCTIONS OF BILE. 195 and discharged, but furthermore that it be discharged into the 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 important ftmction 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 system, 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 ' Op. oit., p. 103. - 1, 196 THE BILE. essentially interfered with, and none of the food was discharged 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. A similar experiment has been successfully performed by Prof. A. Flint, Jr.' In this instance the animal lived for thirty-eight days after the operation, and died finally of inanition ; the symp- toms agreeing in every important particular, with those reported by Bidder and Schmidt. 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. Containing I '^^*^' 874.20 grains. I. Solid residue 633.95 " 1508.15 ' American Journ. Med. Soi., October, 1862. ' Op. cit., p. 217. VAKIATION'S AND FUNCTIONS OF BILE, 197 The solid residue was composed as follows : — Neutral fat, soluble In ether . , 43.710 grains. Fat, with traces of biliary matter . 77.03S " 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 Now, as it has 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, 36.75 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. We 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- 198 THE BILE. 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 the jugular vein, or the 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, -fre 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 pre- cipitable by ether from its alcoholic solution. This substance is often considerably 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 dif&cult, and often impossible, to obtain enough of it, "mixed with ether, for microscopic examination. It dissolves, also, like the biliary sub- stances, 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 considered more abundant in the case of the portal blood ; and yet that from the blood of the vena cava, dis- solved in water, will give Pettenkofer's reaction for bile perfectly, while that of the portal 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 VARIATIONS AND FUNCTIONS OF BILE. 199 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 eecretiori, 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. 200 FOBMATION OF SU.GAS IN THE LIVER. CHAPTER IX. POEMATION OF SUGAE IN THE LIVER. Beside the secretion of bile, tlie liver performs also another exceedingly important function, viz., the production 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 effected 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 Ber- nard' in 1848, and described by him under the name of the glyco- genic function of the liver. It has long been known that sugar may be abundaiitly secreted, under some circumstances, when no vegetable matters have been taken with the food. The milk, for example, of all animals, car- nivorous as well 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 Fonotion dn Foie. Paris, 1853. FORMATION OF SUGAB IN THE LIVER. 201 urine lias often appeared to be altogether out of proportion to tliat whicli 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 other organ in the body may be entirely destitute of sugar, but the liver always contains it in considerable quantity, 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 exa- mination, the liver was found, in each instance, to contain a propor- tion 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 Fonotion du Foie, p. 50. 202 FORMATION OF SUGAR IN THE LIVER. its'origiu 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 (Muriena anguilla and Eaia 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 iii 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: — In man , " monkey " dog . " cat . " rabbit " sheep Pekcentage op Sdgak in the Livek. . 1.68 In ox . . . . 2.30 . 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 J)roperties 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 potassa. It ferments very readily, also, when mixed with 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 Physiologie Experimentale. Paris, 1855, p. 213. FORMATION OF SUGAR IN THE LIVER. 203 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 tisstce, 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 tissue 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, wUl 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 ia 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 process 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. 204 FORMATION OF SUGAR IN THE LIVER. 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 glycogenic matter, or glycogene. This glycogenic 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 glycogenic 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, by which the glycogenic 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 waslied 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 may 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 : — Ci^HjuOij. 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 Troinmer'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. FORMATION OF SUGAR IN THE LIVER. 205 Mood. If a solution of glycogenic matter be mixed with fresh human saliva, and kept for a few minutes at the temperature of 100° ¥., 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, glycogenic 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. Neverthe- less, 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. The formation of sugar in the liver is therefore a function com- ' Journal d« Phyaiologie, Paris, 1858, p. 558. 206 FORMATION OP SUGAR IN THE LIVER. 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 glycogenic matter into sugar, by a process of catalysis and trans- formation. 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 gQCS 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 be easily demonstrated by exa- mining comparatively the two kinds of blood, portal and hepatic, 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 the 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, however, 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 Eobin, that it is at first converted into lactic acid {C^'H. fi^), which decomposes in turn the alkaline carbonates, setting free carbonic acid, and forming FORMATION OF SUGAR IN THE LIVER. 207 lactates of soda and potassa. 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 ingestion of food, according to the investigations of Bernard, the circulation of blood through the portal system and the liver becomes consider- ably accelerated. A larger quantity of sugar is then produced 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 capil- laries; and from four to six hours after the commencement 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 sac- charine character. It does not, however, in the healthy, condition, 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 increase 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. 208 THE SPLEEN. CHAPTER X. THE SPLEEX. 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 cceliao 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 which 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 Ijiat 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 theft 'period. When these periodical congestions take place, ' In Gray, on the Structure and Uses of the Spleen. London, 1854, p. 40. THE SPLBEir. 209 the organ becoming turgid witli blood, the capsule is distended ; and limits, by its resisting power, the degree of tumefection 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 invested capsule can be readily seen in the dog or the cat, by opening the abdomen while digestion is going on, exposing 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, stif^ 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 attaphed to the internal surface of its investing capsule. Thes© fibrous cords, or trabeculse, 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 bifarcation 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- 14 210 THE SPLEEN. aided eye, though they always exist in the spleen in a healthy condition. Their aiverage diameter, according to KoUiker, is jjJj 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 (KoUiker, 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 vas- cular 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 nqt 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 THE SPLEEN. 211 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 suf- ficiently 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 abdomi- nal 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 improve- ment 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, appears to exert any permanently injurious effect upon the 212 THE SPLEEN. animal whicli has been subjected to the operation ; and life inay 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. BLOOD-GLOBULES. 213 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 bhod-gilobuhs, 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 uniform 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 red globules of the blood present, under the microscope, a perfectly circular outline and a smooth exterior. (Fig. 54.) Their size varies somewhat, in human blood, even in the same specimen. The greater number of them have a transverse diameter of -g-^^ of an inch ; but there are many smaller ones to be seen, which are not more than -j^Vtr or even ^jVtt of ^^ i^ch 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 flatwisfe, presents a comparatively broad surface and a circular out- 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 b. The thickness of the globule, seen in this position, is about 214 THE BLOOD. Human Bi.ood-qlobui.es. seeu flatwise. 6. Ked globules, White globule. -a. Eed globnies, seen edgewise, e. TsioTT of an inch, or a little less than one-fifth of its transverse diameter. When the globules are examined lying upon their broad sur- faces, it can be seen that these surfaces are not exactly flat, but that there is on each side a slight "'^' ■ central depression, so that the rounded edges of the blood-globule are evidently thicker than its middle por- tion. This inequality pro- duces a remarkable optical effect. The substance of which the blood-globule is composed refracts light more strongly than the fluid plas- ma. Therefore, when exa- mined "with the microscope, by transmitted light, the thick edges of the globules act as double convex lenses, and concentrate the light ^K.v..^ .^^ .^,^. V-. ^^. Consequently, if the object-glass be carried upward by the adjusting screw of the microscope, and lifted away from the stage, so that F'S: •''S- the blood-globules fall be- yond its focus, their edges will appear brighter. But the central portion 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. An alternating appearance of the blood-globules may, there- fore, be produced by view: ing them first beyond and then within the focus of the instrument. .above the level of the fluid. Red Globules of the Blood beyond tlie t'ucus of the microscope. seen a little BLOOU-GLOBULES. 215 Fig. 56. The same, seen a little within the focus. When beyond the focus, the globules will be seen with a bright rim and a dark 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 tjie bodies' de- signated by it. This term will, consequently, be employed when- ever we have occasion to speak of the blood-globules ^'s- ^'^• in the following pages. Within a minute after being placed under the microscope, the blood-globules, after a fluctuating movement of short duration, very often arrange themselves in slight- 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 ow- ing merely,to the coagulation of the blood, which takes place very rapidly when it is spread out in thin layers and in contact with glass surfaces ; and which, by 6L0{)D-aL0BirLE of coin. adhering together, like rolls 216 THE BLOOD. compressing the globules, forces them into such a position that they may occupy the least possible space. This position is evidently 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 ox 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 semi-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. When a moderate quantity of water is mixed with the blood, the BLOOD-GLOBULES. 217 Bt.ood-globules, ewuliea by the imbibition of edges of the globiiles, being thicker than the central portions, and absorbing water more abundantly, become turgid, and encroach gradually upon the central part. (Fig. 58.) It is very ^'s- ^^■ common to see the central depression under these cir- cumstances, disappear on one side before it is lost on the other, so that the globule, as it swells up, curls over to- wards 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 water" the blood. Dilute acetic acid dissolves the blood-globules more promptly than water, and solu- tions of the caustic alkalies more promptly still. If a drop of blood be allowed partially to evaporate while under the microscope, the globules 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. Blood-globules, slirunken, with their margins crenated. 218 THE BLOOD. The entire mass of tlie blood-globules, in proportion to the rest of the circulating fluid, can only be approximately measured by the eye in a microscopic examination. In ordiaary 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 niore 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 : — Composition op the Buiod-Globuies is 1000 Pakts. Water 688.00 Globuline 282.22 Hsematine lli.lCt Patty substances 2.S1 Undetermined (extractive) matters 2.00 Chloride of soilium " potassium ........ Phosphates of soda and potassa ...... Sulphate " " Phosphate of lime " " magnesia ' . . 8.12 lOUO.UO The most important of these ingredients is the globuline. 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 Tje 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 Verdeil, only becomes opalescent at 160°, and requires for its complete coagulation a temperature of 200° F. . The hwmatine 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, BLOOD-GLOBULES. 219 mechanically deposited in the globuline, but the two substances are iutimately mingled throughout the mass of the blood-globule, just as the fibrin and albumen are mingled in the plasma. Hsematine 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 idirectly 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 haematine 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 camelidas, 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 jVbtt of an inch, and the two-toed sloth {Bradypus didactylus), in which they are jgVxr of an inch in diameter. In the musk deer of Java they are smaller than in any other known species, measuring rather less than y^^ty^ 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 measurement of Mr. Gulliver.' D1A.METEB OP Red Globitles in thi Ape • 5l'iFi!°f *" i'"=l'- Cat . ■ Ij'ffilOf an inch Horse . i^lSH u Fox . IT^Sff u Ox Ts'lff 11 Wolf . sbVit " Sheep . J^HIS tt Elephant iAj It Goat . einjs u Red deer Tn'off It Dog . ■ss'njs u Musk deer • TsivB tt 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 camelidse (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, ' In Works of William Hewson, Sydenham edition, London, 1846, p. 327. 220 THE BLOOD. ■357 reptiles, and fish, the blood-globulea differ so much from the above that they can be readily distinguished by microscopic examination. 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 t^uw of an inch in their long diameter; and in Menohrau' chus, the great water lizard of the northern lakes, ^^^ of an inch. In Prot&us angui- nits they attain the size, ac- cording to Dr. Carpenter,^ of of an inch. Beside the corpuscles de- scribed above, there are glo- bules of another kind found in the blood, viz., the white globules. These globules are very much less numerous than the red ; the proportion between the two, in human blood, being one whitie to two or three hundred red globules. In reptiles, the relative quantity of the white 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 j^Vtr ot 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 Blood-glubul U6 OF Froq. — a. Blood-globule Beea edgewise, b. White-globule. The Microscope and Its Revelations, Philadelphia edition, p. 600. BLOOD-GLOBULES. 221 broken up as the action of the acetic acid upon the globule is longer continued. (Fig. 61.) This collection of. granular matter often assumes a curved or crescentic form, as at a, and sometimes various other irregular shapes. It does not indicate the existence of a nucleus in the white globiile, but it is merely an appearance produced by the coagulating and disintegrating action of ^'?- si- 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 independent 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 ob- served. 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 de- tected by our means of observation,'" it must be acknowledged that the above opinion has no solid foundation. It has been stated by some authors (KoUiker, 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 transition from one form to the other ; but this is not a fact which is generally acknowledged. We have repeatedly examiued, with reference to this point, the fresh blood of the frog, as well as that of the menobranchus, in which the large White Globules of the Blood; altered by dilute acetic acid. ■ EolUker, Handbuch der Grenrebelehre, Leipzig, 1852, p. 582. 222 THE BLOOD. size of the globules would give every opportunity for detecting any such changes, did they really exist ; and it is our unavoidable con- clusion from these observations, that there is no good evidence, even in the blood of rdptiles, of any such transformation taking place. There is simply, as in human blood, a certain variation in size and opacity 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 othfer. The red and white globules are therefore to be regarded as distinct and independent anatomical elemeflts. They are mingled together in the blood, just as capillary bloodvessels and nerves are mingled in areolar tissue ; but there is no other connection between them, so far as their formation is concerned, than that of juxtaposition. 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 ho reason for believing that th,e red globules of the blood are any less permanent, as anatomical forms, than the muscular fibres or the nervous filaments. They undeVgo, 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 which 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. Their ingredients are incessantly altered, by transforma- tion and decomposition, as. they pass through various parts of the vascular system;' but the globules themselves retain their form and texture, and still remain as constituent parts of the circulating fluid. PLASMA. 223 1.55 Plasma. — The plasma of the blood, according to Lehmann, has the following constitution : — Composition of the Plasma in 1,000 parts. Water 902.90 Fibrin . . '. 4.05 Albumen . . . ^ 78.84 Fatty matters 1.73 Undetermined (extractive) matters 3.94 Chloride of sodium " potassium Phosphates of soda and potassa Sulphates " . " " Phosphate of lime " magnesia 1000.00 The above ingredients are all intimately mingled in the blood- plasma, in a fluid form, by mutual solution ; but they may be sepa- rated from each other for examination by appropriate means. The two ingredients belonging to the class of organic substances are the fibrin and the albumen. ^he fibrin, though present in small quantity, is evidently an im- portant 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 presents itself under the form of nearly ^'S- ^^■ white threads and flakes, having a semi-solid consist- ency, and a considerable de- gree 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 /, /.I -Ti 111 /T-1. t\n\ mi CoAoni.ATED FlBKiN, showing itsabrillatedcon- " fibnllated." (Fig. 62.) The diiioo. 224 ■ THE BLOOD. filaments of wliicli it is composed are colorless and elastic, and when isolated are seen to be exceedingly minute, being not more than ^Tshnv or even ^^Uu oi an inch in diameter. They are in part arranged so as to lie parallel with each other ; but are more gene- rally 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 dissolve. They are often interspersed everywhere with minute granular mole- cules, 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 solu- tion, or by prolonged boiling at a very high temperature. These agents, however, produce a complete alteration in the properties 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° F., or by contact with alcohol, the mineral acids, the metallic salts, or with ferrocyanide of potassium in an acidulated solution. It exists natu- rally 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 plasina of the blood, therefore, after the removal of the fibrin, be exposed to the |:emperature of 160° F., it solidifies almost completely; so that only a few drops of water remain that j 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 ^iters 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 Robin and Verdeil, op. oit., vol. ii. p. 202. COAGULATION OP THE BLOOD, 225 way into the blood, and circulates for a time uncnanged. After- ward 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 potas- sium, and the phosphates of soda and potassa 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 dif- ferent, 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 phosphates of soda and potassa 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 oiextraciive matters consjst of a mixture of different ingredients, belonging mostly to the class of organic substances, which have not yet been separated in a state of sufiScient 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 alka- line carbonates, which are held in solution in the serum. It has already been mentioned that while the phosphates are most abun- dant in the blood of the carnivora, the carbonates are most abun- dant in that of the herbivora. Thus Lehmann' found carbonate 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 crystalHza- ble substances derived from the transformation of other ingredients of the blood, or of the tissues through which it circulates. - 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 coagulation or clotting. This process com- mences at nearly the same time throughout the whole mass of the blood. The blood becomes first somewhat diminished in fluidity, ' Op. cit., vol. i. p. 546. * Op. cit., vol. i. p. 393. 15 226 THE BLOOD. 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 case of the human subject, in about fifteen min- utes 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 th.e 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- Fig. 63. nous, trembling, jelly-like mass, which is ^-— ■ 7-^ known by the name of the " crassamentum," ^^I^BaBV/ or "clot." (Fig. 63.) ^^^f^S^HI As soon as the clot has fairly formed, it 1 ^^^^^^V I begins to contract and diminish in size. Ex- \^^B^^ I actly how this contraction of the clot is pro- ^ ^^^^^ y duced, we are unable' to say ;. but it is proba- . -^ ^ bly a continuation of the same process by Bowl of reeentiy coAGu- which its Solidification is at first accomplished, LATED BLOOD showing the ^j. ^^ Jg^^^ ^^^ g-j^^JJ^j. ^ J^ _^g ^^ whole mass uuiformly sohdl- J ■ fled. contraction proceeds, the albuminous fluids begin to be pressed out from the meshes in which they were entangled. A few isolated' drops first appear on the surface of the clot. These drops soon increase in size and be- COAOULATION OF THE BLOOD. 227 Fig. 64.- 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 ipore 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 usually separated into two parts, viz., the cht B(,„, „, coaoblateb which is a red, opaque, dense and resisting blood, after twelve hour«, ^r. p ^ Jl^ • J showinsf the clot cuntracted semi-solid mass, consisting ot the fibrm and and aoating m the fluid serum. 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 :-^ Before coagulation the blood consists of 1st. Globdles ; and 2d. Plasma — containing After coagulation it is separated into f Fibrin, Albumen, Water, 1st. Clot, containing < Fibrin and Globules ; L r Albumen, and 2d. Sebum, containing \ Water, y 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 Robin and Verdeil. 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, iij a full stream, from a large orifice, it remains fluid for a comparatively long time; if it be drawn slowly, from a narrow orifice, it coagulates quickly. Thus it usually happens that in the operation of venesection, the blood drawn immediately after the 228 THE BLOOD. 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 lace- rated 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 pneu- . monia, 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 th» 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- tirely different in their character from the fibres of areolar tissue, or any other normal anatomical elements. They are simply the ulti- COAGULATION OF THE BLOOD. 229 mate 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 differs 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 conditions. But the particular conditions neces- sary for coagulation vary with -the different organic . substances. Thus albumen coagulates 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 mag- nesia, which has no eflect 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 "spontaneous," properly speaking, than that of any other organic substance. Still less does it indicate anything like organ- ization, or even a commencement of it. On the contrary, in the natural process of nutrition, fibrin is assimilated by the tissues and takes part in their organization, only when it is absorbed by them from the bloodvessels 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 in- duce its coagulation. If a ligature be placed upon an artery in the living subject, the blood which stagnates above the ligature coagu- lates, just as it would do if entirely removed from the circulation. If the vessel be ruptured or lacerated, the blood which escapes from it into the areolar tissue coagulates, because here also it is with- drawn 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 chordae tendinesB and the edges of the valves, just as it would do if with- drawn from the body and stirred with a bundle of twigs. In every instance, the coagulation of the fibrin is a morbid phenomenon, de- pendent on the cessation or disturbance of the circulation. 230 THE BLOOD. Fig. 65. Vertical section of a Re- cent CoAGULUM, showing the greater accumulation of blood-globules at the bottom. 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 are thus retained in the position in which they hap- pen to be at the moment that coagulation takes place. If the coagulation, however, be delayed longer than usual, or if the globules sink more rapidly than is cus- tomary, they will have time to subside entirely from the upper por^ tion 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 different portions, viz.j 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 pre- senting 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 contrac- tion 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 colorless portion diminishes very much in size ; and as its edges separate from the sides of the vessel, they curl over toward each other, so that the upper surface of the clot becomes more or less excavated or cup-shaped. (Fig. 66.) Bowl of COAatJIjATRD Blood, showing the clot buffed and cupped. COAGOLATIOX OF THE BLOOD. 231 The blood is then said to be "buffed and cupped." These appear- ances 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 suftered 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. 232 BESPIBATIOlir. CHAPTER XII. RESPIRATION. The blood as it circulates in the arterial system has a bright scariet 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, on 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 respiration. 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 membraije, 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 brcmchise ; that is, delicate filamentous prolongations of some part of the EBSPIKATION. 233 integument or mucous membranes, -which 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 menobranchus (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 „ ^ ■., vy.^u,! ou^ ii, ^^^ v.^ V- Head ahd Gills or Mehobbaxchij;!. 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 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 brailchial 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 234 BKSPIBATION. swallowing movement, and is after a time regurgitated and dis- charged, in order to make room for a fresh supply. In frogs, turtles, 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 commu- nicate with the central pulmonary cavity ; and the partitions^ which join each other at various angles, are all composed of thin, pro- jecting 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 memba-ane is much 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 secondary and tertiary bronchi ; the subdivision continuing, while the bron- chial tubes grow smaller and more numerous, and separate con- stantly 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, according to Kolliker, to the size of ^^ of an inch in diameter ; arid are com- posed only of a thin mucous membrane, lined with pavement epi- thelium, 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 j'j of an inch in diameter, which is termed a " pulmonary lobule." Each pulmonary lobule resembles m its structure the entire frog's lung in miniature. It consists of a LnNG OF Fkoo, showing Ub internal sur- face. RESPIRATION". Fig. 69. 235 Fig. 70. HtTMAN Lartnx, Trachea, Bbonchi, and Lungs; Bliowiug the niniiQcaiioD of Ilia bronchi, and the dlvUion of the lungs into lobules. vascular membrane inclosing a cavity; whicli cavity is divided into a large number of secondary compartments by thin septa or partitions, •whicli project from its internal surface. (Fig. 70.) These secondary cavities are the "pulmonary cells," or " vesicles." Each vesicle is about 7^^ 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- ing to the observations of Kdlliker, are J.l\'::,''^r,\L:LlZ lined everywhere with a layer of pavement . "'•'"' '"h*- '■ cavity of lobuie. .., ,. .. .,1 .1 . • .1 ' e. e. 0. Pulmonary cells, or vesi- epithelium, continuous with that in the cies. 2^36 EESPIRATION, ultimate bronchial tubes. The whole extent of respiratory sur- face in both lungs has been calculated by Lieberkuhn^ 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 '. 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, algae, 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 part's 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 transportatio7i, by which the substances pro- 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 cojjvey the blood from the heart to the different tissues and organs of the body. THE HEART. 265 3d.' The capillaries; a network of minute inosculating tubules, ■which are interwoven with the substance of the tissues, and which bring the blood into intimate contact with the cells and fibres of which they 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 HEAKT. The structure of the heart, and of the ferge vessels connected with it, varies considerably in difterent 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 by anatomical relations with Fig. 76. the organs of circulation, the latter are neoessarily 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 veing unite upon the median line to form the aorta (d) by which the blood is finally distributed throughout the frame. In ClECFLATloK OF F18H. — a. Anricln. b. Ventricle, cc. OiLls. d. Aorta, ee. Veuae cavas. 266 THE CIKCULATIOIT. Fig. 77. these animalg 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. (Fig. 77.) The venae cavse discharge their blood into the right auricle (d), whence it passes into the ventricle (tf). 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 (b), and thence into the ventricle (c), where , it mingles with the venous blood which has just arrived by the venae cavse. 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- ClRCULATION OF KKPTII.KS. — «. Right auricle, 'b. Lertaaricln. «.-. VnDtricle. d. Lungs, e. Aorta. /. Vena cava. THE HBABT. 267 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 arterial system. We have J^'ig- 78- therefore a double cireula- tion, and also a double heart; the two sides of which, though united externally, are separate internally. The mammialian heart consists of a right auricle and ventricle (a, i), receiving the blood from the vena cava (j), 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 perieardiun^ and the pericardial fluid, but capable of a very free lateral and rotatory motion. The aurioles, 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 CiBonr.ATiox iw MahmaiiIahs.— n. KigM auricle, h. Right Teotricle. c. Pnlmonary artery. d. Lunps. 0. Fntmonary vein. /. Left auricle, g. Left Tentricle. K Aorta, i. Vena cava. 268 THE CIRCULATIOX. apex of the heart is constituted altogether by the point of the '. ventricle. Fig. 79. Hdmah H£abt, anterior Yiev.-r a. Bight Tentrlcle. b. Left ventricle, c. Btglit auricle, d. Left auricle, e. .Pttlmonary artery, /. Aorta. Fig. 80. HnHAH Heakt, posterior view.— A. Right vedtricle. &. Left ventricle, e. Bight 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 constrictidns. These constricted orifices, by which the different caviti^. communicate, are known by the names of the Fig. 81. Eight Auricle and Ventricle; Auriculo-veotricular Valves open, Arterial Valveti closed. auricular, auriculo-ventricular, and aortic and pulmonary orifices; the auricular orifices being the passages from the vense cavae and THE HEART. 269 pulmonary veins into tHe right and left auricles; tlie 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 aurict|Io-ventricular, aortic, and pulmonary orifices are fur- nished with valves, which allow the blood to pass readily from the auricles to the veiitricles, 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 VBSTRicrjB; Aurlcnlo-ventricalar Valves closed, Arterial VaWes opea. lungs. Eetuming 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 bipod, 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- 270 THE CIRCULATION. charged into the arteries. Each one of these successive actions is called a beat, or pulsation of the heart. Fig. 83. C0UB8E OP Blood throdoh the Hrart. — n^o. Vena cava, superior and inferinr. 6. Biiflit ventricle, w. Pulmonary artery, d. Fulmiiuary vein. c. 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 over 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 sterrium 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 THE HEART. 271 a pulsation. It is followed by an equal interval of silence ; after whieh 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. -iBt quarter Uj^^j^^^^^ 2d " i 3d " Second sound. 4th " Interval of silence. t 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, directly over the situation of the above-mentioned valves ; 2d, the farther we recede in any direction.from this point, the fainter be- comes the sound ; and 3d, in experiments upon the living animal, often repeated by difierent 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 dis- appears, and remains absent until the valve is again liberated. These valves consist of fibrous sheets, covered with a layer of endocardial epithelium. 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, be- coming 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 closurt of ' Diseases of tdS Heart, Kneelai.d'8 translat:on, Boston. 1846 272 THK CIRCULATION. 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 efiect 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-ventriculai' valves are inserted, and where the sound pro- duced by the tension of these valves would i>e most readily con- (Jucted to the ear. 3d. 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, the difi'erent 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 cardfao cavities than in the larger arteries or veins. 4th. 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 chordae 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 THE HEART. 273 heard a slight _^Wion 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 difierpnt, 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 tbe 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 larynx, 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 carefully cut away from its attachments, and the lungs inflated by insufilation through the trachea. .By keeping up a steady artificial respiration, the move- 18 274 THE CIBCULATION. ments Gf 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 relaooation 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, whifth contract simultaneously with each other, and which cotistitute 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 THE HEAET. 275 such. It is, however, a phenomenon precisely similar to that which takes place in the contraction of a voluntary muscle, which becomes swollen and indurated at the same moment and in the same propor- tion that it diminishes in length. 2. At the time of contraction, the ventricles elongate and the poipt 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 elevation 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 pf 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. P. Sydenham ed., London, 1847, p. 21. • Philadelphia Medical Examiner, No. .44. 276 THE CIBCULATION. can still be readily excited by touching 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 ov Froo la A state uf relaxa- tioa. Heakt op Frog iQ contraction. 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 angle. This contraction is immediately followed by a relaxation ; the point of the heart sinks down, and its sides again bulge out- ward. •Jjet us now see in what manner this change in the figure of the ventricles dusing 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 efiFect 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 Diagram of Simple Looped Fibres, in relaxation and con- traction. THE HEART. 277 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, curi- a's- ^^' ing round the centre of its apex, and then, changing their direction, be- come deep-seated> run upward along Pig. 89. Bullock 'h Hi; art, anterior view, showing the supei ficial m uscular fibres. Left Vrntkicle op Bullock's Heart, show- ing tlie deep fibres. the septum and internal surface of the ventricles, and terminate in the columnas camese, and in the inner border of the aurioulo- ventricular ring. The deepest 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 thatSts sides are drawn together. This efiect is illustrated in the accompanying diagram (Fig. 90), where the white lines show the figure of the heart during relaxation, with the course of its circular 278 THE CIRCULATION. 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 Fig. 90. 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 elonga/tion 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 DiagramofCjRcnT.ABFiBRKs . (Jiagrams^ we shall see that, provided the 0? THE Heart, aod their con- ■,..■,■,.■,. traction. fibres wcre arranged in simple longitudi- nal loops (Fig. 87), their contraction would merely have the eflfect 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, also in a direct line, without deviating or twisting either to the Fig 91. right or to the left. But in. 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 s.uperficial 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 COHVERGING FTBRBR AT THE Apex of the Heart, THE HEAKT. 279 very perceptible to tbe eye at each 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 through 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 js 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, agains 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 peri- staltic motion, except that it is more sudden and vigorous. 280 THE CIRCULATION. 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, n6t 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 contrafction 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 aj'tery ; 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 j;hat 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 tQ 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 inci- dents, 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 accor.dingly 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. oit., p. 31. THE ARTEBIES AND THE ABTEEIAL CIBCULATIOX. 281 protrudes, strikes against the walls of*tlie chest, and twists from left to right, the auriculo-^entricular valves shut back> the first sound is produced, and the blsod 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 alter- nate with each other, and form, by their recurrence, the successive cardiac pulsations. THE ARTERIES AND THE ARTERIAL CIRCULATION. The arteries are a saries 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 -condense'd layers of areolar .tissue. The essential anatomical difference be- tween the 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 man- ner around the vessel, like the circular fibres of the muscular coat of the intestine. In arteries of medium size the middle coat con- tains both muscular and elastic fibres; while in those of the largest calibre it consists of elastic tissue alone. The large arteries, ac- cordingly, 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 vessel ; and therefore that the combined area of all the small arte- ries must be considerably larger than that of the aorta, from which 282 THE CIBCULATIOX. the arterial system originStes. As the blood, consequently, in its passage from the heart outward, flows successively through larger and larger spaces, the rapidity of its circulation must necessarily 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 Mood 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 v the arterial circulation takes place is as follows. The arterial sys- tem is, as we have seen, a vast and connected ramification of tubular canals, which may be regarded as a- great vascular cavity, divided and subdivided from within outward by the successive branching of its vessels, but communicating freely with the heart and aorta at one extremity, and with the capillary plexus at the other; and this vascular system is filled everywhere with the circulating fluid. At the time of the heart's contraction, the muscular walls of the ventricle act powerfully upon its fluid contents. The auriculo- ventricular valves at the same time shutting back and preventing the blood from regurgitating into the ventricle, it is forced out through the aortic orifice. A charge of blood is therefore driven into the arterial ramifications, distending their walls by the addi' tional quantity of fluid forced into their cavities. When the ven- tricle immediately afterward relaxes, the active distending force is removed ; and the elastic arterial walls, reacting upon their contents, 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 blood is therefore urged onward, under the pressure of the arterial elasticity, into the capillary system. When the arteries have thus again partially emptied themselves, and returned to their original dimensions, they are again distended by another contraction of the heart. In this manner a succession of impulses or distensions is produced, which alternates with the reaction or subsidence of the vessels, and which can be felt throughout the body, wherever the arterial ramifications penetrate. This phenomenon is known by the name of the arterial pulse. When the blood is thus driven by the cardiac pulsations into the arteries, the vessels are not only distended laterally, but are elongated as well as widened, and enlarged in every direction. Particularly THE ABTEBIES AND THE ARTERIAL CIRCULATION. 283 when the vessel takes-a curved or serpentine course, its elongation and the increase of its curvatures may be observed at every pulsa- tion. This may be. seen, for example, in the temporal, or even in the radial arteries, in emaciated persons. It is also very well seen in the mesenteric arteries, when the abdomen is opened in the living animal. At every contraction of the heart the curves of the artery on each side become more strongly pronounced. (Fig. 92.) The vessel even rises up partially out of its bed, particularly wjjere 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. But the artCTial pulse has certain other pecu- liarities which deserve a special notice. In the first place, if we place one finger upon the chest at the situation of the apex of the heart, and an- other upon the carotid artery at the middle of the neck, we can distinguish liltle or no difference in time between the two impulses. The disten- Elongation and curva- sion of the carotid seems to take place at the »"" »' "" a»teei i.^. ^ rULSATlON. same instant with the contraction of the heart. But if the second finger be placed upon the temporal artery, instead of the carotid, there is a perceptible interval between the two beats. The impulse of the temporal artery is felt a little, later than that of the heart. In the same way the pulse of the radial artery at the wrist seems a little later than that of the carotid, and that of the posterior tibial at the ankle joint a little later than that of the radial. So that, the greater the distance from the heart at which the artery is examined, the later is the pulsation perceived by the finger laid upon the vessel. But it has been conclusively sh"own, particularly by the investi- gations of M. Marey,' that this difference in time of the arterial pulsations, in different parts of the body, is rather relative than absolute. By the contraction of the heart, the impulse is commu- nicated at the same instant to all parts, of the arterial system ; but the apparent difference between them, in this respect, depends upon the fact, that, although all the arteries begin to be distended at the ' Dr. Brown-Seqnard's Journal de Pliyslologie, April, 18.59. 284 THE CIBCULATION. same moment, yet those nearest the heart are distended suddenly and rapidly, while for those at a distance, the distension takes place more slowly and gradually. Thus the impulse given to the finger, which marks the condition of maximum distension of the vessel, occurs a little later at a distance from the heart, than in its imme- diate proximity. This modification of the arterial pnlse is produced in the follow- ing way : — The contraction of the left ventriqle is a bru^ue, vigorous and sudden motion. The charge of blood, thus driven into the .arterial system, meeting with a certain amount of resistance from the fluid , already filling the vessels, does not instantly displace and force onward a quantity of blood equal to its own mass, but a large proportion of its force is used in expanding the Ostensible walls of the vessels. In the immediate neighborhood, therefore, the expansion of the arteries is sudden and momentary, like the con- traction of the heart itself. But this expansion requires for itB completion a certain expenditure, both of force and time ; so that at a little distance farther on, the vessel is neither distended to the same degree nor with the same rapidity. At the more distant point, accordingly, the arterial impulse is less pow^irful and arrives more slowly at its maximum. ' On the other hand, when the heart becomes relaxed, the artery in its immediate neighborhood contracts upon the blood by its own elasticity ; and as its contraction at this time meets with no other resistance than that of the blood in the smaller Vessels beyond, it drives a portion of its own blood into them, and thus supplies these vessels with a certain degree of distending force even in the inter- vals of the heart's action. Thus the difference in size of the carotid artery, at the two periods of the heart's contraction and its relaxa^ tion, is very marked ; for the degree of its distension is great when the heart contracts, and its own reaction afterward empties it of blood to a very considerable extent. But in the small branches of the radial or ulnar artery, there is less distension at the time of the cardiac contraction, because this force has been partly expended in overcoming the elasticity of the larger vessels; and there .'is less emptying of the vessel afterward, because it is still kept partially filled by the reaction of the aorta and its larger branches. In other words, there is progressively less variation in* size, at the periods .of distension and collapse, for the smaller and distant arteries than for those which are larger and nearer the heart. THE ARTERIK3 AND THE ARTERIAL CIRCULATION. 285 Mr. Marey has illustrated these facts by an exceedingly ingenious and effectual contrivance. He attached to the pipe of a small forcing pump, to be worked by alternate strokes of the piston, a long elastic tube open at the farther extremity. At diflferent points upon this tube there rested little movable levers, which were raised by the distension of the tube whenever water was driven into it by the forcing pump. Each lever carried upon its extremity a small pen- cil, which marked upon a strip of paper, revolving with uniform rapidity, the lines produced by its alternate elevation and depression. By these curves, therefore, both the extent and rapidity of distension .of different parts of the elastic tube were accurately registered. The curves thus produced are as follows : — Fig. 93. Curves op the Arterial PirLBATii>K,aa ilTaBtrated by M, Marey *8 experiment. — 1. Near the dUtendiDg force. 2. At a distance from it. 3. Still farther reraoved. It will be seen that the whole time of pulsation is everywhere of equal length, and that the distension everywhere begins at the same moment. But at the beginning of the tube the expansion is wide and sudden, and occupies only a sixth part of the entire pulsation, while all the rest is taken up by a slow reaction. At the more repiote points, however, the period of expansion becomes longer and that of collapse shorter; until at 3 the two periods are com- pletely equalized, and the amount of expansion is at the same time reduced one-half. Thus, the farther the blood passes from the heart outward, the more uniform is its flow, and the more moderate the distension of the arteries. Owing to the alternating contractions and relaxations of the heart, accordingly, the blood passes through the arteries, not in a steady stream, but in a series of welling impulses ; and the hemorrhage from a wounded artery is readily distinguished from venous or capillary hemorrhage by the fact that the blood flows in successive jets, as well as more rapidly and abundantly. If a puncture be made in the walls of the ventricle, and a slender canula introduced. 286 THE CIKOULATION, 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 con- tinuous ; only it is very much stronger at the time of ventricular contraction, and diminishes, though it does not entirely cease, at the time of relaxation. If the blood were driven through a series of perfectly rigid and unyielding tubes, its flow would be every- where intermittent ; and it would be delivered from an orifice situ- ated at any point, in perfectly interrupted jets. But the arteries are yielding and elastic; and this elasticity, as .we have already explained, moderates the force of the separate arterial pulsations, and gradually fuses them with each other. 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 impefceptible. 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. Whatevar 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 impuls^ 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 dimi- nishes 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 THE ARTERIES AND THE ARTERIAL CIRCULATION. 287 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- 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 passage of the blood through the arterial system takes place under a certain degree of constant pressure. For these vessels being everywhere elastic, and filled with blood, they constantly tend to react, more or less vigorously, and to compress the circulating fluid which they contain. If any one of the arteries, accordingly, be opened in the living animal, and a glass tube inserted, the blood will immediately be seen to rise in the tube to a height of about five and a half or six feet, and will remain at that level ; thus indi- cating the pressure to which it was subjected in the interior of the vessels. This constant pressure, which is thus due to the reaction of the entire arterial system, is known as the m-terial pressure. The degree of arterial pressure may be easily measured by con- necting the open artery, by a flexible tube, with a small reservoir of mercury, which is provided with a narrow upright glass tube, open at its upper extremity. When the blood, therefore, urged by the reaction of the arterial walls, praeses upon the surface of the merctrry in the receiver, the mercury rises in the upright tube, to a corresponding height. By the use of this instrument it is seen, in the first place, that the arterial pressure is nearly the same all over the body. , Since the cavity of the arterial system is every- where continuous, the pressure must necessarily be communicated, by the blood in its interior, equally in all directions. Accordingly, the constant pressure is the same, or nearly so, in the larger and the smaller arteries, in those nearest the heart, and those at a distance. This constant pressure averages, in the higher quadrupeds, six inches of mercury, which is equivalent to from five and a half to six feet of blood. It is also seen, however, in employing such an instrument, that the level of the mercury, in the upright tube, is not perfectly steady. 288 THE CIBCULATION. but rises and falls with the pijlsations of the heart. Thus, at every contraction of the ventricle, the mercury rises fbr about half an inch, and at every relaxation it falls to its previous level. .Thus the instrument becomes a measure, not only for the constant pressure of the arteries, but also for the intermitting pressure of the heart ; and on that account it has received the name of the cardiometer. It is seen, accordingly, that each contraction of the heart is superior in force to the reaction of the arteries by about one-twelfth ; and these vessels are kept filled by a succession of cardiac pulsations, and discharge their contents in turn into the capillaries, by their own elastic reaction. The rapidity 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 fai;ther and farther from the qentre of the circulationi i 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 moves 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- -butes 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 througli 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 tlie 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 blobd-globules can- not be distinguished in it on microscopic examination, but only a THB ARTEEIES AND THK AETKBIAL CIRCULATION, 289 mingled current shooting forward with increased velocity at every cardiac pulsation. Volkmann, 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 divided transversely, and its cardiac extremity fastened to the upper end (b) of the instrument, while its peripheral extremity was Fig. 94. VoiHjrANN'B AppAKATUd for meaaaring thp rapidUy of the artorml circulation. 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- ties of the artery. The instrument, however, was provided, as shown in the accompanying figures, with two transverse cylindrical plugs, also perforated ; and arranged in such a manner, that when, at a 19 290 THE CIRCULATION. 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 ttie blood in a given time could be readily measured upon a scale attached to the side of the glass tube. Volkmann found, as the liverage 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. 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 arteries in containing a much smaller quantity of muscular and elastic fibres, and a larger proportion of simple condensed areolar tissue. They are conseqijently more flaccid and compressible than the arteries, and less elastic and contractile. They are furthermore distin- guished, 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 pjiss readily from the periphery toward the heart, while they prevent altogether its reflux in an opposite direction. Although' the veins are provided with walls which are very much thinner and less elastic than those of the arteries, yet, contrary to what we. might expect, their capacity for resistance to pressure is equal, or even superior, to that of the arterial tubes. Milne Ed- wards' has collected the results of various experiments, which show that the veins will sometimes resist a pressure which is sufficient to rupture the walls of the arteries. In one instance the jugular vein supported, without breaking, a pressure equal to a column of water 148 feet in height ; and in another, the iliac vein of a sheep resisted a pressure of more than four atmospheres. The portal vein was found capable of resisting a pressure of six atmospheres; and in one case, in which the aorta of a sheep was ruptured by a pressure of 158 pounds, the vena cava of the same animal supported a pres- sure equal to 176 pounds. , ' Le9on3 sur la Pliysiologie, &o., toI. iv. p. 301. VENOCra CIRCtTLATIOK. 291 This resistance of the veins is to be attributed to the large pro- portion of white fibrous tissue which enters into their composition ; the same tissue which forms nearly the whole of the tendons and fasciae, and which is distinguished by its density and unyielding nature. The elasticity of the veins, however, is much less than that of the arteries. When they are filled with blood, they enlarge to a certain size, and collapse again when the pressure is taken off; but they do not react by virtue of an elastic resilience, or, at least, only to a slight extent, as compared with the arteries. Accordingly, when the arteries are cut across, and emptied of blood, they still remain open and pervious, retaining the tubular form, on account of the elasticity of their walls ; while, if the veins be treated in the same way, their sides simply fall together and remain in contact with each other. Another peculiarity of the venous system is the abundance of the separate chanridsi which it affords, for the flow of blood from the periphery towards the centre. • The arteries pass directly from the heart outward, each separate branch, as a general rule, going to a separate region, and supplying that part of the body with all the blood which it 'requires ; so that the arterial system is kept constantly filled to its entire capacity with the blood which passes through it. But that is not the case with the veins. In injected preparations of the vascular system, we have often two, three, four, or even five veins, coming together from the same region of the body. ' There are also abundant inosculations between the dif- ferent veins. The deep veins which accompany the brachial artery inosculate freely with each other, and also wiljh the superficial veins of the arm. In the veins coming from the head, we have the ex- ternal jugular communicating with the thyroid veins, the anterior jugular, and the brachial veins. The external and internal jugulars communicate with each other, and the two thyroid veins also form an abundant plexus in front of the trachea. Thus the blood, coming from the extremities toward the heart, flows, not in a single channel, but in many channels ; and as these channels communicate freely with each other, the blood passes some- times through one of them, and sometimes through another. 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. 292 THE CIRCULATION. 1. The force of aspiration of the thorax. — When the chest expands by the lifting of the ribs and the descent of the diaphragm, its movement, of course, tends to diminish the pressure exerted upon its contents, and so 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 the pulmonary vesicles. But the blood in the veins is also drawn into the chest at the same time and by the same force. This force of aspiration, exerted by the expan- sion of the chest, is gentle and uniform in character, like the move- ments of respiration themselves. Accordingly its influence is ex- tended, without doubt, to the farthest extremities of the venous system, the blood being gently solicited toward the heart, at each expansion of the chest, without any visible alteration in the size of the veins, which are filled up from behind as fast as they are emptied in front. But if the movement of inspiration be sudden and violent, instead of gentle and easy, a different effect is produced. For then the walls of the veins, which are thin "and flaccid, cannot retain their position, but collapse under the external pressure too rapidly to allow the blood to flow in from behind. In this case, therefore, the vein is simply emptied in the immediate neighborhood of the chest, but the entire venous circulation is not assisted by the movement. The same difference in the effect of an easy and a violent suction movement, may be readily shown by attaching to the nozzle of an air-tight syringe a flexible elastic tube with thin walls, and placing the other extremity of the tube under water. If the piston of the syringe be "now withdrawn with a gentle and gradual" motion, the water will be readily drawn up into the tube, while the tube itself suffers no visible change; but if the suction movement be made rapid and violent, the tube will collapse instantly under the pres- sure of the air, and will fail to draw the water into its cavity. A similar effect shows itself in the living body. If the jugular or subclavian vein be exposed in a dog or cat, it will be seen that while the movements of respiration are natural and easy no fluc- tuation in the vein can be perceived. But as soon as the respira- tion becomes disturbed and laborious, then at each inspiration the vein is collapsed and emptied ; while during expiration, the chest being strongly compressed and the inward flow of the blood arrested, the vein becomes turgid with blood which accumulates in it from behind. In young children, also, the spasmodic movements of res- VEX0U3 CIRCULATION. 293 piration in crjing produce a similar turgescence and engorgement of the large veins during expiration, while they are momentarily emptied during the hurried and forcible inspiration. In natural and quiet respiration, therefore; the movements of the chest hasten and assist the venous circulation ; but in forced or laborious respiration, they do not assist and may even retard its flow. 2. Tlie contrctction of the voluntarg 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; find 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 accoiint Fig. 96. Pig. 97. VjilN with vulvcd open. Vein with valves closed; stream of blood passiDg off by a lateral cbanael. 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 294 THE CIRCULATION. the pressure of the muscles, may readily find a passage through others, which communicate by cross branches with the first. (Figs. 96 and 97.) ^ S. 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 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 respiratioE 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 flaecidity 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 | ; and would require, on the other hand, a velocity of 2 X, to pass in the same ^ime through an opening equal to one- half a square inch. Now the capacity of the entire venous system. THE CAPILLABY CIBCULATIOX. 295 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 arterial blood nearly as three to two. The velocity of the venous blood, as compared with that of 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 suffi- cient accuracy, the relative velocity of the arterial and venous cur- rents, at corresponding parts, of their conrse. Fig. 98. THE CAPILLAET CIECULATION, 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 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 55*^^ 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 middle or muscu- lar. (Fig. 98, a.) The middle coat then diminishes, in thickness. SmaT'Tj Artf.et, with ito musciilHr tunic (fi), breaking np iuto caplllarieit. Fruui tUe-pla nuiUr. 296 THE CIKCULATION. until it is reduced to a single layer of circular, fusiform, unstriped, muscular fibres, whicli in their turn disappear altogether, as the artery merges at last in the capillaries ; leaving only, as we have 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 ^*8- 93. 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 blood in the capillaries 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 Capillar? Network from web of frog's foot. THE CAPILLAEY CIRCULATION. 297 Fig. 1(10. field by the smaller arteries, shooting through them with great rapidity and in successive impulses, and flowing oflF again by the veins at a somewhat slow rate. In the capillaries themselves 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- 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 witb • 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 difl:erence 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 tlie 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 Cafili.abt CiRODiArioN ill web uf r. og'a fuot. 298 THE CIBCULATIOX. 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 manner, the white globules accumulate in the afifected 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 w:ithout 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, &o., are appropriated in special proportions by the diflPerent 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, >nd converted into new materials, characteristic of the different tissues. In this way are produced the musculine of thQ 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 ibr the nutrition of the body. The physical conditions which influence the movement of the blood in the capillaries, are somewhat different from those which regulate the arterial and venous circulations. We must remember that, as the arteries pass from the heart outward, they subdivide and ramify to such an extent that the surface of the arterial walls is very much increased in proportion to the quantity of blood which they contain. It is on this account that the arterial pulsation is so much equalized at a distance from the heart, since the influence of the elasticity of the arterial coats is thus constantly increased from / within outward. But as these vessels finally reach the confines of the arterial system, having already been very much increased in number and diminished in size, they suddenly break up into TUE CAPILLARY- CIRCULATION. 299 a terminal ramificatiou of still smaller and more numerous vessels, and so lose themselves at last in the capillary network. By this fii|jil increase of the vascular surface, the equalization of the heart's action is completed. There is no longer any intermitting or pjalsatile character in the force which acts upon the circulating fluid; and the blood, accordingly, is delivered from the arteries into the capillaries under a perfectly continuous and uniform pressure. This pressure is sufficient to cause the blood to pass with con- siderable rapidity, through the capillary plexus, into the commence- ment of the veins. This fact was first demonstrated by Prof. Sharpey," of London, who employed an injecting syringe with a double nozzle, one extremity of which was connected with a mercu- rial gauge, while the other was inserted into the artery of a recently killed animal. When the syringe, filled with defibrinated blood, was fixed in this position and the vessels of the animal injected, the defibrinated blood would press with equal force upon the mercury im the gauge and upon the fluid in the bloodvessels ; and thus it was easy to ascertain the exact amount of pressure required to force the defibrinated blood through the capillaries of the animal, and to make it return by the corresponding vein. In this way Prof. Sharpey found that when the free end of the injecting tube was attached to the mesenteric artery of the dog, a pressure of 90 milli- metres of mercury caused the blood to pass through the capillaries of the intestine and of the liver ; and that under a pressure of 130 millimetres, it flowed iif a full stream from the divided extremity of the vena cava. We have also performed a similar experiment on the vessels of the lower extremity. A full grown healthy dog was killed, and the lower extremity immediately injected with defibrinated blood, by the femoral artery, in order to prevent coagulation in the smaller vessels. A syringe with a double flexible nozzle was then filled with defibrinated blood, and one extremity of its injecting tube attached to the femoral artery, the other to the mouthpiece of a cardiometer. By making the injection, it was then found that the defibrinated blood ran from the femoral vein in a continuous stream under a pressure of 120 millimetres, and that it was discharged very freely under a pressure of 130 millimetres. Since, as we have already seen, the arterial pressure upon the ' Todd and Bowmann, Physiological Anatomy and Pliysiology of man, vol. ii. p. 350. 300 THE CIRCULATION. blood is equal to six inches/ or 150 millimetres, of mercury, it is evident that this pressure is sufficient to propel the blood through the capillary circulation. ^ Beside, the blood is not altogether relieved from the influence of elasticity, after it has left the arteries. For the capillaries tjiem- selves are elastic, notwithstanding the delicate texture of their yralls ; and even the tissues of the organs which they, traverse possess, in many instances, a considerable share of elasticity, owing to the minute elastic fibres which are scattered through their sub- stance. These elastic fibres are found in considerable quantity in the lungs, the spleen, the skin, the lobulated glands, and more or less in the mucous membranes. They are abundant, of course, in the fibrous tissues of the extremities, in the fasciae, the tendons, and the intermuscular substance. In the experiment of injecting the vessels of the lower extremity with defibrinated blood, if the injection be stopped, the blood does H not instantly cease flowing from the extremity of the femoral vein, but continues for a short time, until the elasticity of the intervening parts is exhausted. The same thing may be observed even in the liver. If the end of a water-pipe be. inserted into the portal vein, and the liver in- jected with water under the pressure of a hydrant, the liquid will distend the vessels of the organ, and pass out by the hepatic veins. But if the portal vein be suddenly tied or compressed, so as to shut off the pressure from behind, the stream drill continue to run, for several seconds afterward, from the hepatic vein, owing to the re- action of the organ itself upon the fluid contained in its vessels. As a general rule, also, the capillaries do not Suffer any backward pressure from the venous system. On the contrary, as soon as the blood has been delivered into the veins, it is hurried onward toward the heart by the compression of the muscles and the action of the venous valves. The right side of the heart itself continues the same process, by its regular contractions, and by the action of its own valvular apparatus ; so that the blood is constantly lifted away from the capillaries, by the muscular action of the surrounding parts. These are the most important of the mechanical influences under which the blood moves through the continuous round of the circu- lation. The heart, by its alternating contractions and relaxations, and by the backward play of its valves, continually urges the blood forward into the arterial system. The arteries, by their dilatable and elastic walls, convert the cardiac pulsations into a uniform and THE CAPILLARY CIRCULATION. 301 steady pressure. Under this pressure, the blood passes through the capillary vessels; and it is then carried backward to the heart through the veins, assisted by the action of the muscles and the respiratory movements of the chest. * At the same time there are certain phenomena trhich are very important in this respect, and which show that various local in- fluences will either excite or retard the capillary circulation in par- ticular parts, independently of the heart's action. The pallor or suffusion of the face under mental emotion, the congestion of the mucous membranes during the digestive process, the local and de- fined redness produced in the skin by an irritating application, are all instances of this sort. These phenomena are usually explained by the contraction or dilatation of the smaller arteries immediately supplying the part with blood, under the influence of nervous action. As we know that the smaller arteries are in fact provided with organic muscular fibres, this may Undoubtedly have something to do with the local variations of the capillary circulation ; but the precise manner in which these effects are produced is a(? present unknown. 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 examination 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 capil- laries 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 sup- posed 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 capillaries 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 cheraico-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 302 THE CIRCULATION'. slow, yet as the distance to be passed over between the arteries and veins is very small, the blood really requires 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 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 corresporiding 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. By extending these investigations to different animals, it was found that the duration of the circulatory movement varied, to some extent, with the size and species. In the larger quadrupeds, as a general rule, it was longer ; in the smaller, the time required was less. In the Horae,' tlie mean duration was 28 seconds. " Dog " " " " 15 " " Goat " " " " 13 « '• Fox " " " " I2i " " Rabbit '" " « "7 « ' Physical Phenomena of Living Beings, Pereira's translation, Philada. ed., 1848, * p. 317. 2 In Milne Edwards, Lepons aur la Physiologic, &o., vol. iv. p. 364. LOCAL VARIATIONS. 303 "When .these results were first published, it was thought to be doubtful whether the circulation were really as rapid as they would make it appear. It was thought that the saline matter which was injected, "travelled faster than the blood ;" that it became " diffused" through the circulating fluid ; that it transuded through dividing membranes ; or passed round to the point at which it was detected, by some short and irregular route. But none of these explanations have ever been found to be cor- rect. They are all really more improbable than the fact which they are intended to explain. The physical diffusion of liquids does not take place with such rapidity as that manifested by the circulation ; and there is no other route so likely to give passage to the injected fluid, as the bloodvessels and the movement of the blood itself. Beside, the first experiments of Poisseuille and others have not been since invalidated, in any essential particular. It was found, it is true, that certain other substances, injected at the same time with the saline matter, might hasten or retard the circulation to a certain degree. But these variations were not very marked, and never exceeded the limits of from eighteen to forty-five seconds. There is no doubt that the blood itself makes the saime circuit in very nearly the same interval, of time. The truth is, however, that we cannot fix upon any absolutely uniform rate which shall express 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 from the heart at which an organ is situated must modify to some extent the time required for its blood to return again tofthe centre of the circulation. The blood which passes through the coronary arteries and the capillaries of the heart, for example, must be returned to the right auricle in a compara- tively short time ; while that which is carried by the carotids into 301 THE CIKCULATIOX. 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 ; whije that which circulates through the abdominal digestive organs and is then collected by the portal system, to be again dispersed through 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 circuiation 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 produce 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 difierent 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 frofi 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 ; LOCAL VABIATIONS. 305 Fig. 101. in the capillaries of the general system, from red to blue. But its tinge also varies very considerably in different paits 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 simple process, but as made up of many difier- ent circulations, going on simultane- ously 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 repre- sent the pulmonary, the lower the gen- eral circulation. This, however, gives but a very imperfect idea of the entire circulation, as it really takes place. It would be much more accurately re- presented by such a diagram as that in Fig. 101, in which its variations 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 ; another, in circulating through 20 Diagram of the CiBciTLATion'. — 1- Heart. 2. LoDnrs. 3. Head and upper extremities. 4. Spleen. 5. Intestine. 6. Kidney. 7. Lower extremities. 8. Liver. 306 THE CIECULATION. the lungSj exhales the carbonic acid which it has accumulated else- where, and absorbs the oxygen which is afterward transported to distant tissues by the current of arterial blood. The phenomena of the circulation are even liable, as we have already seen, to periodical variations in the same organ ; increasing 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-. IMBIBITION AND EXHALATION. 307 CHAPTEE XV. IMBIBITION AND EXH AL ATI ON.— THE LYMPHATIC SYSTEM. During the passage of the blood through the capillaries of the circulatory system, a very important series of changes takes place by which its ingredients are partly transferred to the tissues by exhalation, and at the same time replaced by others which the blood derives by absorption from the adjacent parts. These phenomena depend upon the property, belonging to animal membranes, of imbibing or absorbing certain fluid substances in a peculiar way. They are known more particularly as the phenomena of endosmosis and exosmosis. These phenomena may be demonstrated in the following way. If we take two different liquids, for example a solution of salt and an equal quantity of distilled water, and inclose them in a glass vessel with a fresh animal membrane stretched between, so that there is no direct communication from one to the other, the two liquids being in contact with opposite sides of the membrane, it will be found after a time that the liquids have become mixed, to a cer- tain extent, with each other. A part of the salt will have passed into the distilled water, giving it a saline taste ; and n part of the water will have passed into the saline solution, making it more dilute than before. If the quantities of the two liquids, which have become so transferred, be measured, it will be found that a comparatively large quantity of the water hag passed into the saline solution, and a comparatively small quantity of the saline solution has passed, out into the water. That is, the water passes inward to the salt more rapidly than the salt passes outward to the water. The consequence is, that an accumulation soon begins to show itself on the side of the salt. The saline solution is increased in volume and diluted, while the water is diminished in volume, and acquires a saline ingredient. This abundant passage of the water, through the membrane, to the salt, is called endosmosis ; and 308 IMBIBITION AND EXHALATION. the more scanty passage of the salt outward to the water is called exosmosis. The mode usually adopte,d for measuring the rapidity of endos- mosis is to take a glass vessel, shaped somewhat like an inverted funnel, wide at the bottom and narrow at the top. The bottom of the vessel, is closed by a thin animal membrane, like the mucous membrane of an ox-bladder, which is stretched tightly over its edge and secured by a ligature. From the top of the vessel there rises a very narrow glass tube, open at its upper extremity. When the instrument is thus prepared, it is filled with a solution of sugar and placed in a vessel of distilled water, so that the animal mem- brane, stretched across its mouth, shall be in contact with pure water on one side and with the saccharine solution on the other. The water then passes in through the membrane, by endosmosis, faster than the saccharine solution passes out. An accumulation therefore takes place inside the vessel, and the level of the fluid rises in the upright tube. The height to which the iluid thus rises in a given time is a measure of the intensity of the endosmosis, and of its excess over exosmosis. By varying the constitution of the two liquids, the arrangement of the membrane, &c., the variation in endosmotic action under different conditions may be easily ascertained. Such an instrument is called an endosmometer. If the extremity of the upright tube be bent over, so as to point downward, as endosmosis continues to go on after the tube has become entirely filled by the rising of the fluid, the saccharine solu- tion will be discharged in drops from the end of the tube, and fall back into the vase of water. A steady circulation will thus be kept up for a time by the force of eiidosmosis. The water still passes through the membrane, and accumulates in the endosmo- meter ; but, as this is already full of fluid, the surplus immediately falls back into the outside vase, and thus a current is established which will go on until the two liquids have become intimately mingled. The conditions which influence the rapidity and extent of endos- mosis have been most thoroughly investigated by Dutrochet, who was the first to make a systematic examination of the subject. The first of these conditions is the freshness of- the membrane itself. This is an indispensable requisite for the success of the experiment. A membrane that has been dried and moistened again, or one that has begun to putrefy, will not produce the desired effect. It has been also found that if the membrane of the endosmometer be THE LYMPHATIC SYSTEM. 309 allowed to remain and soak in the fluids, after the column has risen to a certain height in the upright tube, it begins to descend again as soon as putrefaction commences, and the two liquids finally sink to the same level. The next condition is the extent of contact between the membrane and the two liquids. The greater the extent of this contact, the more rapid and forcible is the current of endosmosis. An endos- mometer with a wide mouth will produce more effect than with a narrow one, though the volume of the liquid contained in it may be the same in' both instances. The action takes place at the surface of the membrane, and is proportionate to its extent. Another very important circumstance is the constitution of> the two liquids, and their relation to each other. As a general thing, if we use water and a saline solution in our experiments, endosmosis is more active, the more concentrated is the solution in the endosmo- meter. A larger quantity of water will pass inward toward a dense solution than toward one which is already dilute. But the force of endosmosis varies with different liquids, even when they are of the same density. Dutrochet measured the force with which water passed through the mucous membrane of an ox-bladder into differ- ent solutions of the same density. He found that the force varies with different substances, as follows :' — Eadosmosis of water, with a solution of albnmen . . 12 " " " sugar . . ..11 " " " gum ..." 5 " " " gelatiue . . 3 The position of the memirane also makes a difference. With some fluids, endosmosis is more rapid when the membrane has its mucous surface in contact with the dense solution, and its dissected surface in contact with the water. With other substances the more favor- able position is the reverse. Matteucci found that, in using the mucous membrane of the ox-bladder with water and a solution of sugar, if the mucous surface of the membrane were in contact with the saccharine solution, the liquid rose in the endosmometer between four and five inches. But if the same surface were turned outward toward the water, the column of fluid was less than three inches in height. Different membranes also act with different degrees of force. The effect produced is not the same with the integument of different animals, nor with mucous membranes taken from different parts of the body. ' In Mattpucci's Liectures on the Physical Phenomena of Living Beings. Philada., 1848, p. 48. 810 IMBIBITION AND EXHALATION. Generally speaking, endosnaosis is more active when the temper- ature is moderately elevated. Dutrochet noticed that an endosmo- meter, containing a solution of gum, absorbed only one volume of water at a temperature of 32° Fahr., but absorbed three volumes at a temperature a little above 90°. Variations of temperature will sometimes even change the direction of the endosmosis altogether, particularly with dilute solutions of hydrochloric acid. Dutrochet found, for example,^ that when the endosmometer was filled with dilute hydrochloric acid and placed in distilled water, at the tern-, perature of 50° F., endosmosis took place from the acid to the water, if the density of the acid solution were less than 1.020 ; but that it took place from the water to the acid, if its density were greater than this. On the other hand, at the temperature of 72° F., the current was from within outward when the density of the acid solu- tion was below 1.003, and from without inward when it was above that point. Finally, the pressure which is exerted upon the fluids and the membrane favors their endosmosis. Fluids tha^s pass slowly under a low pressure will pass more rapidly with a higher one. Different liquids, too, require different degrees of pressure to make them pass the same membrane. Liebig^ has measured the pressure re- quired for several, different liquids, in order to make them pass through the same membrane. He found that this pressure was Ikches of Mebcdrt. For alcohol 52 For oil 37 For solution of salt ....... 20 ^ For water 13 ♦» • There are some cases in which endosmosis takes place without being accompanied by exosmosis. This occurs, for example, when we use water and albumen as the two liquids. For while water readily passes in through the' animal membrane, the albumen does not pass out. If an. opening be made, for example, in the large end of an egg, so as to expose the shell-membrane, and the wEble be then placed in a goblet of water, endosmosis will take place very freely from the w,ater to the albumen, so as to distend the shell- membrane and make it protrude, like a hernia, from the opening in the shell. But the albumen does not pass' outward through the membrane, and the water in the goblet remains pure. After a time, ' In Milne Edwards, Le<;ons sur la Physiologie, Ac, vol. r. p. 164. 2 111 Loiiget's Traite di Physiologie, vol. i. p." 384. THE LYJIPHATIC SYSTEM. 311 however, the accumulatiou of fluid in the interior becomes so ex> oessive as to burst the shell-membrane, and then the two liquids become mixed indiscriminately- together. These are the principal conditions by which endosmosis is influ. enced and regulated. Let us now see what is the nature of the process, and upon what its phenomena depend. Endosmosis is not dependent upon the simple force of diffusion or admixture of two different liquids. F<3r sometimes, as in the , case of albumen and water, all the fluid passes in one direction and none in the other. It is true that the activity of the process de- pends very much, as we have already seen, upon the difference in constitution of the two liquids. "With water and a saline solution, for instance, the stronger the solution of salt, the more rapid is the endosmosis of the water. And if two solutions of salt be used, with a membranous septum between them, endosmosis takes place from the weaker solution to the stronger, and is proportionate in activity to the difference in their densities. From this fact, Dutro- chet was at first led to believe that the direction of endosmosis was determined by the difference in density of the two liquids, and that the current of accumulation was always directed from the lighter liquid to the denser. But we now know that this is not the case. For though, with solutions of salt, sugar, and the like, the current of endosmosis is from the lighter to the denser liquid ; in other instances it is the reverse. With water and alcohol, for example, endosmosis takes place, not from the alcohol to the water, but from the water to the alcohol ; that is, from the denser liquid to the lighter. fcThe difference in density of the liquids, therefore, is not the only condition which regulates the direction of the endosmotic current. In point of fact, the process of endosmosis does not depend princi- pally upon the attraction of the two liquids for each other, but upon the attraction of the animal membrane for the two liquids. The membrane is not a passive filter through which the liquids mingle, but it is the active agent which determines their passage. The membrane has the power of absorbing liquids, and of taking them up into its own substance. This power of absorption, belonging to the membrane, depends upon the organic or albuminous ingredients of which it is composed'* and, with different animal substances, the power of absorption is different. The tissue of cartilage, for exam- ple, will absorb more water, weight for weight, than that of the tendons ; and the tissue of the cornea will absorb nearly twice as much as that of cartilage. 312 IMBIBITION AND EXHALATION. Beside, the power of absorption of an animal membrane is dif- ferent for different liquids. Nearly all animal membranes absorb pure water more freely than a solution of salt. If a membrane, partly dried, be placed in a saturated saline solution, it will absorb the water in larger proportion than the salt, and a part of the salt will, therefore, be deposited in the form of crystals on the surface of the membrane. Oily matters, on the*other hand, are usually absorbed less readily than efther water or saline solutions. Chevreuil has investigated the absorbent power of different animal substances for different liquids, by taking definite quanti- ties of the animal substance and immersing it for twenty-four hours in different liquids. At the end of that time, the substance was rembVed and weighed. Its increase in weight showed the quantity of liquid which it had absorbed. The results which were - obtained are given in the following table : — ' 100 Pakts op Water. Saline Solution. ■ Oil. Cartilage, ■ 231 parts. 125 parts. Tendon, 178 " 114 " 8.6 parts Elastic ligament. absorb in 148 " 30' " 7.2 " Cornea, 24 hours, 461 " 370 " 9.1 " Cartilaginous ligament, 319 " 3.2 " Dried fibrin. . 301 " 154 « The same substance, therefore, will' take up different quantities of water, saline solutions, and oil. Accordingly, when an animal membrane is placed in contact with two different liquids, it absorbs one o^hein more abundantly., than the other ; and that which is absorbed in the greatest quantity is also diffused most abundantly into the liquid on the opposite side of the membrane. A rapid endosmosis takes place in one direc- tion, and a slow exosmosis in the other. Consequently, the least absorbable fluid increases in volume by the constant admixture of that which is taken up more rapidly. The process of endosmosis, therefore, is essentially one of im- bibition or absOTption of the liquid by an animal membrane, com- posed of organic ingredients. We have already shown, in de- scribing the organic proximate principles in a previous chapter, that these substances have- the power of absorbing watery and serous fluids in a peculiar way. In endosmosis, accordingly, the ' In Longet's Traite de Physiologie, vol. i. p. 383. THE LYMPHATIC SYSTEM. 313 imbibed fluid penetrates tbe membrane by a kind of chemical combination, and unites intimately with the substance of which its tissues are composed. It is in this way that all imbibition and transudation take place in the living body. Under the most ordinary conditions, the transu- dation of certain fluids is accomplished with great rapidity. It has been shown by M. Gosselin,' that if a watery solution of iodide of potassium be dropped upon the cornea of a living ra,bbit, the iodine passes into the cornea, aqueous humor, iris, lens, sclerotic and vitreous body, in the course of eleven minutes; and that it will penetrate through the cornea into the aqueous humor in three minutes, and into the substance of the cornea in a minute and a half. In these experiments it was evident that the iodine actually passed into the deeper portions of the eye by simple endosmosis, and was not transported by the vessels of the general circulation ; since no trace of it could be found in the tissues of the opposite eye, examined at the same time. The same observer showed that the active principle of belladonna penetrates the tissues of the eyeball in a similar manner. M. Gos- selin applied a solution 9f sulphate of atropine to both eyes of two rabbits. Half an hour afterward, the pupils were dilated. Three quarters of an hour later, the aqueous humor was collected by puncturing the cornea with a trocar; and this aqueous humor, dropped upon the eye of a cat, produced dilatation and immobility of the pupil in half an hour. These facts show that the aqueous humor of the affected, eye actually contains atropine, which it absorbs from without through the corqea, and this atropine then acts directly and locally upon the muscular fibres of the iris. But in all the vascular organs, the processes of endosmosis and exosmosis are very much accelerated by two important conditions, viz., first, the movement of the blood in circulating through the vessels, and secondly the minute dissemination and distribution of these vessels through the tissue of the organs. The movement of a fluid in a continuous current always favors endosmosis through the membrane with which it is in contact. For if the two liquids be stationary, on the opposite .sides of an animal membrane, as soon as endosmosis commences they begin to ap- proximate in constitution to each other by mutual admixture ; and, as this admixture goes on, endosmosis of course becomes less active, ' Gazette Hebdomadaire, Sept. 7, 1855. 314 IMBIBITION AND EXHALATION. and ceases entirely when the two liquids have become perfectly alike in composition. But if one of the liquids be constantly renewed by a continuous current, those portions of it which have become contaminated are immediately carried away by the stream, a'fid replaced by fresh portions in a state of purity. Thus the . difference in constitution of the two liquids is preserved, and transudation will continue to take place between them with una- bated rapidity. Matteucci demonstrated the effect of a current in facilitating endosmosis by attaching to the stopcock of a glass reservoir filled with water, a portion of a vein also filled with water. The vein was then immersed in a very dilute solution of hydrochloric acid. So long as the water remained stationary in the vein it did not give any indications of the presence of the acid, or did so only very slowly ; but if a current were allowed to pass through the vein by opening fehe stopcock of the reservoir, then the fluid running from its extremity almost immediately showed an acid reaction. The same thing may be shown even more distinctly upon the living animal. If a solution of the extract of nux vomica be in- jected into the subcutaneous areolar tissue of the hind leg of two rabbits, in one of which the bloodvessels of the extremity have been left free, while in the other they have been previously tied, so as to stop the circulation in that part — in the first rabbit, the poison will be absorbed and will produce convulsions and death in the course of a few nainutes; but in the second animal, owing to the stoppage of the local circulation, absorption will be much retarded, and the poison will find its way into the general circulation so slowly, and in such small quantities, that its specific effects will show themselves only at a late period, or even may not be produced at all. The anatomical arrangement of the bloodvessels and adjacent tissues is the second important condition regulating endosrnosis and exosmosis. We have already seen that the network of capil- lary bloodvessels results from the excessive division and ramifica- tion of the smaller arteries. The blood, therefore, as it leaves the arteries and enters the capillaries, is constantly divided into smaller and more numerous currents, which are finally disseminated in the most intricate manner throughout the substance of the organs and tissues. Thus, the blood is brought into intimate contact with the surrounding tissues, over a comparatively very large extent of sur- face. It has already been stated, as the result of Dutrochet's inves- tigations, that the activity of endosmosis is in direct proportion to THE LYMPHATIC SYSTEM. 315 the extent of surface over •which the two liquids come in contact with the intervening membrane. It is very evident, therefore, that it will be very much facilitated by the anatomical distribution of the capillary bloodvessels. It is in some of the glandular organs, however, that the transu- dation of fluids can be shown to take place with the greatest rapi- dity. For in these organs the exhaling and absorbing surfaces are arranged in the form of minute ramifying tubes and follicles, which penetrate everywhere through the glandular substance ; while the capillary bloodvessels form an equally complicated and abundant network, situated between the adjacent follicles and ducts. In this way, the union and interlacement of the glandular membrane, on the one hand, and the bloodvessels on the other, become exceed- ingly intricate and extensive; and the ingredients of the blood are almost instantaneously subjected, over a very large surface, to the influence of the glandular membrane. The rapidity of transudation through the glandular membranes has been shown in a very striking manner by Bernard.' This ob- server injected a solution .of iodide of potassium into the duct of the parotid gland on the right side, in a living dog, and immediately afterward found iodine to be present in the saliva of the correspond- ing gland on the opposite side. In the few instants, therefore, re- quired to perform the experiment, the salt of iodine must have been taken up by the glandular tissue on one side, carried by the blood of the general circulation to the opposite gland, and there transuded through the secreting membrane. We have a;lso found the transudation of iodine through the glandular tissue to be exceedingly rapid, by the following experi- ment. The parotid duct was exposed and opened, upon one side, in a living dog, and a canula inserted into it, and secured by liga- ture. The secretion of the parotid saliva was then excited, by in- troducing a little vinegar into the mouth of the animal, and the saliva, thus obtained, found to be entirely destitute of iodine. A solution of iodide of potassium being then injected into the jugu- lar vein, and the parotid secretion again immediately excited by the introduction of vinegar, as before, the saliva first discharged from the canula showed evident traces of iodine, by striking a blue color on the addition of starch and nitric acid. The processes of exosmosis and endosmosis, therefore, in the living ' Le;ons de Pliysiologie Expfirimentale, Paris, 1856, p. 107. 316 IMBIBITION AND EXHALATION. bodj, are regulated by the same conditions as in artificial experi- ments, but they take place with infinitely greater rapidity, owing to the movement of the circulating blood, and the extent of contact existing between the bloodvessels and adjacent tissues. We have already seen that the absorption of the same fluid is accomplished with different degrees of rapidity by different animal substances. Accordingly, though the arterial blood is everywhere the same in composition, yet its different ingredients are imbibed in varying quantities by the different tissues. Thus, the cartilages absorb from the circulating fluid a larger proportion of phosphate of lime than the softer tissues, and the bones a larger proportion than "the cartilages; and the watery and saline ingredients generally are found in different quantities in different parts of the body. The same animal membrane, also, as it has been shown by experiment, will imbibe different substances with different degrees of facility. Thus, the blood, for example, contains more chloride of sodium than chl6ride of potassium ; but the muscles, which it supplies with nourishnient, contain more chloride of potassium ,than chloride of sodium. In this way, the proportion of each ingredient derived from the blood is determined, in each separate tissue, by its special absorbing or endosmotic power. Furthermore, we have seen that albumen, under ordinary coiidi- tions, is not endosmotic ; that is, it will not pass by transudation through an animal membrane. For the same reason, the albumen of the blood, in the natural state of the circulation, is not exhaled from the secreting surfaces, but is retained within the circulatory system, while the watery and saline ingredients transude in varying quantities. But the degree oi pressure to which a fluid is subjected has gfeat influence in determining its endosmotic action. A sub- stance which passes but slowly under a low pressure, may pass much more rapidly if the force be increased. Accordingly, we find that if the pressure upon the blood in the vessels be increased, by obstruction to the venous current and backward congestion of the capillaries, then not only the saline and watery, parts of the blood pass out in larger quantities, but the albumen itself transudes, and infiltrates the neighboring parts. It is in this way that albumen makes its appearance in the urine, in consequence of obstruction to the renal circulation, and that local oedema or general anasarca may follow upon venous congestion in particular regions, or upon general disturbance of the circulation. The processes -of imbibition and exudation, which thus take THB LYMPHATIC SYSTEM. 317. place incessantly throughout the body, are intimately connected, with the action of the great absorbent or lymphatic system of ves- sels, which is to be considered as secondary or complementary to that of the sanguiferous circulation. The lymphatics may be regarded as a system of vessels, com- mencing in the substance of the various tissues and organs, and endowed with the property of absorbing certain of their ingredi- ents. Their commencement has been demonstrated by injections, naore particularly in the membranous parts of the body ; viz., in the skin, the mucous membranes, the serous and synovial surfaces, and the inner tunic of the arteries and veins. They originate in these situations by vascular networks, not very unlike those of the capillary bloodvessels. Notwithstanding this resemblance in form between the capillary plexuses of the lymphatics and the blood- vessels, it is most probable that they are anatomically distinct from each other. It has been supposed, at various times, that there might be communications between them, and even that the lymph- atic plexus might be a direct continuation of that originating from the smaller arteries ; but this has never been demonstrated, and it is now almost universally conceded that the anatomical evidence is in favor of a complete separation between the two vascular systems. Commencing in this way in the substance of the tissues, by a vascular network, the minute lymphatics unite gradually with each other to form larger vessels ; and, after continuing their course for a certain distance from without inward, they enter and are distri- buted to the substance of the lymphatic glands. According, to M. Colin,' beside the more minute and convoluted vessels in each gland, there are always some larger branches, which pass directly through its substance, from the afferent to the efferent vessels ; so that only a portion of the lymph is distributed to its ultimate glandular plexus. This portion, however, in passing through the organ, is evidently subjected to some glandular influence, which may serve to modify its composition. After passing through these glandular organs, the lymphatic vessels unite into two great trunks (Fig. 43) : the thoracic duct, which collects the fluid from the absorbents of the lower extremities, the intestines and other abdominal organs, the chest, the left upper extremity, and the left side of the head and neck, and terminates in the left subclavian vein, at the junction of the internal jugular ; and the right lymphatic duct, which collects the fluid from the right ' Physiologie comparfie des Animaux domestiques, Paris, 1856, vol. ii. p. 68. 318 IMBIBITION AND EXHALATION, upper extremity and right side of the head and neck, and joins the right subclavian vein at its junction with the corresponding jugular. Thus nearly all the lymph from the external parts, and the whole of that from the abdominal organs, passes, by the thoracic duct, into the left subclavian vein. We already know that the lymphatic vessels are not to be re- garded as the exclusive agents of absorption. On the contrary, absorption takes place by the bloodvessels even more rapidly and abundantly than Tay the lymphatics. Even the products of diges- tion, including the chyle, are taken up from the intestine in large proportion by the bloodvessels, and are only in part absorbed by the lymphatics. But the main peculiarity of the lymphatic system is that its vessels all pass in one direction, viz., from without inward, and none from within outward. Consequently there is no circula- tion of the lymph, strictly speaking, like that of the blood, but it is all supplied by exudation and absorption from the tissues. The lymph has been obtained, in a state of purity, by various experimenters, by introducing a canula into the thoracic duct, at the root of the neck, or into large lymphatic trunks in other parts of the body. It has been obtained by Rees from the lacteal vessels and the lymphatics of the leg in the ass, by Colin from the lacteals and thoracic duct of the ox, and from the lymphatics of the neck in the horse. We have also obtained it, on several different occa- sions, from the thoracic duct of the dog and of the goat. The analysis of these fluids shows a remarkable similarity in constitution between them and the plasma of the blood. They contain water, fibrin, albumen, fatty matters, and the usual saline substances of the animal fluids. At the same time, the lymph is very much poorer in albuminous ingredients than the blood; The following is an analysis by Lassaigne,' of the fluid obtained from the thoracic du(5t of the cow : — Pakts per thousand. Water 964.0 Fibrin . , . 0.9 Albumen 28.0 Fat , . . • 0.4 Chloride of sodium 6.0 Carbonic \ > Phosphate and [ of soda 1.2 Sulphate ■' Phosphate of lime 0.5 1000.0 ' Colin, Physiologie comparge des Animaux domestiquea, vol. ii. p. 111. THE LYMPHATIC SYSTEM. 319 It thus appears that both the fi*briii and the albumen of the blood actually transude to a certain extent from the bloodTessels, even in the ordinary condition of the circulatory system. But this transuda- tion takes place in so small a quantity that the albuminous matters are all taken up again by the lymphatic vessels, and do not appear in the excreted fluids. The first important peculiarity which is noticed in regard to the fluid of the lymphatic system, especially in the carnivorous animals, is that it varies very much, both in appearance and constitution, at different times. In the ruminating and graminivorous animals, such as the sheep, ox, goat, horse, &c., it is either opalescent in appearance, with a slight amber tinge, or nearly transparent and colorless. In the carnivorous animals, such as the dog and cat, it is also opaline and amber colored, in the intervals of digestion, 'but soon after feeding becomes of a dense, opaque, milky white, and con- tinues to present that appearance until the processes of digestion and intestinal absorption are complete. It then regains its original aspect, and remains opaline or semi-transparent until digestion is again in progress. The cause of this variable constitution of the fluid disc*harged by the thoracic duct is the absorption of fatty substances from the intestine during digestion. Whenever fatty substances exist id con- siderable quantity in the food, they are reduced, by the process of digestion, to a white, creamy mixture of molecular fat, suspended in an albuminous menstruum. The mixture is. then absorb.ed by the lymphatics of the mesentery, and transported by them througlf the thoracic duct to the subclavian vein. While this absorption is going on, therefore, the fluid of the thoracic duct alters its appear- ance, becomes white and opaque, and is then called chyh ; so that there are two different conditions, in which the contents of the great lymphatic trunks present different appearances. In the fasting condition, these vessels contain a semi-transparent, or opaline and nearly colorless lymph; and during digestion, an opaque, milky chyle. It is on this account that the lymphatics of the mesentery are called "lacteals." The chyle, accordingly, is nothing more than the lymph which is constantly absorbed by the lymphatic system everywhere, with the addition of more or less fatty ingredients taken up from the intestine daring the digestion of food. The results of analysis show positively that the varying appear- ance of the lymphatic fluids is really due to this cause ; for though Lymph. Chyle. 9(i5.36 902.37 12.00 85.16 1.20 3.70 2.40 3.32 . 13.19 12.33 . traces. 36.01 6 85 7.11 1,000.00 1,000.00 320 IMBIBITION AND EXHALATION. • tte chyle is also richer than the lymph in albuminous matters, the principal difference between them consists in the proportion of fat. This is shown by the following comparative analysis of the lymph and chyle of the ass, by Dr. Eees:' — Water . Albumen Fibrin . , Spirit extract Water extract Fat Saline matter When a canula, accordingly, is introduced into the thoracic duct at various periods after feeding, the fluid whichia discharged varies considerably, both in appearance and quantity. "We have found that, in the dog, the fluid of the thoracic duct never becomes quite transparent, but retains a very marked opaline tinge even so late as eighteen hours after feeding, and at least three days and a half after tKe introduction of fat food. Soon after feeding, however, as we have a,lready seen, it becomes whitish and opaque, and remains so while digestion and absorption are in progress. It also becomes more abundant soon after the commencement of digestion, but diminishes again in quantity during its latter stages. We have found the lymph and chyle to be discharged from the thoracic duct, in the dog, in the following quantities per hour, at different periods of digestion. The quantities are calculated in proportion to the entire weight of the animal. Per Thousand Pabts. 8^ hours after feeding ...... 2.45 7 " " " '. 2.20 13 " " " 0.99 18 " " " 1.15 18J " " " 1.99 It would thus appear that the hourly quantity of lymph, after diminishing during the latter stages of digestion, increases again somewhat, about the eighteenth hour, though it is still considera- bly less abundant than while digestion was in active progress. The lymph obtained from, the thoracic duct at all periods coagu- lates soon after its withdrawal, owing to the fibrin whiclf it contains ' In Colin, op. cit., vol. ii. p. 18. THE LYMPHATIC SYSTEM. 821 iu small quantity. After coagulation, a separation takes place be- tween the clot and serurn, precisely as. in the case of blood. The movement of the lymph in the lymphatic vessels, from the extremities toward the heart, is accomplished by various forces. The first and most important of these forces is that by which the fluids are originally, absorbed by the lymphatic capillaries. Through- out the entire extent of the lymphatic system, an extensive process of endosmosis is incessantly going on, by which the ingredients of the lymph are imbibed from the surrounding tissues, and com- pelled to pass into the lymphatic vessels. The lymphatics are thus filled at their origin ; and, by mere force of accumulation, the fluids are then compelled, as their absorption continues, to discharge themselves into the large veins in which the lymphatic trunks terminate. The movement of the fluids through the lymphatic system is also favored by the contraction of the voluntary muscles and the respiratory motions of the chest. For as the lymphatic vessels are provided with valves, arranged like those of the veins, opening toward the heart and shutting backward toward the extremities, the alternate compression and relaxation of the adjacent muscles, and the expansion and collapse of the thoracic parietes, must have the same effect upon the movement of the lymph as upon that of the venous blood. By these different influence^ the chyle, and lymph are incessantly carried front without inward, and discharged, in a slow but continuous stream, into the returning current of the venous blood. The entire quantity of the lymph and chyle has been found, by direct experiment, to be very much larger than was previously anticipated. M. Colin' measured the chyle discharged from the thoracic duct of an ox during twenty-four hours, and found it to exceed eighty pounds. In other experiments of the same kind, he obtained still larger quantities.' From two experiments on the horse, extending over a period of twelve hours each, he calculates the quantity of chyle and lymph in this animal as from twelve, to fifteen thousand grains per hour, or between forty and fifty pounds per day. But in the ruminating animals, according to his observa- tions, the quantity is considerably greater. In an ordinary-sized cow, the smallest quantity obtained in an experiment extending over 1 Gazette Hetidomadaire, April 24, 1857, p. 2S5. ' Colin, op. oit., vol. ii. p. 100. 21 322 IMBIBITION AND EXHALATIOII. a period of twelve hours, was a, little over 9,000 grains in fifteen minutes ; that is, five pounds an hour, or 120 pounds per day. In another experiment, with a young bull, he actually obtained a little over 100 pounds from a fistula of the thoracic duct, in twenty -four hours. We have also obtained similar results by .experiments upon the dog and goat. In a young kid, weighing fourteen pounds, we have obtained from the thoracic duct 1890 grains of lymph in three hours and a naif. This quantity would represent 540 grains in an hour, and 12,690 grains, or 1.85 pounds in twenty-four hours ; and in a ruminating animal weighing 1000 pounds, this would corre- spond to 132 pounds of lymph and chyle discharged by the thoracic duct in the course of twenty-four hours. The average of all the results obtained by us, in the dog, at dif- ferent periods after feeding, gives very nearly four and a half per cent, of the entire weight of the animal, as the total daily quantity of lymph and chyle. This is substantially the same result as that obtained by Colin, in the horse; and for a man weighing 140 pounds, it would be equivalent to between six and six and a half pounds of lymph and chyle per day. But of this quantity a considerable portion consists of the chyle which is absorbed from the intestines during the digestion of fatty substances. If ^e wish, therefore, to ascertain the total amount of the lymph, separate from that of the chyle, the calculation should - be based upon the quantity of fluid obtained from the thoracic duct in the intervals of digestion, when no chyle is in process of absorption. We have seen that in the dog, eighteen hours after feeding, the lymph, which is at that time opaline and semi-transpa- rent, is discharged from the thoracic duct, in the course of an hour, in a quantity equal to 1.15 parts per thousand of the entire weight ot the animal. In twenty-four hours this would amount to 27.6 parts per thousand ; and for a man weighing 140 pounds this would .give 3.864 pounds as the total daily quantity of the lymph alone. It will be seen, therefore, that the processes of exudation and absorption, which go on in the interior of the body, produce a very active interchange or internal circulation of the animal fluids, which may be considered" as secondary to the circulation of the blood. For all the digestive fluids, as we have found, together with the bile discharged into the intestine, are reabsorbed in the natural process of digestion and again enter the current of the circulation. These fluids, therefore, pass and repass through the mucous membrane of THE LYMPHATIC SYSTEM. 323 the alimentary canal and adjacent .glands, becoming somewhat altered in constitution at each passage, but still serving to. renovate alternately the constitution of the blood and the ingredients of the digestive secretions. Furthermore the elements of, the blood itself also transude in part from the capillary vessels, and are again taken up, by absorption, by the lymphatic vessels, to be finally restored to the returning current of the venous blood, in the immediate neighborhood of the heart. The daily quantity of all the fluids, tjius secreted and reabsorbed during twenty-four hours, will enable us to estimate the activity with which endosmosis and exosmosis go on in the living body- In the following table, the quantities are all calculated for a man weighing 140 pounds. Secbeted and Reabsorbed ddrino 24 hodrs. Saliva 20,164 grains, or 2.880 pounds. Gastric juice 98,000 " " 14.000 " Bile 16.940 " " 2.420 " Pancreatic juice 13,104 " " 1.872 " Lymph 27,048 " " 3.8S4 ■' 25.036 A little over twenty-five pounds, therefore, of the animal fluids transude through the internal membranes and are restored to the blood by reabsorption in the course of a single day. It is by this process that the natural constitution of the parts, though constantly changing, is still maintained in its normal condition by the move- ment of the circulating fluids, and the incessant renovation of their nutritious materials. 324 SECRETIOIT. CHAPTER XVI. SECRETION. We have already seen, in a previous 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 difierent tissues by which they are assimilated. In this way the various tissues of the body, though they have a different 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 teaffs 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 inorgaijic 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 . • SECBETIOX. 325 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 f26 ' SKCBETION. • • has found' that ferrocyanide 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 Steho 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 absprbing 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, and 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. ' Le(;6ns de Physiologie Expgrimeiitale. Paris, 1866, tome ii. p. 96 et seq. 2 Ibid., p. 106. MUCUS. 827 The principal secretions met with in the animal body are as follo»ws : — 1. Mnona. • 6. Saliva. 2. Sebaceous matter. 7. Gastric jnioe'. 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 foUiqles or glandulae, in which the mucus is prepared; These folli- cles are most abundant in the lining membrane of the mouth, nares, pharyns, 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 dUct 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 p. j.„ 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. _ fo,..ic,.b8 op a co„- The mucus, produced in the manner rnuso un-.ova glajbclk. , 't .,-,. _- T, n ' 1 From the Iiiiman subject. (Aftei above described, is aT clear, coloriess fluid, Koiuker.)-", iiembiaDe of ti,e which is poured out in larger or smaller '»>"<:'«■ >>••-■ Epitiieimm of the 1 n n t same. ■ quantity on the surface ot 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 828 SECRETION. tough 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 mucus membrane itself from injury. The composition of mucus, according to the analyses of Nasse," is as follows :-^7 Composition of Pclmonaky Mucns. Water 955.52 Animal matter . Fat Chloride of sodium . Phosphates p{ soda and potassa Sulphates * " " Carbonates " " 33.57 2.89 6.83 1.05 0.65 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 in composition, but differ a little in the cha- racter of their principal organic ingredient, as well as in the pro- portions 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., 184(), p. 352. SEBACEOUS MATTER. 329 glands of tlie 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 with its secretion. Each follicle, as in the Fig- 103. case of the mucous glandules, is lined with epithelium, and its cavity is filled with the secreted sebaceous matter. In the Meibomian glands of the eye- lid (Fig. 103), the follicles are ranged along the sides of an es^retory duct, situated just beneath the conjunctiva, on the posterior surface of the tarsus, and opening ' upon its free edge, a little be- hind the 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 , ,, . , . Mbibomiax Glands, after of the integument Immg the meatus, in ludovic. 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.' CoMFOsiTioH OP Sebaceous Mattes. Animal substances 358 Fatty matters 368 Phosphate of lime .200 Carbonate of lime 21 Carbonate of magnesia X6 Chloride, of sodium 1 . . 37 • Acetate of soda, &c. ' 1000 Owing to the large proportion of ste^rine in the fatty ingredients of the sebaceous matters, they have a considerable degree of con- sistency. Their ofSce 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. 330 SECRETION. Fig. 104. too .rapid evaporation. When 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 hy 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. 3. Pekspibation. — The perspiratory glands of the skin are scat- tered everywhere throughout the integument, being most abundant on the anterior portions of the body. They consist each of a slender tube, about ^J^ of an inch in diamete:^ lined with glandular epi- thelium, 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 cerumi- nous glands of the ear. (Fig. 104.) A network of capillary vessels envelops the tubular coil and sup- plies the gland with the materials necessary to its secretion. The§e 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 bacls 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 perspiratory glands is not less than 2,300,000, and the length of each tubular coil, when unravelled, about yV 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 Persptbatort Gland, with its ves- sel b : magnified 35 times. (After Todd and Bow- maa.) — rt. Glandular coiL b. Fiexas of veijaeln. ' KiJllikeT, Handbuch rler Gewebelehre, Leipzig, 1852, p. 147. PEESPIKATION. 331 It is easy to understand, therefore, that a very large quantity of £uid may be supplied from so extensive a glaudular 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 13,500 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 3J pounds' weight, by both cuta- 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 Ansel- mino,' is as 'follows : — ■ CoMFOSITIOir OF THE PeBRFIBATION. Water 995.00 Animal matters, with lime . .IQ Sulphates, and substances soluble in water .... 1.05 Chlorides of sodium And potassium, and spirit-extract . . 2.40 Acetic acid, acetates, lao^tes, 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 ' Milne Edwards, Leijons snr la Physiologie, &o., vgl. ii. p. 623. ' Piiilosophy of Health, London, 1838, chap. xiii. s Simon. Op. cit., p. 374. 832 SECRETION. 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 inconvenience, 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 pprtion 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. Atiee's translation, Phila., 1854, p. 25. THE MILK. 333 surfaces of the cornea and conjunctiva moist and polished, and to preserve in this way the transparency of the parfs. 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 which would follow from its accumulation and putrefactive alteration. 5. The Milk. — The mammary glands are conglomerate glands resenibling 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 Fig. 105. their successive union, they form larger branches and trunks, until they are reduced in numbers to some 15 or 20 cylindrical ducts, the lactifer- ous 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 contalin 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 amdunt 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- ' Lehmann, op. oit., vol. ii. p. 63. Gr.ANDULAR Structcre of .Mamma 334 SECRETION. btilar or oval bodies, from ttbtt to sin o? an incli in diameter, ■which are termed the "colostrum corpuscles." (Fig. 106.) These bodies are more yellow and Fig- 106. opaque than the true milk- glob ules, as well as being very much larger. They have or 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 j^^j^ to tbVtt of ^^ i^oli ill dia- meter. At the end of a day or two after 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 ijipple, 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 : — Colostrum CoKPrscLES, Trith mllk-globulea ; from a woman, one day after delivery. .*««■' Composition OP Cow'b Milk. Wate* Casein Butter Sugar Soda Chloriiles of sodium and potassium Phosphates of soda and potassa Phosphate of lime " " magnesia " iron . Alkaline carbonates . 870.2 ^14.8 31.3 47.7 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. THE MILK, 335 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- 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 the teniperature of the body, and have a perfectly circular out- line. In the perfect milk, they are very much more abundant and Fig. 107. To" '0° o°o 0. 0„=°„oo Ooq"o'-'^<§) o o o. oO°0°0 ^OO", ooo°oOOoO°oS°°° Oo ^°i,po ; o oo oO, oOo Mii.k-GlobuTiKe>; from same woman as above, four days after delivery. Secretion fully etstablislied. 336 SECRETION. smaller in size than in the colostrum ; as the largest of them are not over jtsVb of an inch in diameter, and the greater number about YsUu 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 butjrine, which gives the peculiar flavor to the butter of milk. The milk-globules have sometimes been described as if each one were separately covered with a thin layer of coagulated casein or albumen. Nosucli 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 (Cj^Hj^Oj^) may he 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 potassa. After the neutralization of these substances has been accomplished, the milk loses its alkaline reaction and begins to turn sour. The fr^e lactic acid then coagulates the casein, and the milk SECEETION OF THE BILE. 337 is curdled. The altered organic matter also acts upon the olea- ginous 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 fir^t 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 qttantity of arterial blood with which it is supplied is extremely small in comparison with that which it receives from ' Op. oit., p. 337. 22 338 SECRETION. Fig. 108. the portal system. The blood which has circulated through the capillaries of the stomach, spleen^ pancreas, and intestine is col- lected bj the roots of the corresponding veins, and discharged into the 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 tAtt of ^^ inch in diameter. These veins, with their terminal branches, are arranged in such a manner as to include between them pentagonal or hexagonal spaces, or portions of the hepatic substance, -j'g to -j-'j 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 "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 the commencing rootlets of the hepatic ducts. As the portal vein, the he- patic artery, and the hepatic duct enter the liver at the transverse fissure, they are closely invested by a fibrous sheath, termed 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 in vestment,, 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 interlobulaE? : spaces ; so that here the lobules are nearly in contact with each Ramification of Poetal Vein in Liter. — a. Twitf of poi'lalvein. &,&, Interlobular veiue. c. Acini. SECRETION OF THE BILE. 339 Other by their adjacent surfaces, being separated only by the inter- lobular veins and the minute branches of the hepatic artery and duct previously mentioned. From the sides of the interlobular veins, and also from their terminal extremities, there are given off 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. LoBuiE OF Liter, ehowing distilbutlon of bloodvessels ; magnifled 22 diameters —a. n. In- terlobular veins, b. latralobnlar vein, a, c, o. Lobular capillary plexus, d, d. Twigs ot inter- lobular vein passing to adjacent lobules. converging capillaries unite into a small vein (6), the " intralobu- lar" vein, which is one of the commencing rootlets of the hepatic vein. These rootlets, uniting successively 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 TuVir 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 340 SECKETIOX. of the cell, sometimes more or less obscured by 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, 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 glaindular cells of the organ, and gives up to them the nutritious materials which are afterward converted into the ingredients of the bile. Hepatic Cklls. Frum the human subject. ' American Journal Meil. Sol., Januarjr, 1848. ' On Some Points in the Minute Anatomy of tlie Liver. London, 1856. EXCRETION, 341 CHAPTER XVII. EXCRETION. » Wk 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 renewal 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, h^matine and the like, 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 342 EXCBETION. ■ , under which 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 capaWe 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 of 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 dischai^ged and expelled from the body. This entire process, made up 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 upcrti 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 frequent 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 which 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 UREA. 343 four days to produce a fatal result. A fatal result,- however, is cer- tain to follow, at I 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 ejdst in the human body are as. follows: — co„ 1. Carbonic acid . 2. Urea . 3. Creatine . 4. Creatinine 5. Urate of soda 6. Urate of potassa 7. Urate of ammonia CsHsNjO, CsH,N302 NaO.CjHNjOj+HO KG.CsHN^Oj . NH40,2C5HNjOj4-HO The physiological relations of carbonic acid have already been studied, at suf&cient length, in the preceding chapters. The remaining excrementitious substances may be examined together with the more propriety, since they are all ingredients of a single excretory fluid, viz., the urine, i Ukea. — This is a neutral, crystallizable, nitrogenous substance, very readily soluble in water, and easily decomposed by various extemial influences. It occurs in the urine in- the proportion Fig- m- of 30- parts per thousand ; in the blood, according to Picard,* in the proportion of 0.16 per thousand. The blood, how- ever, is the source from which this substance is supplied to the urine; and it exists, ac- cordingly, in but small quan- tity in the circulating fluid, for the reason that it is constantly drained off by the kidneys. But if the kidneys be extir- pated, or the renal arteries tied, or the excretion of urine sus- pended by inflammation 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 circumstances, in the propor- TTeea, prepared from arine.and crystallized bv sloir evaporation. (After Lehmann.) ' In Milne Edwards, Lecjons snr la Fhyslologie, &e., Tol. i. p. 297. 344 EXCRETION. tion of 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 is 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 purpose the fresh urine is evaporated 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 decom- posed 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 crystal- lized 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 : — C2HjNj02=Urea. 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 in this way. In order that the conversion of the urea be thus pro- duced, 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. Prof. "William A. Hammond,' who is a very large man, by similar experiments found it to be ' Robin and Verdeil, vol. il. p. 502. ' American Journal Med. Soi , Jan., 1855, and April, 1856. UREA. 345 670 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 appe^arance, and afterward increases in quantity with the development of the 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 ho 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. Hammond has also shown that continued mental applica- tion will raise the quantity of urea above its normal standard, though the muscular system remain comparatively inactive. 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 op Food. Daily Quantity op Urea. Animal . 798 grains. Mixed . ..." 4S7 " Vegetable ........ 337 " Non-nitrogenous 231 " 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 J to 10^ P. M. Urea exists in the urine of the carnivorous and many of the ' New York Jonmal of Medicine. March, 1856. ' Robin and Verdeil, vol. ii p. 500. » Lehmnnn, op. eit., vol. ii. p. 163. * I>oc. cit. 346 EXCRETIOIf. Creatine, crystallized from hot water. (After LehmauuJ herbivorous quadrupeds ; but there is little or none to be found in that of birds and reptiles. Ceeatine. — This is a neutral crystallizable substance, found in the muscles, the blood, and the. urine. It is soluble in water, very slightly soluble in alcohol, and Fig. 112. not at all so in ether. By boil- ing with an alkali, it is either converted into carbonic acid and ammonia, or is decomposed with the production of urea and an artificial nitrogenous crys- tallizable substance, termed sar- cosine. By being heated with strong acids, it loses two equiva- lents of water, and is converted into the substance 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 small pieces, treating it with distilled water, and subjecting it to pressure. Creatine evidently Fig. 113. originates in the muscular tis- sue, is absorbed thence by the blood, and is finally discharged with the urine. Creatinine. — This is also a crystallizable substance. It dif- fers in composition from crea- tine by containing two equiva- lents less of the elements, of water. It is more soluble in water and in spirit than crea- tine, and dissolves slightly also in ether. It has a distinctly Cruati {After Lehmaun.) INE, crystallized from bot water. CREATININE. — URATE OF SODA. 347 alkaline reaction. It occurs, like creatine, in the muscles, the blood, and the urine ; and is undoubtedly first produced in the muscular tissue, to be discharged finally by the kidneys. It is very possible that it originates, not directly from the muscles, but indirectly, by transformation of a part of the creatine ; 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, fur- thermore, while creatine is mores 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. Urate of Soda. — ^As- its name implies, this substance is a neu- tral salt, formed by the union of soda, as a base, with a nitrogenous animal acid, viz., uric acid (CjHNjO^jHO). 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 a free state, in a natural condition of the fluids. When present, it has always been produced by decom- position 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 globu- F'g- 114. lar masses, with projecting, curved, conical, wart-like excrescences. (Fig. 114.) It dissolves readily in the alka- lies ; and by most acid solu- tions it is decomposed, with the production of free uric acid. Urate of soda exists -in the urine and in the blood. It is either produced origin- ally in the blood, or is formed in 'some of the solid tissues, and absorbed from them by the circulating fluid. It is constantly eliminated by the kidneys, in company with the other ingredients of the urine. The average daily quantity of urate of Urate op Soda; from a arinaiy deposit. 348 ■ EXCRETIOX. soda discliarged by the healthy human subject is, according to Lehmann, about 25 grains. This substance exists in the urine of the carnivorous and omnivorous animals, but not in that of the her- bivora. In the latter, it is replaced by another substance, differing somewhat 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 her- bivorous, the urate of soda disappears, and is replaced by the hip- purate. Urates of Potassa and Ammonia. — The urates of potassa and ammonia resemble the preceding salt very closely in their physio- logical 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 closely resemble each other in 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 GENERAL CHAKACTEBS OF THE UBINE. 349 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 are 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 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 prdinary quantity of urine discharged daily by a healthy adult is about 3xxxv, and its mean specific 350 EXCRETION. 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 proportions of water and solid ingredients entering into its constitution. Ordinarily the water of the urine is more than sufficient to hold all the 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 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 off 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 tbe 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 regard to each other ; the former increasing while the latter diminishes, and vice versd. 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 certain 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 DIURNAL VARIATIONS OF THE URINE. 351 quantity of water is often so mncli diminished tliat 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 first discharged in the morning, is usually dense, highly colored, of a strongly acid reaction, and a high specific gravity. That passed during the forenoon is pale, and of a low specific gravity, 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 strongly 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 : — Obseevation First. March 20th. Urine of Ist diapharge, acid, sp. gr. 1025. " 2d " alkaline, " 1015. " 3d " neutral, " 1018. " 4th " acid, « 1018. « 5th " acid, " 1027. Observatios Second. . March 21st. Urine of 1st discharge, acid, sp. gr. 1029. " 2d " nentral, " 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 12.45 352 EXCRKTIOX. 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 : — Composition op the Ueihe. Water 938.00 Urea 30.00 Creatine 1'25 Creatinine 1-50 Urate of soda -i " potaasa \ 1.80 " ammonia J Coloring matter and 1 .... .30 Mueus > Biphosphate of soda Phosphate of soda " potassa " magnesia " lime Chlorides of sodium and potassinm 7.80 Sulphates of soda and potassa 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 certain 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 a very small quantity. The coloring matter, or urosacine, is in solution in a natural condition of the urine, but it 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 mucus of the urine comes from the lining membrane of the EEACTIONS OF THE UKINE. 853 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. 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 form- ation ; 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 potassa, 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 23 354 EXCRETION. 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, potassa, &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 united with them 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- F>g- 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 potassa or soda, be added to the urine so as to neutralize its acid reaotion, 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. Beside 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 pi interfering with the mutual reaction of starch and iodine, and feven TJric Acid ; deputiited from urine. REACTIONS OF THE UBINB. 855 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 3j of iodine water be mixed with a solution of starch, it strikes an opaque blue color ; but if 3j 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 be mixed with four or five times its volume of iodine water, and starch be subsequently added, no union takes place between the starch and iodine, and no blue color is produced. In these instances, the icfdine 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 potassa be afterward added, the mixture takes a dingy, grayish-blue color. On boiling, the cgloi turns yellowish 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 subsljinces are retained upon the filter, while the sugar passes through in solution, and may then be detected as usual by Trommer's test. Accidental Ingredients 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 ^^y a kind of filtration, it is not surprising that most medicinal and poisonous substances, introduced into the circulation, should be ' American Journal Med. Sei., April, 1856. 356 EXCRETION". 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 the Sidneys, either in their original form, or after undergoing cer- tain chemical modifications. The salts of the organic acids, such as lactates, acetates, malates, &c., of soda and potassa, 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 administra- tion 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 potassa. The pure alkalies and their carbonates, according to the same ob- server, produce a similar effect. Bicarbonate of potassa, 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 dissol«?ing 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 deposit. Ferrocyanide of potassium, when introduced into the circulation, appears readily in the urine. Bernard' observed that a solution of ' Physiological Chemiaty, vol. ii. p. 133. ' Lemons de Physiologie ExpSrimentale, 1856, p. 111. ACCIDENTAL INGREDIENTS OF THE URINE. 357 this salt, after being injected into tlie 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, iodi|ie 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 freely, one of them for two months, the other for six 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. Etiier passes out of the circulation in the same way. We have observed the odor of this substance very perceptibly in the urine, after it had been inhaled for the purpose of prpducing anaes- thesia. The bik-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 in- jected and the kind of sugar used for the experiment. If a solution of 16 grains of glucose be jnjected into the areolar tissue of a rabbit weighing a little over two pounds, it is entirely destroyed in the circulation, 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 ■ Le?ons de Phys. Exp., 1855, p. 214 et seq. 358 EXCRETION. sugar used makes a difference in this respect. For while 15 grains of glucose may be injected without .passing out by the kidneys, 7 1 grains of cane sugar, introduced in the same way, fail to be com- pletely destroyed in the cirtiulation, and may be detected in the urine. In certain forms of disease (diabetes), where sugar accu- mulates in the blood, it is eliminated by the same channel ; and a saccharine condition of the urine, accompanied by an increase in its quantity and specific gravity, constitutes the most characteristic 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 suf&cient 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'^ disease, the same symptom occurs, and remains as a permanent condition. In all such instances, however, as the above, where foreign ingredients 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 tilne, 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 qf 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 substance, is liable to putrefaction; and if the temperature to which it is exposed be between 60° and 100° F., it soon becomes altered, ACID FERMENTATION OF THE UEINE. 359 and communicates these alterations more or less rapidly to the super- natant fluid. The first of these changes is called the acid fermenta- tion 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 urine, as we have already stated, contains no free acid, its acid reaction with 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 acid when first passed, may frequently present traces of this substance after some hours' expo- sure 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 freqiiently 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 oxalip acid is also sometimes produced, in a similar manner with tjie lactic. It is very certain that the deposit of oxa- late of lime, far from Tseing 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 either with the kidneys or with 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 360 EXCKETION'. Fig. lie. 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 supernatant fluid remains clear. These crystals are of minute size, transparent, and colorless, and have the 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 decom- position or metamorphosis of urea into carbonate of ammonia. As the alteration of the mucus advaiPces, it loses the power of pro- ducing lactic and oxalic acids, and becomes a ferment capable of acting by cataly^s 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 its mucus, it Oxalate op Lime; deposited from liealtby urine, during the acid fermentation. '' ALKALINE FERMENTATION OF THE URINE. 361 may be afterward kept, for an indefinite time, -witliout alteration. But under ordinary circumstances, the mucus, as soon as its putre- faction has commenced, excites the decomposition of the urea, and carbonate of ammonia begins to be developed. The first portions of the ammoniacal salt thts produced begin to neutralize the biphosphate of soda, so that the acid reaction of the urine diminishes in intensity. This 'reaction gradually becomes weaker, as the fermentation proceeds, until it at last disappears altogether, and the urine becomes neutral. The production 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 is 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 £airly 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 Ijarbonate of ammonia on the phosphates of 362 EXCBETIOSr. soda and magnesia. One of these is the "triple phosphate," phos- phate of magnesia and ammonia (2MgO,]SrHp,POj+2HO). The other is the phosphate of soda and ammonia (Na0,NH40,H0,P0j+ 8H0). The phosphate of magnesia and ammonia is formed from the phosphate of magnesia in the urine (3MgO,POj4-7HO) by the replacement of one equivalent of magnesia by one of am- monia. The crystals of this salt are very elegant and charac- teristic. They show themselves throughout all parts of the mix- ture ; growing gradually in the mucus at the bottom, adhering to the sides of the glass, and scattered abundantly over the film which collects upon the surface. . By their. refract- ive power, they give to this film a peculiar glistening and 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.) Frequently, 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 manner 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 quadrangular form, or some figure derived from it. They are intermingled 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 Phosphate of Magnrsia and Ammonia; deposited from healthy urine, during all^aline termea- tation. EENOVATION BY NUTBITIVE PROCESS. 363 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 fl-eely given ofi' 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. Eenovation of the Body by the Nutritive Process. — We , can now estimate, from the foregoing details, the entire quantity of material assimilated and decomposed by the living body. For we have already seen how much food is taken into the alimentary canal and absorbed by the blood after digestion, and how much oxygen is appropriated!^ from the atmosphere in the process of respiration. We have also learned the-amount of carbonic acid evolved with the breath, and that of the various excretory substances discharged from the body. The following table shows the absolute quantity of these different ingredients of the ingesta and egesta, compiled from the results of direct experiment which have already been given in the foregoing pages. Absorbed dubinq 24 HOURS. DiSCHAHOED DUUIlfG 24 HOUBS. Oxygen . 1.019 lbs. Carbonic acid . 1.535 lbs Water 4.735 " Aqueous vapor 1.155 " Albuminous matter . .396 " Perspiration . 1.930 " Starch . .660 " Water of the urine 2.020 " Fat . .220 « • Urea and salts .110 " Salts .040 " Feces .320 •• 7.070 7.070 Eather more than seven pounds, therefore, are absorbed and dis- charged daily by the healthy adult human subject ; and, for a man having the average weight of 140 pounds, a quantity of material, equal to the weight of the entire body, thus passes through the system in the course of twenty days. It is evident, also, that this is not a simple phenomenon of the passage, or filtration, of foreign substances through the animal frame. The materials which are absorbed actually combine with the tissues, and form a part of their substance ; and it is only after undergoing subsequent decomposition, that they finally make their appearance in the excretions. None of the solid ingredients of the food are discharged under their own form in the urine, viz., as 364 EXCRETION. starch, fat, or albumen ; but they are replaced by urea and other crystallizable substances, of a different nature. Even the carbonic acid exhaled by the breath, as experience has taught us, is not pro- duced by a direct oxidation of carlJon ; but originates by a steady process of decomposition, throughout the tissues of the body, some- what similar to that by which it is generated in the decomposition of sugar by fermentation. These phenomena, therefore, indicate an actual change in the substance of which the body is composed, and show that its entire ingredients are incessantly renewed under the influence of the vital operations. SECTIO?^ II. NERYOUS SYSTEM chAptek I. GBNEEAL STEUCTURB AND FUNCTIONS OP THE NEBVOUS SYSTEM. In 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 ( 365 ) 366 GENEBAL STRUCTURE AND FUNCTIONS parts of the body, and to cause them to act in harmony with each other. This object may be more fully exemplified as follows : — Bach organ and tissue in the body has certain properties peoiiliar 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 irritability. We' have often had occasion to notice this pro perty of irritability, in experiments related in the foregoing pages, We have seen, for. ejcample, 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. Very often, two organs which are situ- ated in distant parts of the body are connected with each other by such a sympathy, that the activity of one is influenced by the condition of the other. 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 adjacent or remote. Thus the peristaltic action of the mus- cular coat of the intestine commences when the food is brought in OF THE NERVOUS SYSTEM. 367 contact with its mucous membrane. The lachrymal gland is excited to increased activity by anything -which causes irritation of the conjunctiva. 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 stimulus applied to one organ may excite the activity of 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 harmony with each other, and at the right time, and in the right direction. 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 disturb- ance must inevitably follow. When the muscular system is ex- cited by unusual 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 nerv- ous system, there would immediately follow deficiency of aeration, vascular congestion, and derangement of the circulation. 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, wtiere 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. 368 GENERAL STRUCTURE AND FUNCTIONS 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 distinguished from each other by their color, their structure, and their mode of action. One of these is known as the white substance, or the fibrom tissue. It constitutes the whole of the substance of the nervous trunks and branches, and is found in large quantity on the exterior of 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 diam^eter of xT^uTy of an inch. In the trunks and branches of the nerves they average jtjVt 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 very similar to it in refractive power. In the interior of this tubular membrane there is contained a thick, semi-fluid nervous matter, which is white and glistening by reflected light, and is called the "white substance of Schwann." Finally, running longitudinally 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 external tubu- lar membrane. When nervous matter is prepared for the microscope and exa- mined by transmitted light, two remarkable appearances are ob- served 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 OF THE NERVOUS SYSTEM. Fig. 118. preparation, produces an irregularly bulging or varicose appearance in them at various points, owing to the readiness witii which the semi-fluid white substance in their interior is displaced in diflerent directions. (Fig. 11&.) 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 mem- brane may be still seen, pass- ing across from one portion to the other. When a nerv- ous filament is torn across under the microscope and subjected to pressure, a cer- tain 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), scattered over the field of the microscope. The varicose appear- ance above alluded to is more frequently seen in the smaller nerv- ous filaments from the brain and spinal cord, owing to their soft consistency and the readiness with which they yield to pressure. The second efiect produced by the artificial preparation of the nervous matter is a partial coagulation of the white substance of Schwann. 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 whitf: " substance of Schwann. The coagulating process, however, subsequently goes on, and 24 WFBVOua Filaments froin white aubstance of braia, — a, u, a. Sort subntance of the fllamenta preHHCj} out, and floating in irregulurlj rounded drops. 370 GENERAL STBUCTCRK AND FUNCTIONS gradually advances from the edges of the filament toward its centre, until^ its entire thickness after a time presents the same appearance. The effect of this process can also be seen in those portions of the white substance which have been pressed out from the interior of the filaments, and which float about in the form of drops. (Fig. 118, a.) These Pig.. 119. drops are always covered with a layer of coagulated material which is thicker and more opaque in propor- tion to the Jength 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 condensed 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 substance 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. Thesfs pri- mary bundles are ag^dn united into secondary, the secondary into NEi^Togs FtT.AMEifTs from sciaticnerve, i^howJug their cuagulntioii. — At li, the torn extremity of a nervous filament with the axis cylinder {h) protruding from it. At c, the white substance of Sohwaan is nearly separated by accideotai compression, but the axis- cylinder 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 NKBVOUS SYSTEM. 871 Fig. 120. tertiary, &o. A nerve, therefore, consists of a large bundle of ulti- 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 branched 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 originates, for ex- ample, from the spinal cord in the region of the neck, and runs down the upper extremity, dividing and subdividing, to be finally distri- buted to the integument and mus- cles of the hand, contains at its point of origin all the filaments' into which it is afterward divided, and which are merely separated at successive points from the main bundle. ' The ultimate fila- ments, accordingly, are continu- ous throughout, and do not them- selves divide at any point between their origin and their final distri- bution. 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 Division of a Nrkvk, phowlnj? portion of nervous trunk ("), aud the separation of its filaments (ft, c, d, e). 372 GKNEBAL STBUCTCRK AND FUNCTIONS origin to its termination, and will always tave the same character and function in every part of its course. The other variety of nervous matter is known as the gray sub- stance. It is sometimes called " cineritious matter," and sometimes Fig. 121. InoHCulation of N b r t k s . "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 Fig. 122. NRRT8 Cbli.b, Intermingled with fibres; from Bemilunar ganglion of cat. the external portions of the cere,brum and cerebellum. It also constitutes the sub- stance of all the ganglia of the great sympathetic. Ex- amined by the microscope, it consists of vesicles or cells, of various forms and sizes, im- bedded in a grayish, granular, intercellular substa'nce, and containing, also, very fre- quently, granules of grayish pigmentary matter. It is to the presence of this granular pigment that this kind" of OF THE NERVOUS SYSTEM. 373 nervous matter owes the ashy or " cineritious" color from which it derives its name. The cells composing it vary in size, according to Kolliker, from ^Atr to ^^^ 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 sometimes divided into two or three smaller branches. These c6lls are inter- mingled, in all the collections of gray matter, with nervous filaments, and are entangled with their extremities in such a manner that it is exceedingly difficult to ascertain the exact nature of the anato- mical relations existing between them. - It is certain that in some instances the slender processes running out from the nervous vesi- cles 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 Fig. 123. 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, so as to form a circular series of similar parts, each organ being repeated at difierent nbrtoos system of staefibh. 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 immedi- 374 GENERAL STBUCTUBE AND FUNCTIONS ately beneath it the stomach or digestive cavity, which sends pro- longations into every one of the projecting limbs. There is also contained in each limb a portion 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 con- nected with each other by commissures, so as to form a nervous collar or chain, surrounding the orifice of the digestive cavity. Each gailglion 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 nianner : — 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 gahglion 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 myscular 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 repollected that this action does not necessarily indicate any sensation or volition, nor even any consciousness on tbe 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 p&rception. 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 ^nd contain the same organs, their action will also be the same ; and the effects of this communication of the stimulus from one to OF THE NERVOUS SV8TEM. 375 the other by means of commissures -will be a repetition, or rather a simultaneous production of similar movements in different parts 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 caWed sensitive fibres. The othet set run from the ganglion, to the muscles, and carry the nervous impression outward. These are called motor fibres. "In the starfish, where 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 orgians, situated in dif- ferent regi6i3S,'we find th'at ' the nervous ' ganglia, presiding over the function of these organs, present a corresponding degree of dissimilarity. In Aplysia, 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 some- what convoluted intestine. The liver is large, and placed on one side of the body, while the gills, in the form of vascular laminffi, 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 mantle, which ex- pands at the ventral surface into a tolerably well-developed "foot," or organ of locomo- tion, by which the animal is enabled to change its position and move from one nbrtocs ststhm op ■t -I'. , .y Api.tbia. — 1. Dij^estive or locality to another. oesophageal ganRlion. 2. C«re. The nervous system of this animal is con- *"*' ganglion. 3, 3. Podai .,r , . , locomototy ganglia. 4. Eespi- structed upon a plan correspondmg with ratoiygangiioW. Fig. 124. 376 GESEEAL STBOCTUBE ANDi FUNCTIONS Fig. 125. that of the entire body. (Fig. 124.) There is a small gianglion (i) situated anteriorly, which sends nerves to the conimencement of the digestive apparatus, and is regarded as the oesophageal or digestive ganglion. Immediately behind it is a larger one (2) ca,lled 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 locoraotory ganglia, which supply the muscular mantle and its foot-like expansion, and which . regulate the movement of these organs. Finally, another ganglion (4), situ- ated 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 accord- ingly 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 centipedes, or scolopendra. Here the body is composed of twenty-two successive and nearly similar articu- lations, each of which has a pair of legs attached, and contains a portion of the glandular, respira- tory, digestive and reproductive apparatuses. The animal, therefore, consists of a repetition of similar compound parts, arranged in a longitudi- nal chain or series. The only exceptions to this similarity are in the first and last articulations. 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 Nl^RTOlTS STBTEK OF Centipede, OF THE NERVOUS SYSTEM. 377 the alimentary canal. Each pair of ganglia is connected with the integument and muscles of its own articulation by sensitive and motor filaments ; and with those which precede and follow hy a double cord of longitudinal commissural fibres. In the first articu- 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 im- pression received by 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 backward, and produce simultaneous movements in several neighboring articula- tions ; 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 ventra,l gan^ia, 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 oeeupyinig 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 378 GENERAL 8TRUCTUBK AND FUNCTIONS 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 kmount 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 commis- sures so as to form a compound mass. As the organs of locomo- tion, furthermore, instead of being distributed, as in the centipede, throughout 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 secre- tory and generative functions are situated together, in the cavity of the abdomen. 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 different 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, together 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 unsymmetrical and irregular, as in the molluscs. The organs of respiration, however, are nearly symmetl-ical 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 struc- tural arrangement of the organs under its control. That portion which presides over the locomotory, respiratory, sensitive, and in- tellectual functions forms a system by itself, called the cerebrospinal system. This system is arranged in a manner very similar to that of the articulata. It is composed of two equal and symmetrical halves, running along the' median line of the body, the different OF THE NERVOUS SYSTEM. 379 parts of ■wliicli 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. 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 heck and thorax; but is uusymmetrical in the abdomen, where it attains its largest size and its niost complete developmdnt- 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 system 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 ani- mal, very much greater than that contained in the system of the great sympathetic; and this, preponder- ance increases, in the higher classes, just in proportion to their supe- riority in intelligence, sensation, power of motion, and other func- tions 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. CSREBRO-BPfNAL ST8TEH OF MAN. ^1. Corehrum. 2. Cereb'-llnm. 3,3,3. Spin.tl cord and nerveA. 4, 4. Brachial nerveii S, fi. Sacral nerves. 380 GENEBAI, STRUCTURE AND FUNCTIONS by an anterior and, posterior median fissure, into two lateral halves, which still remain connected with each other by a central mass or commissure. Its inner portions are occupied by gray, matter, which forms a continuous ganglionic chain, running from one ex- tremity of the cord to the other. Its oijter portions are 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 classeSj 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 remiarkable plexuses, before entering their corre- spond ing limbs, viz., the bra- ^'^' ^^^' chial plexus above, and the sacral plexus below. The • cOrd itself, moreover, pi-e- sents two enlargements at the point of origin of these nerves, viz., the cervical en- largement from which the brachial nerves (4, 4) are given off, and the lumbar TranBTerso Section of Spikaj, CoBn.-n,*. Spinal enlargement from wMch the nerves of riglit and left Bides Rho^rlnt,' tlielr two rootB. gaCral ncrVCS (s, s) Originate, rf. Origin of anterior root. «. Origin of posterior root. Ti? T. ■ l J ^ioned. By the eruxphahn 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, 382 GENERAL gTRUCTUBE AND FUNCTIONS Fig. 128. 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 o/nervous ganglia, connected with each other and with the spinal cord hy transverse and longitudinal commissures. . The number and relative size of these ganglia, in different kinds of animals, depend upon the perfection of the hodily 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 thei more complex. The brain of the Alligator (¥\g. 128) consists of five pair of ganglia,;ranged one behind the other in the interior of the cranium. The first of theSe.are iwoi rounded masses (i), lying just above and behind the nasal cavities, which distri- bute their nerves upon the Schneiderian .mucous membrane. These are the olfac- tory ganglia. The f are connected with tl^e rest of the brain by two long and slender commissures, the " olfactory com- rtissures." The next pair (s) are some- what larger and of a triangular shape, .wheR. viewed from above downward. They are termed the "cerebral ganglia," or the hemispheres. Immediately follow- ing them are two quadrangular masses (a) which give origin to the optic nerves, and are therefore called the optic ganglia. They are termed also the " optic tiiber- cles ;" and in some of the higher animals, where they present an imperfect division into four nearly equal parts, they are known as the " tubercula quadrigemina." Behind them, we have a single triangular collection of nervous matter(4), which is called the cerebellum. Finally, the upper por- tion 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 (s), 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 "pneumogastric" or " respiratory'* ganglia. It will be seen that the posterior columns of the cord, as they GATOR.— 1, Bbaik of a factory ganglia. 2. Hemispheres. 3. Optic tubercles. 4. Cerebellum. 5. Medulla oblongata. OP THE NERVOUS SYSTEM. 383 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 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 successive ganglia which compose the brain, being arranged in pairs as above described, are separated from each other on Jhe 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 cerebel- lum 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 throughout. In birds, the hemispheres are of much larger size than in rep- tiles, and partially conceal the optic ganglia.. The cerebellum, also, is very well developed in 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 here extends so far backward as almost completely to conceal the medulla oblongata and the fourth ven- tricle. In the quadrupeds, the hemis- pheres and cerebellum attain a still greater size in proportion to the remaining parts of the brain. There are also two other pairs of ganglia, situated beneath the he- mispheres, and between them and the tubercula quadrigemina. These are the corpora striata in front and the opiic thalami hehini. In Fig. 129 is shown the brain of the rabbit, with the heqiispheres laid open and turned aside, so as to show the internal parts in their natural situatiou. The olfactory ganglia are seen in front (i) con- Brain op Rabbit, viewed from above. — 1. Olfactory gant^lia. 2. HemifipliereB, tara(>d aside. 3. Corpora Ktriata. . 4. Optic thalarni. 6. Tubercula quadrigemica. 6. Cerebellutii. 384 GENERAL STBUCTUBE AND FUNCTIONS nected with the remaining psnts by the olfactory commissures. The separation of the hemispheres (s, s) shows the corpora striata (s) and the optic thalami (4). Then come the tubercula quadrigemina (i), which are here composed, as above mentioned, of four rounded masses nearly equal in size. The cerebellum (e) is considerably en- larged 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 nume- rous 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 best be studied in the following order. As the spinal cord, in the human subject, passes upward into the cranial cavity, it en- larges into the medulla oblongata as already described. The medulla oblongata presents on each side three projections, two anterior a,iid one posterior. The middle projections on its anterior surface (Fig. 130, 1, 1), which are called the anterior pyramids, 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 external surface, to terminate in the gray matter of the hemispheres. The projections immediately on the outside of the anterior pyramids, in the medulla oblongata, are the olivary Mies (i, 2). They contain in their in- Fig. 130. Medtm^a OBtasnATA OF Hi'HAN Brain, aote- rior view — 1, 1. Anterior py- ramids: 2,2. Olivary bodies. '3, 3. Restil'tfrm bodies. 4. De- cii-ssatibn of 'the anterior co- iiimiis. Tlie medulla oblonfr- ftta is seen terminated above by the transvertie fibres of the pons Varolii. OF THE NERVOUS SYSTEM. 385 terior 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. The anterior columns of xhe 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 each side of the fourth ventricle, form the posterior and lateral projections of the medulla oblongata (3, 3). They are sometimes called the " res- tiform 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 25 GENKEAL STRUCTURE AND FUNCTIONS or tuberosity, which is known by the name of the tuber annulare. In the deeper portions of this protuberance there is situated, among the longitudinal fibres, another collectM)n of gray matter, which though not of large size, has very important functions and connec- tions. This is known as the ganglion of the tuber annulare. Situated almost immediately above these parts we have the cor- pora striata in front, and the optie thalami behind, nearly equal in size, and giving passage, as abov% described, to the fibres of the anterior and posterior columns. Behind them still, and on a little lower level, are the tuber^ula quadrigemina, giving origin to the optic nerves. The offactory 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 ganglia occur in tlie following order, counting from before back- ward: 1st. The olfactory gan- glia. 2d. The cerebrum or hemi- spheres. 8d. 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 Fig. 131. Diagram of Humah Bbaiit, in vertical sec- tion ; showing the situation of the different gan- glia, and the course of the fibres. 1. Olfactory ganglion. 2. Hemisphere. 3. Corpus striatum. 4.. Optic thalamus. 5. Tubercula quadrigemina. 6. Cerebellum. 7. Ganglion of tuber annulare. 8. Ganglion of medulla oblongata. OF THE NERVOUS SYSTEM. 387 to bef seen in tlie accompanying figure. A portion of the anterior fibres, we have already observed, pass upward and backward, with- the restifbrm bodies, to the cerebellum ; whUe the remainder run forward through the tuber annulare and the corpus striatum, and- then radiate to the gray matter of the cerebrum. The posterior fibres, constituting the restiform body, are distributed partly to the cerebellum, and then pass forward, as previously described, under- neath the tubercula quadrigemina to the optic thalmi, 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." It 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" hfis 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, besides supplying the corresponding parts of the head, preside also over the organs of special sense, and perform various other functions of a purely nervous cliaracter. 388 OF NEEVOUS IBBITABILITY CHAPTEE II. OF NERVOUS IRRITABILITY AND ITS MODE OP ACTION. "We have already mentioned, in a previous chapter, that every organ in the body is endowed with the property of 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 muscular fibre itself. The existence of muscular irritability cannot be ex- plained by 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 AND ITS MODE OP ACTION. 389 these properties to a certain extent for some time afterward. It is only when the constitution of the tissues has become altered by 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 fibres 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 consider- ably 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 in birds the muscular irritability continues only a few minutes after the death of the animal. In quadrupeds it lasts somewhat longer ; while in reptiles it remains, under favorable cir- Fig- 132. cumstances, 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 nutrition going on with rapidity, the constitution of the muscular fibre becomes so rapidly altered after the circulation has ceased, that its irritabibty soon disappears. 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 cir- culation can therefore be dispensed with for a longer period, before the constitution of the tissues be- comes so much altered as to destroy altogether froo's lkg, wuh their vital properties. i""'"' "' sawanic bat- _. ,. I- • ft 1J1.1JJ '"y applied to the Owing to this peculiarity or the cola-blooaea nmsciea at a, s. 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 390 OF NERVOUS IRRITABILITY removed, and the poles of a galvanic apparatus applied to the sur- face of the muscle (a, h), a contraction takes place every time the circuit is completed and a discharge passed through the tissues of the limb. The leg of the frog, prepared in this way, may be em- ployed for a long time for the purpose of exhibiting the effect of various kinds of stimulus upon the muscles. All the mechanical and chemical irritants which we have mentioned, pricking, pinching, cauterization, galvanism, &o., act with more or less energy and promptitude, though the most ef&cient 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 habituail excitant 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 pbisonous substances have the power of destroying the irritability of the muscles by a direct action upon their tissue. Sulphocyanide of potassium, for example, introduced into the cir- culation in suf&oient 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 dis- tributed to a glandular organ produces increased secretion ; one dis- tributed 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 AND ITS MODE OF ACTION. 391 Fig. 133. M glandular organs, owing to the stoppage of the circulation, disappears also very rapidly, or at least cannot readily be demonstrated. The 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. (Pig. 133.) If the two poles, of a galvanic appa- ratus be now placed in contact with different points (ab) 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. 132. In that experiment the galvanic discharge passes through 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 nerve. 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 irri- tability of the latter, and may be used as a convenient measure of its intensity. 27ie irritability of the nerve continues after death. The knowledo-e 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. Frog's Leg, with sciatic uei-ve (N) at* tached. — a b. Poles of galvanic battery, ap- plied to nerve. 392 ' OF NERVOUS lEBITABILITY also, as that given above with regard to the muscks, nervous irri- tability lasts much longer after death in the cold-blooded than in 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, aU 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 attache^, 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 irri- tability 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 con- tinued for a longer period. Nervous irritability, like that of the muscles, is exhausted hy 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 contrac- tion will again ensue on the application of a stimulus to the nerve. Exhausted a second time, and a second time allowed to repose, it will again recover itself; and this may even be repeated several times in succession. At each repetition, however, the recovery of AND ITS MODE OF ACTION. 393 nervous irritability is less complete, until it finally disappears alto- gether, and can no longer be recalled. Various accidental circumstances tend to diminish or destroy nervous irritability. The action of the woorara poison, for example, destroys at once the irritability of the nerves ; so that in animals killed by this substance, no muscular contraction takes place on irritating the nervous trunk. Severe and sudden mechanical inju- ries 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 sjeparated 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 moment^, however, if kept under favorable conditions, it recovers from the shock, and regains its natural irritability. The action of the galvanic current upon the nerves, 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 towards its distribution, as from a to 6 in Fig. 133, it is called the direct current. If it be made to pass in the contrary direction, as from h to o, 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 3M OF NERVOUS IBEITABILITY nervous connection hanging between. (Pig. 134.) If the positive pole, a, of the battery be now placed in the vessel which holds leg No. 1, and the negative pole, b, 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, b, be reversed, the effects of the current will be changed in a corre- sponding 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 th& 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 AND ITS MODE OF ACTION. 395 usually adopted in the galvanic machines used in medical practice, for the treatment of certain paralytic affections. In these machines, the electric circuit is alternately formed and broken with 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 irritability 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 instafice 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 woorara, 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 throjigh the muscles, contrac- tion at once takes place. The muscular irritability has survived that of the nerves, and must therefore be regarded as essentially 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. — The special endowment by which a nerve acts and manifests its vitality is a peculiar one, inherent in the anatomical structure and constitution of the nervous tissue. It is 396 OF NERVOUS IRRITABILITY manifested, in the foregoing experiments, by its effect upon the con- tractile muscles. But we shall hereafter see that this is, in reality, only one of its results, and that it shows itself, during life, by a variety of other influences. Thus it produces, in one case, sensa- tion; in another, muscular contraction; in another, increased or modified glaiidular activity ; in another, alterations in the pheno- mena of the circulation. The force, however, which is exerted by a nerve in a state of activity, and which brings about these changes, is not directly appreciable in any way by the senses, and can be judged of only by its secondary effects. "We understand enough of its mode of operation, to know that it is not identical with the forces of chemical affinity, of mechanical action, or of electricity. And yet, by acting upon the organs to which the nerves are distributed, it will finally produce phenomena of all these different kinds. By the intervention of the muscles, it results in mechanical action ; aind by its infiilfence upon the glands and bloodvessels, it causes chemical alterations in the animal fluids of the most import- ant character. It will even produce well-marked electrical phenomena, which in some cases are so decided, as to have long attracted the attention of physiologists. It has been fully demonstrated 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 con- ductors for electricity, and is stopped by those which are non-con- ductors of the same.' All the ordinary phenomena produced bj the electric current, viz : the heating and melting of a fine con- ducting 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 generated by these animals. There is, accordingly, no room for doubt as to its nature. The electrical phenomena, in these cases, are produced by certain organs which are called into activity by the nervous influence. The electrical organs of the gymnotus and torpedo occupy a con- siderable 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 suffer if the nerves distributed to them AND ITS MODE OP ACTION, 397 were subjected to a similar violence. The electricity produced by these animals, accordingly, is not supplied by the nerves, but by a special generating organ, the action of which is regulated by nerv- ous influence. Moreover, the experiments of Longet and Matteucci* have shown that no electrical current is to be detected in a living nerve, even when in a state of activity. The electrical phenomenon, when it exists, is only a secondary effect, and is not the active force residing in the nervous tissue. This force is special in its nature, and is regulated by laws peculiar to itself. ' Longet, Traits de Physiologie. Paris, 1850, vol. ii. p. 130. S98 THE SPINAL COBD. CHAPTEE III. THE SPINAL COED. We have already seen that the spinal cord is a long ganglion, 'covered with longitudinal bundles of nervous filaipents, 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. Sensation. — The power of sensation, or sensibility, is the power by which we are enabled to receive impressions from external objects. These impressions are usually of such a nature that we can derive from them some information in regard to the qualities of external objects and the effect which they may produce upon our own systems. Thus, by bringing a foreign body into contact with the skin, we feel that it is hard or soft, rough or smooth, cold or warm. We can distinguish the separate impressions produced by several bodies of a similar character, and we can perceive whe- ther either one of them, while in contact with the skin, be at rest or in motion. This power, whrich is generally distributed over the external integument, is dependent on the nervous filaments rami- fying in its tissue. For if the nerves distributed to any part of the body be divided, the power of sensation in the corresponding region is immediately lost. SENSATioiir. 899 The sensibility, thus distributed over the integument, varies in, its acuteness in different parts of the body. Thus, the extremities of the fingers are more sensitive to external impressions than the general surface of the limbs and trunk. The surfaces of the fingers which lie in contact with each other are more sensitive than their dorsal or palmar surfaces. The point of the tongue, the lips, and the orifices of most of the mucous passages are endowed with a sensibility which is more acute than that of the general integument. If the impression to which these parts are subjected be harsh or violent in its character, or of such a nature as to injure the texture of the integument or its nerves, it then produces a sensation oipain. It-is essential to notice, however, that the sensation of pain is not a mere exaggeration of ordinary sensitive impressions, but is one of quite a different character, which is superadded to the others, or takes their place altogether. Just in proportion "as the contact of a foreign body becomes painful, our ordinary perceptions of its phy- sical properties, are blunted, and the sense of suffering predominates over ordinary sensibility. Thus if the integument be gently touched with the blade of a knife we easily feel that it is hard, cold, and smooth ; but if an incision be made with it in the skin, we lose all distinct perception of these qualities, and feel only the suffering produced by. the incisi9n. We perceive, also, the difference in temperature between cold and warm substances brought in contact with the skin, so long as this difference is moderate in degree ; but if a foreign body be excessively cold or excessively hot, we can no longer appreciate its temperature by the touch, but only its injurious and destructive effect. Thus the sensation caused by touching frozen carbonic acid is the same with that produced by a red-hot metal. Both substances blister the surface, but their actual temperatures cannot be distinguished. It is, therefore, a very important fact in this connection, that the sensibility to pain is distinct from the power of ordinary sensation. This distinction was first fully established by M. Beau, of Paris, who has shown conclusively that the sensibility to pain may be diminished or suspended, while ordinary sensation remains. This is often seen in patients who are partially under the influence of ether > or chlo- roform. The etherizfition may be carried to such an extent that the patient may be quite insensible to the pain of a surgical opera- tion, and yet remain perfectly conscious, and even capable of feeling the incisions, ligatures, &c., though he does not suffer from them. It not unfrequently happens, also, when opium has been adminis- 400 THE SPINAL COED. tered for the relief of neuralgia, that the pain is completely abolished by the influence of the drug, while the patient retains completely his Consciousness and his ordinary sensibility. In all cases, however, if the influence of the narcotic be pushed to its extreme, both kinds of sensibility are suspended together, and the patient becomes entirely unconscious of extesrnal impressions. Motion. — Wherever muscular tissue exists, in any part of the body, we find the power of motion, owing to the contractility of the muscular fibres. But this power of motion, as we have already seen, is dependent on the nervous system. The excitement which causes the contraction of the muscles is transmitted to them by the nervous filaments ; and if the nerve supplying a muscle or,a limb be divided or seriously injured, these. parts are at once paralyzed and become incapable of voluntary movement. A nerve which, when irritated, acts directly upon a muscle, producing contraction, is said to be excitable; and its excitability, acting through the mus- cle, produces motion in the part to which it is distributed. The excitability of various nerves, however, oftein acts "during life upon other organs, beside the muscles ; and the ultimate effect varies, of course, with the properties of the organ which is acted upon. Thus, the nervous excitement transmitted to a muscle pro- duces contraction, while that transmitted to a gland produces an increased secretion, and that conveyed to a vascular surface causes congestion. In all such instances, the effect is produced by an influence transmitted by the nerve directly to the organ which is called into activity. But in all the external parts of the body muscular contraction is the most marked and palpable effect produced by the direct influence of nervous excitement. "We find, therefore, that so far as we have yet examined it, the nervous action shows itself princi- pally in two distinct and definite forms ; first, as sensibility, or the power of sensation, and second, as excitability or the power of pro- ducing motion. 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 apo- pleixy, concussion or compression of the brain or spinal cord, or a DISTINCT SEAT OF SENSATION AND MOTION. 401 wound of any kind involving the nerves or nervous centres, insen- sibility and loss of motion usually appear simultaneously. It is difftcult, therefore, 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 remov^ by examining sepa- rately different parts of the nervous system. In the instances mentioned above, the injury whiqh is inflicted is comparatively an extensive one, and involves at the s^me time many adjacent parts. But 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 Lpnget, 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 26 402 THE SPINAL COKD. nerve, in the living animal, convulsive movements are produced ini 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 therfe 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 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 (Fig. 135) at a b, and an irritation applied Diagram of Spisai Coup and Nekveb. The posterior root is sees divided at a, b, the anterior at c, d. * to the separated extremity (a), no effect will be produced; but if the irritation be applied to the attached extremity (6), 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 SENSIBILITY AND EXCITABILITr IN SPINAL COBD. 403 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 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 theit charac- ter and function, viz., the " sensitive" filaments, or those which convey sensation, and the " motor" filaments, or those which excite movement. These filaments are never confounded with each other in their action, nor can they perform each other's functions. The sensitive 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 tirne over the functions of movement and sensation. Distinct Seat of Sensibility and Excitability in the Spinal Cobd. — Various experimenters have demonstrated the fact that different parts .of the spinal cord, like the two roots of the spinal nerves, are separately endowed with sensibility and excita- bility. 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 produpes immediately convulsions in the limbs below ; but there is no indication of pain. On the other hand, signs of 404 THE. SPINAL CORP. acute pain become manifest whenever the irritation is applied to the posterior columns ; 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 transverse section, the application of any irritant to the anterior surface of the separated portion produces at once con- vulsions below ; while if applied to the posterior columns behind the point of division, it has no sensible effect whatever. The an- terior and posterior columns of the cord are accordingly, so far, analogous in their properties to the anterior and posterior roots of the spinal nerves, arid are plainly composed, to a greater or less ex- tent, 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 pos- terior root, to the spinal cord ; then upward, along the longitudinal fibres of the cord to the brain, where it produces a sensation corres- ponding in character with the original irritation. A motor im- pulse, 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, following the motor filaments of the nerve through its trunk and branches, produces at last a muscular contraction at the point of its final distribution. Crossed Action of the Spinal Cord. — As the anteripr columns of the cord pass upward to join the medulla oblongata, a decussa- tion 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 m^ be readily shown (as in Fig. 130) by gently separating the anterior columns from each other, at the lower extremity 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 communicate by any further decussation. ■ Traits de Physiolpgie, toI. ii. part 2, p. 8. CBOQSED ACTION OF THE SPINAL COBD. 405 If the anterior columns of the spinal cord, therefore, be wounded at any point in the cervical, dorsal, or lumbar region, a paralysis 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 tlie lefib side of the brain will be fol- lowed 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-S^quard,' 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 opgosite 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 sensation 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. ' Experimental Researches applied to Physiology and Pathology. New York, 18r)3. " Memoirs eur la Physiologie de la Moelle 4pini4re ; Gazette M^dicale de Paris, 1855. 406 THE SPINAL CORD.' There are certain important facts whicli sfill remain to be noticed regarding the mode of action of the spinal cord and its nerves. They are as follows : — 1. An irritation applied 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 hav6 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 integumeitt to which its filaments are distributed. Thus, if the ulnar nerve he accidentally struck at the point wheje it lies behind the inner con- dyle of the humerus, a sensation of tingling and numbness is pro- duced in the last two fingers of the corresponding hand. It is common to hear patients who have suffered amputation complain of painful sensations in the amputated limb for weeks or months, and sometimes even for years after the operation. They assert that th6y 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 irrita- tion, conveyed to the brain by the sensitive fibres, will produce 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 for the cure of tic douloureux. If the cause of the difficulty be seated upon the trunk of the nerve, between its point of emergence froifi 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 disappears from within out- INDEPENliKNCE OP NEBVOUS FILAMENTS. 407 ward, that of the sensitive filaments from without inward. Immedi- ately after tlie separation of the frog's leg from the body, irritation of the nerve at any point produces muscular contraction in the 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 also, in turn, it afterward disappears j receding in this man- ner, farther and farther toward the terminal distribution of the nerve, where it finally disappears altogether. ,0n 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. JSach nervous filament acts independently of the rest throughout its entire length, 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 pf 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 imitation 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 frofii 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, ■wfhich 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 408 THE SPINAL COBD. 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 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 he looked for in some special endowment of the nervous centres from which they originate. Eeflex Action of the Spinal Cobd. — The spinal cord, as we have thus far examined it, may be regarded 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 production of conscious sensation and voluntary motion. Beside its nervous 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 KEFLEX ACTION OF THE SPINAL COKD. 409 • 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 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 integvment 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, irritation of the skin below is no longer followed by any muscular contrac- tion. If either the anterior or posterior roots of the nerve be divided, the same want of action results ; and finally, if, thfe 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 ^^^^^^Jg^36^^^^^^ irritated or mutilated to* any extent, ^^^^^U^^^^^^^^f without exciting the least muscular. ^^^^P^^^^j^^^^^^l contraction. It is evident, therefore, ^^^^ ■ B^^^^^f that the spinal cord acts, in this case, ^^^| 1^ ^S^^^l as a nervous centre, through which ^^^H ■ iH^^^H the irritation applied to the skin is ^^^H I, ^^K^^l communicated to the muscles. The ^^^H ■ m^^^^^B irritation first passes upward, as shown ^^^H ■ ^^^^^^H in the accompanying diagram (Fig. ^^^^^^^^^^^^^^^M 136), along the sensitive fibres of the ^^^^^^^^^^^^^^^B ^ . , Diagram of SPlKAL Cord im Vik- posterior root (a) to the gray matter tic*!, skchoj., showing reflex action. of the cord, and is then reflected back, -"• !'''»»•"<" ""' <•' •p"'»i -""•■ »• Anterior root of spinal nerve. along the motor fibres of the anterior root (6), until it finally reaches the musclQp, and produces a contrac- 410 THE SPINAL COBD. • tion. This action is known, accordingly, as the reflex action of the spinal cord. It will be remembered that this reflex action of the cord is not accompanied by 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 animalsv 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 by strychaine 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 movementfOf 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'Sondition, 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- ninei, 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 frog be poisoned with a moderate dose of strychnine, the body and limbs will remain quiesqpnt so long as there is no external source EKFLEX ACTION OF THE SPINAL CORD. 4.11 of excitement ; but the limbs*are at once thrown into convulsions by the slightest irritation applied to the skin, as, for example, the contact of a hair or a feather, or even ■ the jarring of the table on which the animal is placed. That the convulsions iu cases of poisoning by strychnine are always of a reflex character, and never spontaneous, is shown by the following fact first noticed by Ber- nard,' viz., that if a frog be poisoned after division of the posterior roots of aU the spinal nerves, while the anterior roots are left un- touched, death takes place as usual, but is not preceded by any con- vulsions. In this instance the convulsions are absent simply because, owing to the division of the posterior roots, external irri- tations cannot be communicated to the cord. The reflex action, above described, may be seen very distinctly in the human smbject, 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 ana- tomical lesion. , Under these conditions, the patient is incapable of making any muscular exertion in the paralyzed parts, and is uncon- scious 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 twitch- ing of the toes will often take place, and even retractile movements of the leg and thigh, altogether without the patient's knowledge^ 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 dis- ease of the spinal cord, where the paralysis and insensibility 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 dbmonstrated,' 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- ■ ' Iie(;ons surles effets des Substances toxlques et ipedicamentenses, Paris, 1857, p. 357. > Ibid., Cbaps. 23 and 24. 412 THE SPINAL COBD. tered; and 2d, thabthe motor and s^sitive nervous filaments may be paralyzed independently of each other. The above facts are shown by the three following experiments : — 1. In a.living frog (Fig. 137), the sciatic nerve {N) is exposed in the back part of the thigh, after which a ligature is passed under- neath it and drawn tight around the bone and the remaining soft parts. In this way the circulation is entirely cut off from the limb ((£), which remains in opnnection with the trunk only by means of the sciatic nerve. A solution of sulphocyanide of potassium is ftien introduced beneath the skin ^'^- ^^^- of the back, at I, 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, h, c ; while it is pre- vented from entering the limb d, by the ligature' which has been placed about the thigh. Sulphocyanide of potassium produces 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, b, and c, produces no contraction in them, while the same stimulus, applied to d, is followed by a strong and healthy reaction. But at the moment when the irritation is applied to the poisoned limbs a, b, 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 inte- gument of «, b, and c, and transmitted, by sensitive and motor fila- ments, through the cord to d. While the muscles of the poitoned KEFLEX ACTION OF THE SPINAL CORl). 413 limbs, therefore, have been directly paralyzed, thd nerves of the same parts have retained their irritability. ' 2. If a frog be poisoned with woorara by simply placing the 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 s^sitive 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 I, when the limb d is irritated its own muscles react, while no movement takes place in a, b, or c; but if the irritation be applied to a, b, or c, reflex movements are imme- diately produced in d. In the poisoned limbs, therefore, 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 mator filaments and the mvscles have retained their irritability. "We now come 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 414 THE SPINAL COED. even fairly understand the cause of the involuntary act. If the body by any accident suddenly and unexpectedly loses its balance, the limbs are thrown into a position calculated to protect the ex- 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 %lso 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 alloived to escape. Whea any external irritation is applied to the anus, or whenever the feces present themselves internlally, 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 urgent with the increased distension of the . rectum, becomes irre- sistible;' 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 are 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 KEFLEX ACTION OF THJ! SPINAL CORD. 415 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 distension of the rectum is announced by the usual sensation, but the reflex impulse to evacuation is so urgent that it cannot be controlled by 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 paralyzed ; 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 elas- ticity of the protecting fibres, by simple force of accumulation, and afterward dribbles away as fast as it is excreted by the kidneys. Paralysis of the bladder, therefore, first causes a permanent 'disten- sion 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 it^ 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 almost invariably follow 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 de- pendent 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 416 THE SPINAL CORD. 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 of the urine contained in the bladder, converts it into an irri- tating and ammoniacal liquid, which reacts upon the mucous mem- brane 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 BEAIN. 417. CHAPTER IV. THE BRAIN. By the brain, or enc^hahn, 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 al], 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 surface 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 the base of the brain which contain prolongations of the posterior columns of the cord. 27 41^ TSE BKAIN. The most central portions of the nervous system, therefore, ai^ 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 dififerent ganglia of which the brain is composed. Olfactoey GtANGLIA. — These ganglia, which in some of the lower animals are very large, corresponding in size with, the ex- tent of the olfactory 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 beneath 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 olfaptory ganglia with their commissures are sometimes spoken of as the " olfactory nerves." They are not nerves, however, but ganglia, since they are mostly composed 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 olfactory membrane. It has been found difficult to determine the function of these ganglia by direct experiment on the lower animals. They may he 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 speciail sense of smell, and are not connected, in any appreciable degree, with ordinary sensibility, nor with the production of voluntary move- ments. Optic Thalami. — These bodies are not, as their name would imply, the ganglia of vision. Longet has found that the power of sight and the sensibility of the pupil both remain, in birds, after CORPORA STRIATA. — HEMISPHERES. 419 the optic thalami Iiave been thorouglily disorganized ; and that arti- ficial 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 voluntary 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 in- stances 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 compa- rative 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 substance 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 connec- tion is at present altogether unknown. Hemispheres. — The hemispheres, or the cerebral ganglia, con- stitute in the human subject about nine-tenths of the whole mass of the brain. Throughout their whole extent they are entirely destitute, as we have already mentioned, of both sensibility arid ex- citability. Both the white and gray substance may be wounded, burned, lacerated, crushed, or galvanized in the living animal, with- out exciting any convulsive movement or any apparent sensation. In the human subject a similar insensibility has been observed when the substance of the hemispheres has been exposed by acci- dental violence, or in the operation of trephining. Very 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 Prof. William Detmold, of New York,' in « > Am, Journ. of Med. Sci., January, 1850. 420 THE BRAIN, 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, however, headache and drowsiness came on ; and the latter symptom, becom- ing 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 sub- stance, making a wound one inch in length and half an inch in depth, when the abscess was reached and over two ounces of pus discharged. The patient immediately aroused from his comatose condition, so that he was able to speak ; and in a few days reco- vered, to a very considerable extent, his cheerfulness, intelligence, and appetite. Subsequently, however, the collection of pus re- turned, 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 pointeS iron bar, three feet and a half in length, and one inch and a quarter in diameter, was driven through the patient's head by the ipremature 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 i^he 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 rec&Vered 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 HEMISPHEBKS. 421 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 similar 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- Fig. 138. Pigeon, after Removal of the Heuiepheres. mage is smooth and glossy, but is uniformly expanded, by a kind of erection of the feathers, so that thfe body appears somewhat puffed out, and larger than natural. Occasionally the bird opens his eyes with a vacant stare, stretches his neek, 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 422 THE BRAirr. hearing, or of ordinary sensibility. All these functions remain, as •well as that of voluntary motion. If a pistol be discharged behind 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 ; hut he immediately becomes quiet again, and pays no farther attention to it. Sight is also retained, since the bird will sometimes fix its 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 sensorium. 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 sensation 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, because apparently it lacks its usual mental stimulus and direction. The powers which 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 redognized as the ordinary consequence of lesions of the brain. In cases q{ 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. HEMISPHEEES. 423 His mental derangfement 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 difi'erent 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- 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 unvaTying, 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 propor- tion 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 424 THE BEAIN. in them. For wliile mau is superior in general intelligence to all the lower animals, he, is inferior to many of them in the acutenesa of the 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 bf 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 sourqes, 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 in the power of perceiving' external objects and events, And of re- cognizing the connection between them, as in that of drawing con- clusions from one fact to another, and of adapting to new combina- tions 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 HEMISPHERES. 425 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 hemispheres. 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, ^he 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 and impressibility of organization which take the place, to a certain 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 some years ago. in various parts of the United States, under the name of the "Aztec children." They were a boy and a girl, aged respectively about seven and five years. The boy was 2 feet 9f inches high, and weighed a little over 20 pounds. The girl was 2 feet 5J inches high, and weighed 17 pounds. Their bodies were tolerably well proportioned, but the cranial cavities, as shown by the accompany- ing portraits, were extremely small. The antero-posterior diameter of the boy's head was only 4J inches, the transverse diameter less than 4 inches. The antero- 426 THE BRAIN. posterior diameter of the girl's head was 4J inches, the transverse diameter only 3f inches. The habits of these children, so far as regards feeding and taking care of themselves, were those of chil- 'Pi Fig. 139. Aztec Chii.dkeh. — Taken from life, at five and seven years of age. 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 rgpiarkably 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 meaningcould 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 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 different facul- ties 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 HEMISPHERES. 427 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 endowment? of the in- dividual 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 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 428 THE BRAIN. 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, like those 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 suf&cient 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 mistakes. It is extremely improbable, therefore, that either Gall or Spurzheim could, in a single lifetime, have accomplished this comparison 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 tie 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, where- ever it exists. Gall and Spurzheim located all the mental faculties in those parts of the brain which are accessible to external exploration. An examina- tion of different sections of the . brain will show, however, that the greater por- tion of the gray substance is so placed, that its quantity cannot be estimated by an external examination through the skull. The only portions which are exposed to such an examination are the upper and lateral portions of the con- vexities of the hemispheres, together with the posterior edge and part of the under surface of the cerebellum. (Fig. 140.) A very extensive Fig. 140. Diagram of the Br AiH ik 6ITD, showing those portions which are ex- posed to examination. HKMISPHEEES, 429 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 ante- rior and middle lobes (i, 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 alaove the medulla oblongata (5) ; and the two opposite convoluted surfaces in the fissure of Sylvius (e, 1), where the anteiior 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 examination. The whole of the convoluted surface of the brain must, however, be regarded as of equal im- portance in the distribution of the mental qualities; and yet it is evident that not liiore than one-third or ©ne-quarter of this surface is so placed that it can be examined by external manipulation. It must further- more be recollected that the gray matter of the cerebrum and cerebellum is everywhere convoluted, and that the convolutions pene- trate to various depths in the substance of the brain. Ev-en 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 convolu- tions penetrate, as upon the prominence of their edges. While phrenology, therefore, is partially founded upon acknow- ledged physiological facts, there are yet essential deficiencies in its scientific basis, as well as insurmountable difBculties in the way of its practical application. Transverse section of B r a I N, showing depth of great longi- tudinal flssare, at a. Cerebellum. — The cerebellum is the second ganglion of the encephalon, in respect to 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 cerebellum therefore is smaller, as a whole, than the 430 THE BKAIN. cerebrum, it contains, in proportion to its size, a mucli larger quan- tity 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 haitily 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 exiperimental 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 Elourens, 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- 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. Bveti 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 efliciency of the muscular apparatus. The opinion Which locates the above harmonizing or associating power in the cerebellum was first suggested by the effects observed CEREBELLUM. 431 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, bilt can neither walk nor fly. Fig. 142. PlOEOK, AFTER REBOTAI. OF THE CEREBELLUM. 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 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 432 THE BKAIN. moves sluggishly and unwillingly, but otherwise acts 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 eaisily terrified, and endeavors, frequently with violent struggles, to escape the notice of those who are watching him; but his movements are sprawling and unnatural, and are evidently no longer under the effectual control of -the will. (Fig. 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 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. "We have often performed the above operation, and always with the same effect. Indeed there are few experiments that have been tried upon the nervous system, which give results so uniform and so constant as this. Taken by themselves, these results would invariably sustain the theory of Flourens, which, indeed, is founded entirely upon them. But we have met with another very important fact, in this respect, which has hitherto escaped notice. That is, that birds, which have lost their power of muscular co-ordination from injury of the cere- bellum, may recover this power in process of time, notwithstanding that a large portion of the cerebellum has been permanently removed. Usually such an operation upon the cerebellum, as we have men- tioned above, is fatal within twenty-four hours, probably on account of the close proximity of the medulla oblongata. But in' some instances, the pigeons upon which we have operated have survived, and in these cases the co-ordinating power became re-established. In the first of these instances, about two-thirds of the cerebellum was taken away, by an opening in the posterior part of the cranium. Immediately after the operation, the animal showed all the usual effects of the operation, being incapable of flying, walking, or even standing still, but reeled and sprawled about in a perfectly helpless manner. In the course of five or six days, however, he had regained a very considerable control over the voluntary movements, and at the end of sixteen days his power of muscular co-ordination was so nearly perfect, that its deficiency, if any existed, was impercep- tible. He was then killed ; and on examination, it was found that his cerebellum remained in nearly the same condition as immediately after the operation ; about two-thirds of its substance being deficient. CKBEBELLUM. 433 and no attempt Laving been made at regeneration of the lost parts. Tlie accompanying figures, 143 ta 146, show the appearances, in this case, as compared with the brain of a healthy pigeon. We have also met with three other cases, similar to the above, in which about one-half of the cerebellum was removed by operation. The loss of co-ordinating power, immediately after the operation, though less complete than in the instance above mentioned, was perfectly well marked in character ; and in little more than a fort- Pig. 143. Fig. 144 Braik of HEiLTHT F I OE OS— Profile Bkaim op Operated Pioeos— Tiew. — 1- HtimUphere. 2. Optic tubercle, 3. Profile view — showing the matilatioa Cerebeilam. 4. Optic uerTe. 5. Medalla ob- of cerebellam. longata. Fig. 145. Fig. 146. Braim of Healthi Pioeos— Foste- Braik of Operated PioEoir— rior view. Fo9teri# view — showing the matila- tioa of cerebellum. night the animals had nearly or quite recovered the natural control of their motions. These instances show, accordingly, that a large portion of the cerebellum may be wanting without a porresponding deficiency of the co-ordinating power. If the theory of Flourens be correct, therefore, these cases can only be explained by supposing that those parts of the cerebellum which remain gradually become en- abled to supply the place of those which are removed. It is more probable, however, that the loss of co-ordinating power, which, is immediately produced by taking away a considerable portion of this nervous centre, is to be regarded rather as the effect of the sudden injury to the cerebellum as a whole, than as due to the mere removal of a portion of its mass. 28 434 THE BEAIN, Morbid alterations of the cerebellum, furthermore, particularly of a chronic nature, such as slow inflammations, abscesses, tumors, &c., have often been observed in the human subject, without giving rise to any marked disturbance of the voluntar/ movements. On the other hand, many facts derived from comparative anatomy seem to favor the opinion of Flourens. If we compare different classes of animals with each other, as fish with reptiles, or birds with quadrupeds, in which the developnjent and activity of the entire nervous system vary extremely, the results of the comparison ■ will be often contradictory. But if the comparison ,be made be- tween different species in which the general structure and plan of organization are similar, we often find the development of the cere- bellum to correspond very closely with the perfection and variety of the voluntary movements. The frog, for example, is an aquatic reptile, provided with anterior and posterior extremities; but its, movements, though rapid and vigorous, are exceedingly simple in character, consisting of little else than flexion and extension of the posterior limbs. The cerebellum in this animal is exceedingly small, as 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 movements of swimming, diving, progression, &c., are accomplished by the consentarfBbus action of anterior and posterior extremities, and where the motions of the head and neck are also much more varied than in the frog. In this instance the cerebellum is very much more bighly developed than in the former. lit the alligator, 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. The complete function of the cerebellum, accordingly, as a nerv- ous centre, cannot be regarded as positively ascertained ; but so far as we may rely on the results of direct experiment, this organ has evidently such an intimate and peculiar connection with the volun- tary movements, that a sudden and extensive injury inflicted upon its substance is always followed by an immediate, though tempo- rary, disturbance of the co-ordinating power. TuBEBCDLA 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 TUBEECULA QUADRIGEMINA. 435 the sense of sight ; on -which account they are also known by thd name of the " optic ganglia." Their development corresponds very closely with that of the external organs of vision. Thus, they ard large in fish, reptiles, and birds, in which the eyeball is for th«f most part very large in proportion to the entire head; and are small in quadrupeds and in man, where the eyeball is, comparatively, speaking, of insignificant siae. Direct experiment also shows tha 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, produces complete blindness ; and destruc- tion of the tubercles themselves 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 un- touched, 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 iminjured 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 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 mucTi 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 436 THE BBAIK. 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 %ith the brain, mechanical irritation of the optic nerve, as we have shown above, causes contraction of the pupil ; but if the nerve be divided, and the extremity which remains, in connection with the ieyeball be subjected to irritation, no effect upon the pupil is pro- duced. 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 at 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. 147.) The two Fig. 147. Fig. 148. Inferior Surface of Bbatit OF Cod.— 1. Rlglit optic nerve. 2. Left optic nerve. 3. Right optic tubercle. 4. Left optic tabercle. 5, 6. Hemispheres. 7. Medulla oblongata. Inferior Surface of Brain o? Vowh.—l. Right optic nerve. 3 Left optic nerve. 3. Right o|>tic tubercle. 4. Left optic tubercle, ff, 6. Hemitipheres. 7. Me- dulla oblongata. nervous cords are here totally distinct from each other throughout their entire length ; and are only connected, at the point of cross- TUBEBCULA QUADBIGEMINA. 437 ing, by intervening areolar tisSue. ' Impression* 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. 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 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. 148.) 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 side on which the injury was inflicted. Fie. 14fl. ConKBE OP Optic Nerves iH Man.— 1, 2. Eight and left eyeballs. 3. Decossltion or optic nerves. 4, 4. Tnbercala qnadrigemina. In the human %ilbject, on the other hand, where the visual axes are parallel, and where both eyes are simultaneously directed toward 438 THE BRAIN. the same object, the optic nerves dedtssate with each other hi 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. 149.) This decussation, which is somewhat complicated, takes place in the following manner. From each optic tubercle three different bundles or " tracts" of nervous fibres are given ofl". One set passes 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 forwal-d 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 direction, so th^t the parallelism of their axes is lost, and the image no longer falls upon corresponding parts of the two retinae. Tuber Ankulake. — 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 sensa- tion and voluntary niotion. "We have already seen that these func- tions are not destroyed by taking away the cerebrum, and that they also remain after removal of the cerebellum. According to the ex- periments of Longet, even after complete removal of the olfactory ganglia, the cereibrum, 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 en- deavor by voluntary 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 MEDULLA OBLONGATA. 48® exist. The only movements whieh then follo-w 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 voluntary character. The animal, under these circum- stances, is to all appearance 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, con- veyed 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 taike 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 fe^t 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 arid 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 the cerebral hemispheres ; the latter, in the gan- glion of the tuber annulare. Medulla Oblongata.— The last remaining ganglion of the en- cephalon is that of the medulla oblongata. This ganglion, it will be remembered, is imbedded in the substance of the restiform body, MO THE BBAIN. occupying the 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 pre- servation 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 imme- diate and necessary destruction of life ; but so soon as the medulla oblongata is broken up, and its ganglion destroyed, respiration ceases instantaneously, and the circulation also soon comes to an end. Eemoval of the medulla oblongata produces, therefore, as its immediate and direct result, a stoppage of respiration : and death takes place principally 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- portionally 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 J;he 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 thg greater part of the time, in a perfectly quiet and regular manner, without our volition and even without our consciousness. They continue uninterruptedly through the deepest' slumber, and even in a con- dition 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 MEDULLA OBLONGATA. 441 reflected 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. Eespiration, 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 designated 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 satilfied, the muscles relax, anS the chest collapses. In a few seconds the previous condition recurs and tbfcame 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 442 THE BRAIN. and third cervical nerves, botli 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 ineffectual 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 strug- gle, 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 pro- cess 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 hemi- spheres, corpora striata, or optic thalami, is the seat of the hemor- rhage, the patient generflly lives at least twelve hdurs; but if the hemorrhagqjpikes place into the medulla itself, or at the base of th% brain in its immediate neighborhood, so as to compress its sub- stance, 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- pa^f'ed 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- MEDULLA OBLOifGATA. 443 # 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 coimection 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 different 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|bervous impression is of a special character, and its influence is finally exert^ 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 tbe 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 444 THE BRAIN. 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 sensation of hunger > and thirst. All actions of this nature are termed instinctive. 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 Ifeve 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 Hf 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 theapimal 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 circufti- stances. The third kind of reflex action requires the co-operation of the hemisphereg. 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 MEDULLA OBLONGATA. 445 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 dfiapted, 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. 44:6 THE CRANIAL NEBVES. CHAPTER V. • THE CRANIAL NERVES. In examining the cranial nerves, we shall find that although they dt 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 cra- nial 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, Olfactoey 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 finally that their destruction, together with that of the olfactory ganglia, does not occasion any paralysis nor loss of ordinary sensibility. THE CRANIAL NERVES. 447 Optic Nerves. — We have already given some account of these nerves and their decussations, in connection with the history of the tubercula quadrigemina.r They consist of rounded bundles of white fibres, running between the tubercles and the retina. As the reti- nae themselves are membranous expansions consisting mostly of vesicular or cellular nervous matter, the optic nerves, or "tracts," must be regarded as commissures connecting the retinae with the tubercles. We have also seen that they serve, by some of their fibres, to connect the two retinae 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 Longet, the optic nerves are absolutely insensible to pain through- out 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 lumi- nous 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 ^jrections ; and a sudden stroke upon the eyeball will often give rise to an apparent discharge . 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 ol&ctory, 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 the 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 > Traits de Physiologie, vol. ii. p 286. , 448 THE CRANIAL NERVES. 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 or sensitive, properly speaking; and that they are distinct in their nature from the ordinary spinal nerves. The remainder of the cranial nerves, however, present the ordinary qualities belonging to the spinal nerves. 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, cor- responding 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 : — 1st FAIB. 2d FAIB. 3d FAIB. J Cbaniai. Xebtes. Nerves of Special Sense. 1. Olfaetory. 2. Optio. 3. Anditoiy. Motor nerves. Sensitive Nerves. ' Motor ocnii com. Patheticus Motor oc. eKtemns Small root of 5th pair Facial Distributed to Large root of 5th pair. Face. J Hypoglossal Spinal accessory Glossq-pharyngeal. Fneumogastric. Tongue. Neck, &c. The above arrangement of the cranial nerves is not absolutely perfect in all its details. Thus, while the hypoglossal 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 he found extremely useful as an assistant in the study of their func- tioi^s. 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 THE CRANIAL NERVES. 449 bundle only at a certain distance from their point of origin. The 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 trunk 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 Prof. 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. 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 integu- ment or to mucous membranes; secondly, their division causes muscular paralysis; and thirdly, mechanical irritation applied at their origin produces muscular contraction in the parts to which they are distributed, but does not give rise to a painful sensa- tion. In several instances, nevertheless, the motor nerves, though insensible at their origin, show a certain degree of sensibility when irritated after their exit from the skull, owing to fibres of com-, munication which they receive from the purely sensitive nerves. In this respect they resemble the spinal nerves, the motor arid sensitive filaments of which are at first distinct in the anterior and posterior roots, but afterward mingle with each other, jon leaving the cavity of the spinal canal. The three sensitive nerves originating from the brain are the ' Nervous System of Rana pipiens ; published by the Smithsonian Institution. Washington, 1853. 29 450 THE CRANIAL NERVES. large root of tlie fiftb 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 ^fifth 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 gan- glia, are intermingled with others of a motor origin. The large, root of the fifth pair, which 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 spir nal accessory, becoming consequently a mixed nerve outside the crania] 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. Motor Oculi Communis.— This nerve, which is sometimes known by the more convenient name of the ocuh-motorius, originates from the inner edge of the crus cerebri, passes into the cavity of the orbit by the sphenoidal fissure, and is distributed to the levator palpebrse 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 FIFTH PAIB. 451 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. PATHETIC0S. — This nerve, which supplies the superior oblique musclfe 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 Exteenus. — 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. Fifth Pair. — This is one of the most important and remarkable in its properties of all the cranial nerves. It is the. great sensitive nerve of the face, and of the adjoining mucous membranes. Its larger 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, passing forward in com- pany with the others, 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. In front of the anterior and external border of the Casserian ganglion, the fifth nerve separates into three principal divisions, viz., the ophthalmic, the superior maxillary, and the inferior maxillary. The first of these divisions, . 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 gan- glion of the sympathetic, to the lachrymal gland, the conjunctiva, 452 THE CRANIAL NERVES. Fig. 150. and the mucous membrane of the lachrymal sac. It also sends off a small branch (nasal branch) which penetrates into the nasal pas- sages and supplies the Schneiderian mucous membrane. It then emerges upon the face by the supra-orbital foramen, and is distri- buted to the integument of the forehead and side of the head as far back as the vertex. The second 'division of this nerve, or the superior maxillary, passes out by the foramen rotundum, and runs along the longitu- dinal canal in the floor of the orbit, giving off branches during its passage to the teeth of the upper jaw and to the mucous membrane of the antrum maxillare. It finally emerges upon the middle of the face by the infra-orbital foramen, and is distributed to the integu- ment of the lower eyelid, nose, cheek, and upper lip. The third, or inferior maxillary division of the fifth pair, which is the largest of the three, leaves the cavity of the cranium by the foramen ovale. It comprises a con- siderable portion of the large root of the nerve, and all the fibres of the small root. This division is therefore a mixed nerve, containing both motor and sensitive fibres, while the two former are exclu- sively sensitive. It is distributed^ accordingly, both to muscles and to the sensitive surfaces. Soon after emerging from the foramen ovale • it sends branches to the temporal muscle, to the masseter, the bucci- nator, and to the internal and ex- ternal pterygoids; that is, to the muscles which are particularly con- cerned in the movements of the lower jaw. It also sends sensitive filaments to the integument of the temple, to that of a portion of the external ear and external audi- tory meatus. The third division of the fifth pair, then passing downward and forward, gives off a branch of considerable size, the lingual branch, which is distributed to the mucous membrane of the anterior two-thirds of the tongue, and which also sends filaments to the arches of the palate and to the mucous membrane of the cheek. DiSTKIBlTTIOS OP FjPTH NeRTB UPON THE Face. — a. Casserian j;f)iQglion. I. Ophthalmic divisioa. ' 2. Superior maxil- lary divisiOD. 3. Inferior maxillary division. FIFTH PAIR. 453 The remaining portion of the third division4 after giving a'few branches to the mylo-hyoid muscle and to the anterior belly of the digastric, then enters the inferior dental canal, sends filaments 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, and to the muscles of mastication. A few of its fibres are sent also to the superficial muscles of the face, such as the buccinator and the orbicularis oris ; 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 various branches of this nerve be irritated by a galvanic current, no con- vulsive movements whatever are produced in those superficial muscles of the face, which it supplies with filaments; but if its smaller or non-ganglionio root be irritated in the same way, con- tractions instantly follow in the muscles of mastication. The fifth pair is the most acutely sensitive nerve in the whole body. Its irritation by mechanical means always causes intense pain, and even though the animal be nearly unconscious from the influence of ether, any severe injury to its large root is almost invariably followed by cries which indicate the extreme sensibility of its fibres. If this nerve be completely divided, in the living animal, within the cranium, at the situation of the Casserian ganglion, the operation is followed by total loss of sensibility in the skin of the face and in the adjacent mucous membranes. The conjunctiva, upon the affected side, is then completely insensible, and may be touched with the point of a needle or the blade of a knife, without exciting any un- easiness, and even without the consciousness of the animal. Probes and needles may be passed into the nostril, and the lips or the cheek may be pinched, pierced or cut, without exciting the least sign of sensibility. The animal is entirely indifferent to all me- chanical injuries upon the affected side, though upon the opposite side the parts retain their natural sensibility. Owing to the paralysis of the lingual nerve, also, after this ope- ratioii, the tongue, in its anterior two-thirds, becomes insensible to ordinary irritations, and loses, beside, the power of taste. Another peculiar effect of the division of the fifth pair depends 454 THK CRANIAL NERVES. Upon the paralysis of its motor fibres, which are distributed, as we have seen, to the muscles of mastication. In many of the lower animals, consequently, the movements of mastication become ex- ceedingly enfeebled upon the affected side. In the cat, for example, an animal in which mastication is usually very thoroughly per- formed, this process becomes excessively laborious, so that the animal after this operation cannot masticate solid meat, but requires to be fed with that which has already been cut in pieces. The fifth pair, beside supplying the sensibility of the integument of the face, has a peculiar and important influence on the organs of special sense. JThis influence appears to consist in some connection between the action of the fifth pair and the processes of nutrition ; so that when the former is injured, the latter very soon become 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. Now as the nutrition of the organ is, to a certain extent, under the control of the fifth 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 fifth pair, distributed s throughout its middle and lower portions, by which it is supplied with ordinary sensibility. Since the fifth pair, accordingly, supplies general sensibility, to the nasal passages, this property wiU remain after the, special sense of smell has been destroyed. If, however, the fifth pair itself he divided, not only is general sensibility destroyed in the Schneiderian mucous membrane, but 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 mucous membrane, under these circumstances, becomes injected and swollen, and the nasal passage is obstructed by an accumulation of puriform mucus. According to Longet, the mucous memT3rane also 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 FIFTH PAIR. 455 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 secretions 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 fifth pair within the cranium, or of its ophthalmic branch, is fol- lowed by an inflammation of the corresponding eye, which usuliUy 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 fifth pair, when performed at the level of the Casserian g-anglion, 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 fifth pair on the nutrition of the eyeball does not reside in its own proper fibres, but in some filaments of the sympathetic nerve which join the fifth 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 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 positive manner. 456 THE CRANIAL NERVES. Division of the fiftli 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 accu- mulation of cerumen to take place after inflammation of this part, so as to block up the auditory canal and produce partial or com- plete deafness. It has not been ascertained, however, whether division of the fifth pair is usually followed by similar changes in this part. The lingual branch of the fifth 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 different nerves ; in its anterior two-thirds, by the lingual branch of the fifth pair ; in its posterior third, by the fibres of the glosso-pharyngeal. The facial branches of the fifth 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 sensi- tive filaments which that nerve receives from the fifth pair. FACiAL.^This nerve was known to the older anatomists as the "portio dura of the seventh pair." It 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 former, they have received the names respectively of the "portio mollis" and " portio dura" of the seyenth 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 different branches. It then sends its filaments forward in a diverging course, and is finally distributed to the muscles of the external ear, to the frontalis and superciliaris mupcles, to the orbicularis oculi, the compressors and dilators of the nares, the orbicularis oris, and to the elevators and depressors FACIAL NERVE, I 457 Fig. 151. FAOlAf. NekvB. of the lips ; that is, to the superficial muscles of the face, which are concerned in the production of expression. (Fig. 151.) ; The facial, consequently, is the- motor nerve of the face. It has nothing to do with transmitting sensitive impressions, since it has been frequently shown that after section of the fifth pair, the facial remaining entire, the sensibility of the face is completely lost ; set that the integument may be cut, pricked, pierced, or lacerated, without any sign of pain being exhibited by the animal. The facial,, therefore, does not transmit sensation from these parts ; and its division, which was formerly resorted to in cases of tic douloureux, is accordingly alto- gether incapable of relieving neuralgic pains. This nerve, however, is directly connected with muscular action, since mechanical or galvanic irritation of its fibres produces con- vulsive twitching in the ears, nostrils, lips, and cheeks. If the facial nerve be divided in one of the lower animals, as, for example, in the cat, immediately after its emergence from the etylo-mastoid foramen, it will be found that complete muscular paralysis has occurred in all those parts to which the nerve is dis- tributed, while the power of sensation remains unimpaired. The animal is incapiable of moving the ear, which remains constantly in the same position. There is also incapacity of closing the eyelids, owing to paralysis of the orbicularis oculi, and the eye accordingly remains constantly open, even when the opposite eye is closed; as during sleep, or in the act of winking. If the conjunctiva be touched, the animal feels the irritation, and endeavors to escape from it ; but the eyeball is only drawn partially backward into the socket by the action of the recti muscles, and the third eyelid pushed partly across the cornea. The complete closure of the eye is impossible. It will be observed, accordingly, that precisely oppo- site effects are produced upon the eyelids by -paralysis of the oculo- motorius nerve, and by that of the facial. In the former instance, owing to the paralysis of the levator palpebr^ superioris, the eye is always partially closed ; in the latter, owing* to paralysis of the 458 • THE OEANIAL NERVES. orbicularis, it is always partially open. The movements of the nares are also suspended on the side of the injury, and if the angle of the mouth be examined on that side, it will be found to hang down lower than on the opposite side, and to be constantly partly open, owing to the paralysis of the orbicularis oris and the eleva- tors of the angle of the mouth. These are the only inconveniences which follow the division of the facial nerve in the cat, but in some other of the lower animals, where various muscular organs in this region are particularly de- veloped, the consequences are more tr&ublesome. Thus, in the rabbit, the ear, upon the affected side, falls down, and cannot be raised or pointed in different directions ; and as the movements of the ear are important in these animals, as aids to the hearing, the per- fection of this sense must be considerably impaired by paralysis of the facial nerve. In the horse, it has been noticed by Bernard,' that division of the facial on both sides is fatal by suffocation. For this animal breathes exclusively through the nostrils, which open widely at the time of inspiration, to allow the admission of air. If these movements be suspended, by paralysis of the facial nerve, the nostrils immediately collapse, and the animal dies by suffocation. In the human subject, the facial nerve is occasionally paralyzed upon one side, sometimes from sympathetic irritation^ sometimes from organic disease in the petrous portion of the temporal bone, or within the cranial cavity near the origin of the nerve. In either case, an extremely well-marked affection is the result, known as " facial paralysis." This condition is chiefly characterized by an entire absence of expression on the affected side of the face. The lower eyelid sinks downward, from paralysis of the orbicularis muscle, and cannot be closed. The corner of the mouth also falls downward, 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 appear- ance. The principal inconvenience, however; suffered by the human subject in facial paralysis, depends upon the want of action of the muscles about the lips and cheek. In drinking, the fluids escape ' Lemons sur la Pliysiologie et la Patkologie da Syst^me Nerreux, Paris, 1858, Tol. ii. p. 38. GLOSSO-PHABYNaEAL NEBVE. 459 by the corner of the mouth, and in mastication the food has partly a tendency to escape "by the same opening, and partly accumulates, on the affected side, between the gums and the cheek, owing to the paralysis of the buccinator muscle, which receives its motor fila- ments from the facial nerve. Thus, the action of all this superficial facial muscles is suspended, the expression of the face is destroyed, and the movements of the lips and the prehension of the food seriously interfered with. 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 these branches are irritated iu 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 their own, but lipon those which they derive 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 experimenter, then, upon another animal, divides the fifth pair within the skull, leaving the facial untouched; and afterward, on irritating as before the ex- posed branches of the latter nerve, he finds that its sensibility has entirely disappeared. It is by filaments, accordingly, derived from the fifth pair, that a certain degree of sensibility is communicated to the branches of the facial. 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. Glosso-Phabyngeal. — This nerve originates from the lateral portion of the medulla oblongata, passes outward, and enters the ' Traitfi de Physiologie, vol. ii. pp. 354-357. 460 THE CBANIAL NERVES, 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 distributed 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 him, as well as by various other observers, to be both sensitive and motor, 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 ofiice of this nerve is to impart the sense of taste to the posterior third of the tongue, to which it "is distributed. It alSo presides over the general sensibility of this part of the tongue, as well as that of the fauces and pharynx. Dr. John Eeid,' who has performed a great variety of experiments upon this nerve, comes to the following conclusions in regard to it. First, that it is essentially a sensitive nerve, since there are unequi- vocal signs of pain when it is pricked, pinched, or ctft. Second, that irritation of this nerve produces convulsive movements of the throat and. lower part of the face ; but that these movements are, in great measure, not direct, but reflex in their character, since they will take place equally well after the glosso-pharyngeal has been divided, if the irritation be applied to its cranial extremity. Third, that this nerve supplies the special sensibility of taste to a portion of the tongue ; but that it is not the exclusive nerve of this sense, since the power of taste remains, after it has been divided on both sides. There are certain reflex actions, furthermore, which take pl^ce 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 • ' In Todd's Cyclopaedia of Anatomy and Physiology, article Glosso-pharyngeal Nerve. PNEUMOGASTRIC NERVE. 461 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 ttie 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. Pneumogasteio. — 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, luiigs, stomach and liver, as well as to several other parts of secondary importance, it has been often known by the name^of the par vagum. The pneumogastric arises, by a number of separate filaments, from the lateral portion of the medulla oblongata, in the groove between 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 pneumogastric nerve." Immediately below the level of this ganglion* 'the nerve receives an important branch of communication from the spinal accessory, and afterward from the facial, the hypoglossal, and the anterior branches of. the first and second cervicals. At its origin, the pneumogastric is exclusively 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 filaments which it has received from the above-men- 462 THE CRANIAL NERVES. Fig. 152. tioned motor nerves. It becomes, 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 bel&ngs. • The ordinary sensibility of the pneu- mogastric nerve, however, as all experi- menters have observed, is exceedingly dull, in comparison with that of the other sensitive cranial nerves. We have often divided this nerve in the middle of the neck, without any distinct manifestation of pain being given by the animal ; and thougB Bernard has found that at some times its sensibility is well marked, while at others it is very indistinct, he is not able to say upon what special physio- logical (3onditions this difference depends. While the pneumogastric, however, is decidedly deficient, as a general rule, in ordinary sensibility, it possesses, as we shall see hereafter, a sensibility of a pecu- liar kind, which is exceedingly important for the maintenance of the vital func- tions. In passing down' the neck, this nerve sends branches to the mucous membrane and muscular coat of the pharynx, ceso- 'phagus, and respiratory passages. Among the most important of these branches are the two laryngeal nerves, viz., the supe- rior and inferior. The superior laryngeal - nerve, which is given off from the trunk of the pneumogastric jnst after it has emerged from the cavity of the skull, passes downward and forward, penetrates the larynx by an opening in the side of the thyro-hyoid membrane, and is distributed to the mucous mem- brane of the larynx and glottis, and also to a single laryngeal mus- cle, viz., the crico-thyroid. This branch is therefore partly mus- cular, but mostly sensitive in its distribution. The inferior laryngeal branch is. given off just after the pneumogastric has entered the Diagram of Pneumogastric N E R y E, wi th its principal branclieH. — I. Pharyngeal brancli. 2. Supe- rior laryngeal. 3 Inferior laryn- geal. ' 4. Pulmonary branches. 6. Stomach. 6. Liver. PNEUMOGASTBIC NERVE. 463 cavity of the chest. It curves round the subclaviaii 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 pos- terior edge of the thyroid, and is distributed to all the muscles of the larynx, with the exception of the crico-thyroid. This branch is, therefore, exclusively muscular in its distribution. The trunk of the pneumbgastrio, after supplying the above branches,, as well as sending numerous filaments to the trachea and oesophagus in* the neck, 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 but and ^re 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 continued in the lower portion of the pharynx and throughout the oesophagus by the aid of the pneumogastric. As the food is compressed by the superior constrictor muscle of the pharynx and forced downward, it excites the mucous membrane with which it is brought in contact and gives rise to another contraction of the middle ^constrictor. The lower constrictor is then brought into action in its turn in a similar manner ; and a wave-like or peristaltic contraction is thence pro- pagated throughout the entire length of the oesophagus, by which the food is carried rapidly from above downward, and conducted at last to the stomach. Bach successive portion of the mucous mem- brane, 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 ii;;i the paralyzed oesophagus, into which it is forced by the voluntary 464 THE CRANIAL NERVES. movements of 'the mouth and fauces, and by the continued 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 sufiicient 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 pro- duces 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 th§ 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, by opening the pharynx and oesophagus on one side, and turning' the larynx for- ward, it will be seen that so long as the vocal chords preserve their usual relaxed condition during expiration, no sound is heard, PNEUMOGASTRIC NERVE. 465 except tlie ordinary faint whisper of the air passing gently through the cavity of the larynx. "When a vocal sound, however, is to be produced, the ch,ords are suddenly made tense and applied closely to* each other, so as to diminish, very considerably 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 pro- duce the sound required. The tone, pitch, and intensity of this sound, vary with the conformation of the larynx, the degree of ten- sion and approximation 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 mouth, 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 paralyzing these muscles, should produce a loss of voice. It has been sometimes found that in very young animals the crico-thyrpid muscles, which are the only ones not affected by division of the inferior laryngeal nerves, are still Buf&cient 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 found 30 466 THE CRANIAL NEBVES. by Bischoff and by Bernard' that if all the roots of the spinal acces- sory 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 laryn- geal itself had been destroyed. All the motor fibres of the pneu- mo'gastrio, 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 superior laryngeal nerve, is reflected back upon the expiratory muscles of the chest and abdomen, by which 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 jn a previous chapter, are of the greatest importance to the preservation of life. "We have seen that 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 that in expiration the muscles and the vocal chords are both relaxed, and the air allowed to pass out readily through 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 inspiratioft.. 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- ' Recherehes Exp6rimentales sur les fonctions du nerf spinal. Paris, 1851. PNEUMOGASTRIC NERVE. 46? 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 off only after the main trunks have entered the cavity of the chest. 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 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 two days old almost instantly suffocated after section of the two inferior laryngeal nerves, We have found that, in pups 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 ' In Longet's Traite de Physiologie, toI. ii. p. 324. 468 THE CRANIAL 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 accessory, the facial, the hypoglossal, 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 princ;ipally 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 they 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 ligaturp, may be instantaneously divided, and the effects of the operation readily appreciated. Immediately after the division of the nerves, when performed ia the above manner, the respiration is hurried and difficult, 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- PNEUMOGA.STSIC NERVE. 469 tion is a diminished frequency 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 efibrt, 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 paii;. 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 470 THE CRANIAL NERVES. corresponding with the different degrees of. alteration exhibited bj 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 suiBcient. It accounts very well for the diminished frequency of 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. PNECJIOGASTEIC NBBVB. 471 therefore, to understand wHy respiration should be retarded, after section of the pneumogastrics, since the chief source of the stimulus to respiration is cut ofi'; 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, is no doubt 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 evi- dently 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 sec- tion 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 difficulty, either alone or combined with the diminished fre- quency of respiration, must have a very considerable effect in im- peding 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 difference in the condition of the two animals. There was the same obstruction of the breath (owing to closure of the glottis), the same gasping and sucking inspiration, and' the same frothing at the mouth. Very soon, however, in pup No. 1, the respiratory movements 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 respiration continued frequent as well as laborious, and the general signs of agitation and discomfort were kept up for one or two hours. 472 THE CRANIAL NEKVJfiS. i 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 pneumogastric 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 less frequently than usual. From these two causes combined, the blood is imperfectly arterialized. But the usual consequence of such a condition, viz., an increased rapidity of the respiratory movements, does not follow. The imperfect arterialization of the blood 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 in the tissues begins to exert a narcotic effect, diminishing the sensibility of the nervous centres, and tending to retard still more the movements of respiration. Thus all these causes react upon and aggravate each other ; because the connec- tion, naturally existing between imperfectly arterialized blood and the stimulus to respiration, is now destroyed. The narcotism and PNEUMOGASTBIO NEBVE. 473 pulmonary 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 pneumogastrio 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 pneumogastrio 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. Stomach, and Digestive Function. — After division of the pneumo- gastrio 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 pneumogastrio nerve. Digestion, however, very seldom 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 act 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 474 THE CBANIAL NEBVES. 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, whUe 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 its 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 nerve, 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 efiect 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. Spinal Acoessoet. — This nerve originatfes, by many filaments, from the side of the medulla oblongata, below the level of the pneumogastric, and also from the lateral portions of the spinal cord, between the anterior and posterior roots of the upper five or six cervical nerves. These fibres of spinal origin pass upward, uniting into a slender rounded filament, which enters the cavity of the cranium by the foramen magnum, and is then joined by the fibres which originate from the medulla oblongata. The spinal accessory nerve, thus constituted, passes qut from the cavity of the skull by the posterior foramen lacerum, in company with the glosso-pharyn- geal and pneumogastric nerves. Immediately afterward it divides SPINAL ACCESSORY. 475 into two principal branches: First the internal or anastomotic branch, which joins the pneumogastric nerve, and becpmes mingled with its fibres; and, secondly> the external or muscular branch, which passes downward and outward, and is distributed to the sterno-mastoid and trapezius muscles. The spinal accessory is essentially a motor nerve. It has been found, both by Bernard and Longet, to be insensible at its origin, like the anterior roots of the spinal nerves ; but if irritated after its exit from the skull, it gives signs of sensibility. This sensibi- lity it acquires from the filaments of inosculation which it receives from the anterior branches of the first and second cervical nerves. Though its external branch, accordingly, is exclusively distributed to muscles, as we have already seen, this branch contains some sensi- tive fibres, which have the same destination. The reason for this anatomical fact, viz., that motor nerves are supplied during their course with sensitive fibres, becomes evident when we reflect that the muscles themselves possess a certain degree of sensibility, though less acute than that which belongs to the skin. The sensibility of the muscles is undoubtedly essential to the perfect performance of their function ; and as the motor nerves are incapable, by them- selves, of transmitting sensitive impressions, they are joined, soon after their origin, by other filaments which communicate to them this necessary power. The most important result which has been obtained by experi- ment upon the spinal accessory nerve, is that its internal or anasto- motic branch is directly connected vdth the vocal movements of the glottis. It has been found by Bischofif, by Longet, and by Bernard, that if the spinal accessory nerves on both sides, or their branches of inosculation with the pneumogastric, be divided or lacerated, the pneumogastric nerves themselves being left entire, the voice is instantly lost, and the animal becomes incapable of making a vocal sound. We have also found this result to follow, in the cat, after the spinal accessory nerves have been torn out by their roots, through the jugular foramen. The animal, after this operation, can no longer make an audible sound. At the same time the respira- tory movements of the glottis go on undisturbed, and most of the other animal functions remain unaffected. The fibres of communication, therefore, derived from the spinal accessory, pass to the pneumogj^stric nerve and become entangled with its other filaments, so that they can no longer be traced by anatomical dissection. They pass downward, however, aiid become 476 THE CRANIAL NERVES. a part of the motor fibres of the inferior laryngeal or recurrent branches of the pneumogastric ; being finally distributed to the muscles of the larynx, which they supply with those nervous influ- ences which are required for the formation of the voice. The special function of the external or muscular branch of the spinal accessory is not so fully understood. This branch, as we have seen, is distributed to the stemo-mastoid and trapezius mus- cles. But these muscles also receive filaments from the cervical spinal nerves ; and, accordingly, they still retain the power of mo- tion, to a certain degree, after the external branches of the spinal accessory have been divided on both sides. The spinal accessory is, accordingly, a n^rve of very peculiar distribution. For it partly supplies motor fibres to the pneumo- gastric nerve, and is partly distributed to two muscles, both of which also receive motor nerves fifom another source. Sir Charles Bell, noticing the close connection between this nerve and the pneumogastric, regarded the two as associated also in their func- tion, as nerves of respiration. He considered, therefore, the exter- nal branch of the spinal accessory as destined to assist in the movements of respiration, when these movements become unusu- ally laborious, by bringing into play the sterno-mastoid and trape- zius muscles, in aid of the action of the intercostals. He therefore called this nerve the " superior respiratory nerve." But the most satisfactory explanation of this peculiarity is that proposed by M. Bernard. According to this explanation, whenever a muscle, or set of muscles, derive their nervous influence from two different sources, this is not for the purpose of assisting them in the performance of the same function, but of enabling them to perform two different functions. We have seen this already exemplified in the muscles of the larynx. For these muscles perform certain movements of respiration for which they receive indirectly filaments from the facial hypoglossal, and cervical nerves. But they also perform the movements necessary to the formation of the voice, the nervous stimulus for which is derived altogether from the spinal accessory. The internal branch of the spinal accessory, accordingly, excites, in the parts to which it is distributed a function which is incompa' tible with respiration. For the movements of respiration cannot go on while the voice is sounded; and a necessary preliminary to the production of a vocal sound, is the temporary stoppage of respiration. The movements of respiration, therefore, and the HYPOGLOSSAL. 477 movements of the voice alternate with each other, but are never simultaneous ; so that the internal branch of the spinal accessory is antagonistic to the motor fibres of the larynx derived from other nerves. It is thought by M. Bernard, that the fibres of the external branch of the spinal accessory, have also a function which is anta- gonistic to respiration. For respiration is naturally suspended in all steady and prolonged muscular efforts. In these efforts, such as those of straining, lifting, and the like, the movements of respira- tion cease, the spinal column is made rigid by the contraction of its muscles, and the head and neck are placed in a fixed position, principally by the contraction of the sterno-mastoid and trapezius muscles. The function of the spinal accessory, in both its branches, is therefore regarded as destined to excite movements which are incompatible with those of respiration ; and which accordingly come into play only when the ordinary movements of respiration have been temporarily suspended. Hypoglossal. — The hypoglossal nerve originates from the ante- rior 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 affect- ing directly the sensibility of its mucous membrane. This nerve, acQordingly, is the motor nerve of the tongue. If irritated at its origin, the hypoglossal 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 sensibility, like that of the facial, is consequently derived from its , inosculation with other sensitive nerves, after its emergence from the skull. 4:78 THE SPECIAL SENSES. CHAPTER VI. THE SPECIAL SENSES. Geneeal and Special Sensibility. — We have already seen that there exists, in the general integument, a power of sensation, by which we are made acquainted with surrounding objects and some of their most important physical qualities. By this power we feel the sensations of heat and cold, and are enabled to distinguish between hard and sofl; substances, rough bodies and smooth, solids and liquids. This kind of power is termed General Sensibility, because it resides in the general integument, and because by its aid we obtain information with regard to the simplest and most material properties of external objects. The general sensibility, thus existing in the integument, is an endowment of the sensitive nerves derived from the cerebro-spinal , system. These nerves ramify in the substance of the skin, and by subsequent inosculation form a minute plexus in the superficial portions of the tissue of the corium. Froni this plexus, the ulti- mate filaments, reduced to an exceedingly minute size, pass up- ward into the conical ' papillae with which the free surface of the corium is covered. In the papilla the nervous filaments terminate, sometimes by loops returning upon themselves, and sometimes ap- . parently by free extremities. The papillae are also supplied with looped capillary bloodvessels, and are capable of receiving an abundant vascular injection. These papillae appear to be the most essential organs of general sensation, since the sensibility of the skin is most acute where they are most abundant and most highly developed, as, for exampl6, on the palm of the hand and the tips of the fingers. The best method of measuring accurately the sensibility of dif- ferent regions is that adopted by Professors Weber and Valentin. They applied the rounded points of a pair of compasses to the integument of difierent parts, and found that if they were held very near together they could no longer be distinguished as sepa- GENERAL AND SPECIAL SENSIBILITY. 479 rate points, but the two sensations were confounded into one. The distance, however, at which the two points failed to be distinguished from each other, was much shorter for some parts of the body than for others. 'Prof. Valentin's measurements,^ which are the most varied and complete, give the following as the limits of dis- tinct perception in various parts : — Paris Lihe. At the tip of tongue .483 " palmar surface of tips of fingers .... .723 " " " of second phalanges . . . 1.558 " " " of first phalanges .... 1.650 " dorsum of tongue 2.500 " dorsal surface of fingers 3.900 " cheek 4.541 " back of hand 6.966 " skin of throat 8.292 " dorsum of foot 12.525 " skin over sternum 15.875 " middle of back . . • 24.208 This method cannot, of course, give the absolute measure of the acuteness of sensibility in the different regions, since the' two points might be less easily distinguished from each other in any one re- gion, and yet the absolute amount of sensation produced might be as great as in the surrounding parts ; still it is undoubtedly a very accurate measure of the delicacy of tactile sensation, by which we are enabled to distinguish slight inequalities in the surface of solid bodies. We find, furthermore, that certain parts of the body are particularly well adapted to exercise the function of general sen- sation, not only on account of the acute sensibility of their integu- ment, but also owing to their peculiar formation. Thus, in man, the hands are especially well formed in this respect, owing to the articulation and mobility of the fingers, by which they may be adapted to the surface of solid bodies, and brought successively in contact with all their irregularities and depressions. The hands are therefore more especially used as organs of touch, and we are thus enabled to obtain by their aid the most delicate and precise information as to the texture, consistency,. configuration, &c., of foreign bodies. But the hands are not the exclusive organs of touch, even in the human subject, and in some of the lower animals, the same func- ' In Todd's Cyclopaedia of Anatomy and Physiology, vol. iv., article on Touch, by Dr. Carpenter. 480 THE SPECIAL SENSES. tion is fully performed by various otter parts of the body. Thus in the cat and in the seal, the long bristles seated upon the lips are used for this purpose, each bristle being connected at its base with a highly developed nervous papilla : in some of the monkeys the extremity of the prehensile tail, and in the elephant the end of the nose, which is developed into a flexible and sensitive proboscis, is employed as an organ of touch. This function, therefore, may be performed by either one part of the body or another, provided the accessory organs be developed in a favorable manner. About the head and face, the sensibility of the skin is dependent mainly upon branches of the fifth pair. In the neck, trunk, and extremities it is due to the sensitive fibres of the cervical, dorsal, and lumbar spinal nerves. It exists also, to a considerable extent, in the mucous membranes of the mouth and nose, and of the pas- sages leading from them to the interior of the body. In these situations, it depends upon the sensitive filaments of certain of the cranial nerves, viz., the fifth pair, the glosso-pharyngeal, and the pneumogastric. The sensibility of the mucous membranes is most acute in those parts supplied by branches of the fifth pair, viz., the conjunctiva, anterior part of the nares, inside of the lips and cheeks, and the anterior two-thirds of the tongue. At the base of the tongue and in the fauces, where the mucous membrane is supplied by filaments of the glosso-pharyngeal nerve, the general sensibihty is less perfect; and finally it diminishes rapidly from the upper part of the oesophagus and the glottis toward the stomach and the lungs. Thus, we can appreciate the temperature and consistency of a foreign substance very readily in the mouth and fauces, but these qualities are less distinctly perceived in the oesophagus, and not at all in the stomach, unless the foreign body happen to be excessively hot or cold, or unusually hard and angular in shape. The general sensibility, which is resident in the skin and in a certain portion of the mucous membranes, diminishes in degree from with- out inward, and disappears altogether in those organs which are not supplied with nerves from thq cerebro-spinal system. It is particularly to be observed, however, that while the general sensibility of the skin, and of the mucous membranes above men- tioned, varies in aouteness in different parts of the body, it is every- where the same in kind. The tactile sensations, produced by the contact of a foreign body, are of precisely the same nature whether they be felt by the tips of the fingers, the dorsal or palmar surfaces of the hands, the lips, cheeks, or any other part of the integument. TASTE. 4.81 The only difference in tlie sensibility of these parts lies in the de- gree of its development. But there are certain other sensations which are different in kind from those perceived by the general integutnent, and which, owing to their peculiar and special character, are termed special sensations. Such are, for example, the sensation of light, the sensation of sound, the sensation of savor, and the sensation of odors. The special sensibility which enables us to feel the impressions derived from these sources is not distributed over the body, like ordinary sensi- bility, but is localized in distinct organs, each of which is so con- stituted as to receive the special sensation peculiar to ity and no other. Thus we have, beside the general sensibility of the skin and mucous membranes, certain peculiar faculties or special senses, as they are called, which enable us to derive information from ex- ternal objects,*which we could not possibly obtain by any other means. Thus light, however intense, produces no perceptible sen- sation when allowed to fall upon the skin, but only when admitted to the eye. The sensation of sound is perceptible only by the ear, and that of odors only by the olfactory membrane. These different sensations, therefore, are not merely exaggerations of ordinary sensibility, but are each distinct and peculiar in their nature, and are in relation with distinct properties of external objects. In examining the organs of special sense, we shall find that they each consist — First, of a nerve, endowed with the special sensibility required for the exercise of its peculiar function ; and, Secondly, of certain accessory parts, forming an apparatus more or less compli- ' cated, which is intended to assist in its performance and render it more delicate and complete. We shall take up the consideration of the special senses in the following order. First, the sense of Taste ; second, that of Smell ; third, that of Sight ; and fourth, that of Hearing, Taste. — We begin the study of the special senses with that of Taste, because this sense is less peculiar than any of the others, and differs less, both in its nature and its conditions, from the ordinary sensibility of the skin. In the first place, the organ of taste is no other than a portion of the mucous membrane, beset with vascular and nervous papillEe,' similar to those of the general integument. 31 482 THE SPECIAL SENSES. Secondly, it gives us impressions of such substances only as are actually in contact with sensitive surfaces, and can establish no communication with objects at a distance. Thirdly, the surfaces which exercise the sense of taste are also endowed with general sen- sibility ; and Fourthly, there is no one special and distinct nerve of taste, but this property resides in portions of two different nerves, viz., the fifth pair; and the glosso-pharyngeal ; nerves which also supply general sensibility to the mouth and surrounding parts. The sense of taste is localized in the mucous membrane of the tongue, the soft palate, and the fauces. The tongue, which is more particularly the seat of this sense, is a, flattened, leaf-like, muscular organ, attached to the inner surface of the symphysis of the lower jaw in front, and to the os hyoides behind. It has a vertical sheet or lamina of fibrous tissue in the median line, which serves as a framework, and is provided with an abundance of longitudinal transverse and radiating muscular fibres, by which it can be elon- gated, retracted, and moved about in every direction. The mucous membrane of the fauces and posterior third of the tongue, like that lining the cavity of the mouth, is covered with minute vascular papillae, similar to those of the skin, which are, however, imbedded and concealed in the smooth l^yer of epithe- lium forming the surface of the organ. But about the junction of its posterior and middle thirds, there is, upon the dorsum of the tongue, a double row of rounded eminences, arranged in a V-shaped figure, running forward and outward, on each side, from the situa- tion of the foramen caecum ; and, from this point forward, the upper surface of the organ is everywhere covered with an abundance of thickly-set, highly' developed papillae, projecting from its surface, and readily visible to the naked eye. These lingual papillae are naturally divided into three different sets or kinds. First, the filiform papillae^ which are the most nume- rous, and which cover most uniformly the upper surface of the organ. They are long and slender, and are covered with a some- what horny epithelium, usually prolonged at their free extremity into a filamentous tuft. At the edges of the tongue these papilla are often united into parallel ranges or ridges of the mucous mem- brane. Secondly, the fungiform papillae. These are thicker and larger than the others, of a rounded club-shaped figure, and covered with soft, permeable epithelium. They are most abundant at the tip of the tongue, but may be seen elsewhere on the surface of the organ, scattered among the filiform papillae. Thirdly, the circum- TASTE. 483 vallate papillae. These are the rounded eminences whicli form the V-shaped figure near the situation of the foramen cseoum. They are eight or ten in number. Each one of them is surrounded by a circular wall, or circumvallation, of mucous membrane, which gives to them their distinguishing appellation. The circumvalla- tion, as well as the central eminence, has a structure similar to that of the fungiform papillsB. The sensitive nerves of the tongue, as we have already seen, are two in number, viz., the lingual branch of the fifth pair, and the lingual portion of the glosso-pharyngeal. The lingual branch of the fifth pair enters the tongue at the anterior border of the hyo- glossus muscle, and its fibres then run through the muscular tissue of the organ, from below upward and from behind forward, with- out any ultimate distribution, until they reach the mucous mem- brane. The nervous filaments then penetrate into the lingual papillae, where they finally terminate. The exact mode of their termination is not positively known. According to Kolliker, they sometimes seem to end in loops, and sometimes by free extremities. The lingual portion of the glosso-pharyngeal nerve passes into the tongue below the posterior border of the hyo-glossus muscle. It tlien divides into various branches, which pass through the mus- cular tissue, and are finally distributed to the mucous membrane of the base and sides of the organ. Pig. 153. DlAQRAM OF ToNODB, With itJiseD-sItive nervesand papUljB.— 1. Lint'ual branch of fifth pnir, 2. Glosao-pharjogeaL nerve. The mucous membrane of the base of the tongue, of its edges, and its under surface near the tip, as well as the mucous membrane of the mouthvand fauces generally, is also supplied with mucous follicles, which furnish a viscid secretion by which the free surface of the parts is lubricated. 484 THE SPECIAL SENSES. Finally, the muscles of the tongue, it will.be remembered, are animated exclusively by the filaments of the hypoglossal nerve. The exact seat of the sense of taste has been determined by placing in contact with different parts of the mucous membrane a small sponge, moistened with a solution of some sweet or bitter substance. The experiments of Verni^re, Longet and others have shown that the sense of taste resides in the whole superior surface, the point and edges of the tongue, the soft palate, fauces, and part of the pharynx. The base, tip, and edges of the tongue seem to possess the most acute sensibility to savors, the middle portion of its dorsum less of this sensibility, and its inferior surfaces little or none. Now as the whole anterior p^t of the organ is supplied by the lingual branch of the fifth pair alone, and the whole of its posterior portion by the glosso-pharyngeal, it follows that the sense of taste, in these different parts, is derived from these two different nerves. Furthermore, thd tongue is supplied, at the same time and by the same nerves, with general' sensibility and with the special sensibility of taste. The general sensibility of the anterior portion of the tongue, and that of the branch of the fifth pair with which it is supplied, are sufficiently well known. Section of the fifth pair destroys the sensibility of this part of the tongue as well as that of the rest of the face. Longet has found tfiat after the lingual branch of this nerve has been divided, the mucous membrane of the anterior two- thirds of the tongue may be cauterized with a hot iron or with caustic potassa, in the living animal, without" producing any sign of pain. Dr. John Eeid, on the other hand, together with other experi- menters, has determined that ordinary sensibility exists in a marked degree in the glosso-pharyngeal, and is supplifed by it to the parts to which this nerve is distributed. Accordingly we must distinguish, in the impressions produced by foreign substances taken into the mouth, between the special impressions derived from their sapid qualities, and the general sensa- tions produiced by their ordinary physical properties. As the tongue is exceedingly sensitive to ordinary impressions, and as the same body is often capable of exciting both the tactile and gustatory functions, these two properties are sometimes liable to be confounded with each other by careless observation. The truly sapid qualities, however, the only ones, properly speaking, which we perceive by the sense of taste, are such savors as we designate by the term sweet, bitter, salt, sour, alkaline, and the like. But there are many TASTE. 485 Other properties, belonging to various articles of food, which belong really to the class of ordinary physical qualities and are appre- ciated by the ordinary sensibility of the tongue, though we usually speak of them as being perceived by the taste. Thus a siaircTiy, viscid, watery, or oleaginous taste is merely a certain variety of con- sistency in the substance tasted, which may exist either alone or in connection with real savors, but which is exclusively perceived by means of the general sensibility. So also with a pungent or burning' taste, such as that of red pepper or any other irritating powder. The quality of piquancy in the preparation of artificial kinds of food is always communicated to them by the addition of some such irritating substance. The styptic taste seems to be a corabination of an ordinary irritant or astringent effect with a peculiar taste, which we always associate with the former quality in astringent sub- stances. There is also sometimes a liability to confoniid the real taste of certain substances with their odorous properties, or Jlavors. Thus in most aromatic articles of food, such as tea and coffee, and in various kinds of wine, a great part of what we call the taste is in rea,lity due to the aroma, or smell which reaches the nares during the act of swallowing. Even in many solid kinds of food, such as freshly cooked meats, the odor produces a very important part of their effect on the senses. We can easily convince ourselves of this by holding the nose while swallowing such substances, or by recol- lecting how much a common catarrh interferes with our perception of their taste. The most important conditions of the sense of taste are the fol- lowing : — In the first place, the sapid substance, in order that its taste may be perceived, must be brought in contact with the mucous mem- brane of the mouth in a state of solution. So loijg as it remains solid, however marked a savor it -may possess, it gives no other impression than that of any foreign body in contact with the sensi- tive surfaces. But if it be applied in a liquid form, it is then spread over the surface of the mucous membrane, and its taste is imme- diately perceived. Thus it is only the liquid and soluble portions of our food which are tasted, such as the animal and vegetable juices and the soluble salts. Saline substances which are insoluble, Hich as calomel or carbonate of lead, when applied to the tongue, produce no gustatory sensation whatever. The mechanism of the sense of taste is, therefore, in all proba- THE SPECIAL SENSES, bility, a direct and simple one. The sapid substances in solution penetrate the lingual papillae by endosmosis, and, coming in actual contact with the terminal nej-vous filaments, excite their sensibility by uniting with their substance. We have already seen that the rapidity with which endosmosis will take place under certain con- ditions is sufficiently great to account for the almost instantaneous perception of the taste of sapid substances, when introduced into the mouth. It is on this account that a free secretion of the salivary fluids is so essential to the full performance of the gustatory function. If the mouth be dry and parched, our food seems to have lost its taste ; but when the saliva is freely secreted, it is readily mixed with the food in mastication, and assists in the solution of its sapid ingredi- ents ; and the fluids of the mouth, thus impregnated with the savory substances, are absorbed by the mucous membrane, and excite the gustatory nerves. An important part, a,lso, is taken in this process by the movements of the tongue ; for by these movements the food is carried from one part of the mouth to another, pressed against the hard palate, the gums, and the cheeks, its solution assisted, and the penetration of the fluids into the substance of the papillae more rapidly accomplished. If a little powdered sugar, or some vege- table extract be simply placed upon the dorsum of the tongue, but little effect is produced ; but as soon as it is. pressed by the tongue against the roof of the mouth, as naturally happens in eating or drinking, its taste is immediately perceived. This effect is easily explained; since we know how readily movement over a free sur- face, combined with slight friction, will facilitate the imbibition of liquid substances. The nervous papillae of the tongue may there- fore be regarded as the essential organs of the sense of taste, and the lingual muscles as its accessory organs. The full effectfof sapid svhstances is not obtained until they are actu- ally swallowed. During the preliminary process of mastication a sufScient degree of impression is produced to enable us to perceive the presence of any disagreeable or injurious ingredient in the food, and to get rid of it, if we desire. But it is only when the food is carried backward into the fauces and pharynx, and is compressed by the constrictor muscles, of these parts, that we obtain a complete perception of its sapid qualities. For at that time the food is spread out by the compression of the muscles, and brought at once in contact with" the entire extent of the mucous membrane possessing gustative sensibility. Then, it is no longer under the control of the TASTE. 487 will, and is carried by the reflex actions of the pharynx and oeso- phagus downward to the stomacb. The impressions of taste made upon the tongue remain for a cer- tain time afterward. When a very sweet or very bitter substance is taken into the mouth, we retain the taste of its sapid qualities for several seconds after it has been ejected or swallowed. Conse- quently, if several different savors be presented to the tongue in rapid succession, we soon become unable to distinguish them, and they produce only a confused impression, made up of the union of various different sensations ; for the taste of the first, remaining in the mouth, is mingled with that of the second, the taste of these two with that of the third, and so on, until so mauy savors become confounded together that we are no longer able to recognize either of them. Thus it is notoriously impossible to recognize two or three different kinds of wine with the eyes closed, if they be repeat- edly tasted in quick succession. K the stibstance first tasted have a particularly marked savor, its taste will preponderate over that of the others, and perhaps pre- vent our recognizing them at all. This effect is still more readily produced by substances which excite the general sensibility of the tongue, such as acrid or stimulating powders. In the same manner as a painful sensation, excited in the skin, prevents the nerves, for the time, from perceiving delicate tactile impressions, so any pungent or irritating substance, which excites unduly the general sensibility oft the tongue, blunts for a time its special sensibility of taste. This effect is produced, however, in the greatest degree, by substances which are at the same time sapid, pungent and aromatic, like sweet- meats flavored with peppermint. Advantage is sometimes taken of this in the administration of disagreeable medicines. By first taking into the mouth some highly flavored and pungent substance, nauseous drugs may be swallowed immediately afterward with but little perception of their disagreeable qualities. A very singular fact, in connection with the sense of taste, is that it is sometimes affected in a marked degree by paralysis of the facial nerve. No less than six cases of this kind, occurring in the human subject, have been collected by M. Bernard, and we have also met with a similar instance in which the peculiar phenomena were well marked. M. Bernard has furthermore seen a similar effect upon the taste produced in animals by division of the facial nerve within the cranium. The result of' these experiments and observations is as follows : When the facial nerve is divided or seriously injured 483 THE SPECIAL SENSES. by organic disease, before its emergence from the stylo-mastoid foramen, not only is there a paralysis of the superficial muscles of the face, but the sense of taste is diminished on the corresponding side of the tongue. If the tongue be protruded,.and salt, citric acid or sulphate of quinine be placed upon its surface on the two sides ' of the median line, the taste of these substances is perceived on the affected side more slowly and obscurely than on the other. It is not, therefore, a destruction, but only a diminution of the sense of taste, which follows paralysis of the facial in these instances. At the same time the general tactile sensibility of the tongue is unal- tered, retaining its natural acuteness on both sides of the tongue. The exact mechanism of this peculiar influence of the facial nerve upon the sense of taste is not perfectly understood. It may be considered as certain, however, that it is derived through the medium of that branch of the facial nerve known as the chwda tympani. This filament , leaves the facial at the intumescentia gangliformis, in the interior of the aqueduct of Fallopius, enters the cavity of the tympanum, passes across the membrane of the tym- panum, and then, emerging froji the cranium, runs downward and , forward and joins the lingual branch of the fifth pair. It then ac- companies this nerve as far as the posterior extremity of the sub- maxillary gland. Here it divides into two portions ; one of which passes to the submaxillary ganglion, and, through it, to the sub- stance of the submaxillary gland, while the other continues onward, still in connection with the lingual branch of the fifth pair, and, in company with the filaments of this nerve, is distributed to the tongue. The chorda tympani thus forms the only anatomical connection between the facial nerve and the anterior part of the tongue. When the facial, accordingly, is divided or injured after its emergence from the stylo-mastoid foramen, no effect is produced upon the sense of taste ; but when it is injured during its course through the aqueduct of Fallopius, and before it has given off the chorda tym- pani, this nerve suffers at the same time, and the sense of taste is diminished in activity, as above described. It is probable that this effect is produced in an indirect way, by a diminution in the activity of secretion in the lingual follicles, or by some alteration in the vascularity of the parts. Smell. — The main peculiarity of the sense of smell consists in the fact that it gives us intelligence of the physical chariacter of bodies in a gaseous or vaporous condition. Thus we are enabled to SMELL. 489 perceive the existence of an odorous substance at a distance, and when it is altogether concealed from sight. The minute quantity of volatile material emanating ttora it, and thus pervading the atmosphere, comes in contact with the mucous membrane of the nose, and produces a peculiar and special sensation. The apparatus of this sense consists, first, of the olfactory mem- brane, supplied by the filaments of the olfactory nerve, as its special organ ; and secondly, of the nasal passages, with the tur- binated bones and the muscles of the anterior and posterior nares, as its accessory organs. At the upper part of the nasal fossae, the mucous membrane is very thick, soft, spongy and vascular, and is supplied with mucous follicles which exude a secretion, by which its surface is protected and kept in a moist and sensitive condition. It is only this portion of the mucous membrane of the nares which is supplied by filaments of the olfectory nerve, and which is capable of receiving the impressions of smell ; it is therefore called the Olfactorp membrane. Elsewhere, the nasal passages are lined with a mucous membrane which is less vascular and spongy in structure, and which is called the Schndderian membrane. The filaments derived from the olfaetory ganglia, and which penetrate through the cribri- form plate of the ethmoid bone, are distributed to the mucous membrane of the su- perior and middle turbinated bones, and to that of the upper part of the septum nasi. The exact mode in which these filaments terminate in the ol- factory membrane has not been definitely ascertained. They are of a soft consistency and gray color, and, after di- viding and ramifying freely in the membrane, appear to become lost in its substance. It is these nerves which exer- cise the special function of smell. They are, to all appearance, incapable of receiving ordinary impressions, and must be regarded as entirely peculiar in their Fig. 154. Distribution of Nertes ik the Nabal Passaoeb. — 1, Olfactory ganglion, withita nerves 2. Naitat branch of fifth pair. 3. Spheno-palatiue ganglion. 490 THE SPECIAL SENSES. nature and endowments. The nasal passages, however, are supplied with other nerves beside the olfactory. The nasal branch of the ophthalmic division of the fifth pair, after entering the, anterior part of the cavity of the nares, just ia advance of the cribriform plate of the ethmoid bone, is distributed to the mucous membrane of the in- ferior turbinated bone and the inferior meatus. Thus the organ of smell is provided with sensitive nerves from two different sources, viz., at its upper part, with the olfactory nerves proper, deriyeid from the olfactory ganglion (Fig. 154, i), which are nerves of special sensation ; and secondly, at its lower part, with the nasal branch of the fifth pair (a), a nerve of general sensation. Beside which, the spheno-palatine ganglion of the great sympathetic (3) sends, fila- ments to the mucous membrane of the whole posterior part of the nasal passages, and to the levator palati and azygos uvulae muscles. Finally, the muscles of the anterior nares are supplied by filaments of the facial nerve. The conditions of the sense of smell are much more special in their nature than those of taste. For, in the first place, this sense is excited, not by actual contact with the foreign body, but only with its vaporous emanations ; and the quantity of these emanations, sufi&cient to lexcite the smell, is often so minute as to be altogether inappreciable by other means. We cannot measure the loss of weight in an odorous body, though it may affect the atmosphere of an entire house, and the senses of .all its inhabitants, for days and weeks together. Secondly, in the olfactory organ, the special sensibility of smell and the general sensibility of the mucous mem- brane are separated from each other and provided for by different nerves, not mingled together and exercised by the same nerves, as is the case in the tongue. In order to produce an olfactory impression, the emanations of the odorous body must be drawn freely tKrough the nasal passages. As the sense of smell, also, is situated only in the upper part of these passages, whenever an unusually faint or delicate odor is to be per- ceived, the air is forcibly directed upward, toward the superior turbinated bones, by a peculiar inspiratory movement of the nos- trils. This movement is very marked in many of the lower .animals. As the odoriferous vapors arrive in the upper part of the nasal passages, they are undoubtedly dissolved in the secretions of the olfactory membrane, and thus brought into relation with its nerves. Inflammatory disorders, therefore, interfere with the sense of smell, both by checking or altering the secretions of the part, and by SMELL. 491 producing an unnatural tumefaction of the mucous membrane, whicli prevents the free passage of the air through the nasal fossae. As in the case of the tongue, also, we must distinguish here between the perception of true odors, and the excitement of the general sensibility of the Schneiderian mucous membrane by irri- tating substances. Some of the true odors are similar in their nature to impressions perceived by the sense of taste. Thua we have sweet and sour smells, though none corresponding to the alkaline or the bitter tastes. Most of the odors, however, are of a very peculiar nature and are difficult to describe ; but they are always distinct from the simply irritating properties which may belong to vapors as well as to liquids. Thus, pure alcohol has little or no odor, and is only irritating to the mucous membrane ; while the odor of wines, of cologne water, &c., is commlinicated to them by the presence of other ingredients of a vegetable origin. In the same way, pure acetic acid is simply irritating ; while vinegar has a peculiar odor in addition, derived from its vegetable impurities. Ammonia, also, is an irritating vapor, but contains in itself no odoriferous principle. The sensations of smell, like those of taste, remain for a certain time after they have been produced, and modify in this way other less strongly marked odors which are presented afterward. As a general thing, the longer we are exposed to a particular odor, the longer its effect upon our senses continues ; and in some cases it may be perceived many hours after the odoriferous substance has been removed. Odors, however, are particularly apt to remain after the removal or destruction of the source from which' they were derived, owing to their vaporous character, and the facility with which they are entangled and retained by porous substances, such as plastered walls, woollen carpets, and hangings, and woollen clothes. It is supposed to be in this way that the odor of a post- mortem examinatioii will sometimes remain so as to be perceptible for several hours or even an entire day afterward. But this alone does not fully explain the fact. For if it depended simply on the retention of the odor by porous substances, it would afterward be perceived constantly, until it gradually and continuously wore off; while in point of fact, the physician who has made an autopsy of this kind does not afterward perceive its odor constantly, but only occasionally, and by sudden and temporary fits. The explanation is probably this. As the odor ' remains con- stantly by us, we soon become insensible to its presence, as in the 492 THE SPECIAL SENSES. case of all other continuous and unvarying impfessions. Our at- tention is only called to it when we meet suddenly with another and familiar odor. This second odor, we find, does not produce its usual impression, because it is mingled with and modified by the other, which is more persistent and powerful. Thus we are again made aware of the former one, to which we had become insensible by reason*of its constant presence. The sense of smell is comparatively feeble in the human species, but is excessively acute in some of the lower animals. Thus, the dog will not only distinguish different kinds' of game in the forest by this sense, and follow them by their tracks, but will readily dis- tinguish particular individuals by their odor, and will recognize articles of dress belonging to them by the minute quantity of odor- iferous vapors adhering to their substance. Sight. — The sight undoubtedly occupies the first rank in the list of special nervous endowments. It is the most peculiar in its operation, and the most immaterial in its nature, of all the senses, and it is through it that we receive the most varied and valuable impressions. The physical agent, also, to which the organ of sight is adapted, and by which its sensibility is excited, is more subtle and peculiar than any of those which act upon our other senses. For the senses of touch, taste, and smell require, for their exercise, the actual contact of a foreign body, either in a solid, liquid, or aeriform condition ; and even the hearing depends upon the me- chanical vibrations of the atmosphere, or some other sonorous medilim. But the eye does not need to be in contact with the luminous body. It will receive the impressions of light with per- fect distinctness, ev,en when they are transmitted from an immea- surable distance, as in the case of the fixed stars ; and the light itself is not only immaterial in its nature, so far as we can ascertain, but is also capable of being transmitted through space without the intervention of any material conducting medium, yet discoverable. Finally, the apparatus of vision is more complicated in its struc- ture than that of any other of the^speoial senses. This apparatus eonsists, first, of the retina, as a special sensitive nervous membrane ; and secondly, of the vitreous body, crystalline lens, choroid, scle- rotic, iris and cornea, together with the muscles moving the eye- ball and eyelids, lachrymal gland, &c., as accessory organs. The arrangement of the parts, constituting the globe of the eye, is shown in the following figure. (Fig. 155.) SIGHT. 493 The filaments of the optic nerve, after running forward and pene- trating the posterior part of the eyeball, spread out into the sub- stance of the retina (3), thus forming a delicate and vascular nerv Fig. 155. Vertical Section of the Eyeball. — 1. Sclerotic. 2. Choroid. 3. Retina. 4. Leos. 5. Hyalciil membrane. 6. Goraea. 7. Iris, 8. Ciliary maBCle and processes. ous expansioh, in the form of a spheroidal bag or sac, with a wide opeiiing in front, where the retina terminates at the posterior mar- gin of the ciliary body. This expansion of the retina is the essen- tial nervous apparatus of the eye. It is endowed with the special sensibility which renders it capable of receiving luminous impres- sions; and, so far as we have been able to ascertain, it is incapable of perceiving any other. On the outside, the retina is covered by the choroid coat (s), a vascular membrane, which is rendered opaque by the presence of an abundant layer of blackish-brown pigment-cells, and which thus absorbs, the light which has once passed through the retina, and prevents its being reflected in such a way as to confuse and dazzle the sight. Inside the retina is the vitreous body, a transparent spheroidal mass of a gelatinous consistency, which is surrounded and retained in position by a thin, structureless mem- brane, called the hyaloid membrance (s), lying immediately in contact with the internal surface of the retina. The lens (4) is placed in front of the vitreous body, in the central axis of the eye- ball, enveloped in its capsule, which is continuous with the hyaloid membrane. Just at the edge of the lens, the hyaloid membrane divides into two laminae, which separate from each other, leaving between them a triangular canal, the canal of Petit, which can be seen in the above figure. In front of the lens is the iris (7), a nearly 494 THE SPECIAL SENSES, vertical muscular curtain, formed of radiating and concentric fibres, pierced at its centre with a circular opening, the pupil, through which the light is admitted, and covered on its posterior surface with a continuation of the choroidal pigment, which excludes the passage of any other rays than those which pass through the pupil. At the same time, the whole globe is inclosed and protected by a thick, fibrous, laminated tunic, which in its posterior and middle portions is opaque, forming the sclerotic (i), and in its anterior.por- tion is transparent, forming the cornea (s). The muscles of the eye- ball are attached to the external surface of the sclerotic in such a way that the cornea may be readily turned in various directions ; while the eyelids, which may be opened and closed at will, protect the eye from injury, and, with the aid of the lachrymal secretion, keep its anterior surfaces moist, and preserve the transparency of the cornea. The organ of vision is supplied with nerves of ordinary sensi- bility by the ophthalmic branch of the fifth pair. The filaments of this nerve which terminate about the eye are distributed mostly to the conjunctiva, lachrymal gland, and skin of the eyelids ; while a very few of them run forward in company with the ciliary nerves proper, and are distributed to the ciliary circle and iris. All these parts, therefore, but more particularly the conjunctiva and skiii;.§f the eyelids, possess ordinary sensibility, which appears to be totally wanting in the deeper parts of the eye. The ophthalmic ganglion gives off the ciliary nerves, which, are distributed to the iris and ciliary muscle. Finally, the muscles moving the eyeball and eye- lids are supplied with motor nerves from the third, fourth, sixth and seventh pairs. Of all the properties and functions belonging to the diflferent structures of the eyeball, the most peculiar and characteristic is the special sensibility of the retina. This sensibility is such that the retina appreciates both the intensity, and the quality of the light- that is to say, its color and the different shades which this color may present. On account of the form, also, in which thc'retina is constructed, viz., that of a spheroidal membranous bag, with an opening in front, it becomes capable of appreciating the direction from which the rays of light have come, and, of course, the situation of the luminous body and of its different parts. For the rays which enter through the pupil from below can reach the retifia only at its upper part, while those which come in from above, can reach it only at its lower part; so that in both instances the rays strike the SIGHT. 495 sensitive surface perpendicularly, and thus convey the impression of their direction from above or below. But beside the sensibility of the retina, the perfection and value of the sense of sight depend very much on the arrangement of the accessory organs, the most important of which is the crystalline lens. The fwmtion of the crystalline lens is to produce distinct perception offm-m and outline. For if the eye consisted merely of a sensitive retina, covered with transparent integument, though the impressions of light would be received by such a retina they could not give any idea of the form of particular objects, but could only produce the sensation of a confused luminosity. This condition is illus- trated in Fig. 156; where the arrow, a, h, represents the luminous object, and the vertical dotted line, at the right of the diagram, represents the retina. Rays, of course, will diverge from every point of the object in every direction, and will thus reach every part of the retina. The different parts of the retina, consequently, 1, 2, 8, 4, will each receive rays coming both from the point of the arrow, a, and from its butt, b. Thei'e will therefore be no distinc- tion, upon the retina, between the different parts of the object, and no Fig. 156. .. Fig. 157. definite perception of its outline. But if, between the object and the retina, there be inserted a double convex refracting lens, with the proper curvatures and density, as in Fig. 157, the effect will be dif- ferent. For then all the rays emanating from a will be concentrated at X, and all those emanating from b will be concentrated at y. Thus the retina will receive the impression of the point of the arrow separate from that of the butt ; and all parts of the object, in like manner, will be distinctly and accurately perceived. This convergence of the rays of light is accomplished to a certain extent by the other transparent and refracting parts of the eyeball ; but the lens is the most important of all in this respect, owing to its superior density and the double convexity of its figure. The distinctness of vision, therefore, depends upon the action of the 496 THE SPECIAL SENSES. lens in converging all the rays of light, emanating from a given point, to an accurate focus, at the surface of the retina. To accomplisli this, the density of the lens, the curvature of its surfaces, and its distance from the retina, must all b6 accurately adapted to each other. For if the lens be too convex, and its refractive power con- sequently too great, the rays will be converged to a focus too soon, and will not reach the retina until aftef they have crossed each other and become partially dispersed, as in Fig. 158. The visual impression, therefore, coming from any particular point in the object is not concentrated and distinct, but diffused and dim, from being dispersed more or less over the retina, and interfering with the impressions coming from other parts. This is the condition which is present in myopia, or near-sightedness. On the other hand. Fig. 158.. Fig. 159. Miopia. Pkebbtopia. if the lens be too flat, and its convergent power too feeble, as in Fig. 159, the rays will fail to come together at all, and will strike the retina separately, producing a confused image, as before. %ia is the defect which exists in presbyopia, or long-sightedness. In both cases, the immediate cause of the confusion of sight is the same, viz., the rays coming from the same point of the object striking the retina at different points ; but in the first instance, this is because the rays have actually converged, and then crossed ; in the second, it is because they have only approached each other, but have never converged to a focus. Another important particular in regard to the action of the lens • is the accommodation of the eye to distinct vision at different distances. It is evident that the same arrangement of the refractive parts, in the eye, will not produce distinct vision when the distance of the object from the eye is changed. If this arrangement be such that the object is seen distinctly at a certain. distance, as in Fig. 160, and the object be then removed to a remoter point, as in Fig. 161, the image wiU become confused ; for the, rays will then be con- SIGHT. 497 verged to a focus at a point in front of the retina ; because, being less divergent, when they strike the lens, the same amount of re- fraction will bring them together sooner than before. On the other hand, if the object be moved to a point nearer the eye, the rays, becoming more divergent as they strike the lens, will be converged less rapidly to a focus, and vision will again become indistinct. This may easily be seen by the aid of a very simple experiment. If two needles be placed upright, at different distances from the eye, one for example at eight and the other at eighteen inches, but nearly in the same linear range, and if then, closing one eye, we look at them alternately, we shall find that we cannot see both dis- tinctly at the same time. For when we look at the one near- Fig. 160. Fig. Itil, est the eye, so as to perceive its form distinctly, the image of the more remote one becomes confused ; and when we see the more re- mote object in perfection, that which is nearer loses its sharpness of outline. This shows, in the first place, that the same condition of the eye will not allow us to see two objects at different distances with distinctness at the same time ; and secondly that, on looking from one to the other, there is a change of some kind in the focus of the eye, by which it is adapted to difierent distances. Indeed we are conscious of a certain effort at the time when the point of vision is transferred from one object to the other, by which the eye is adapted to the new distance ; and this alteration is not quite instan- taneous, but requires a certain interval of time for its completion. This accommodation of the eye to different distances is un- doubtedly efiected by an antero-posterior movement of the lens within the eyeball. It will at once be perceived, on referring to Fig. 161, that if the lens were moved a little backward toward the 32 498 THE SPECIAL SENSES. retina, at the same time that the object is removed to a greater dis- tance from the eye, the focus of the convergent rays would still 6,11 upon the retina, and the image would still be distinct. In the op- posite case, where the object is brought nearer the eye, a similar movement of the lens forward would again secure perfect vision. Thus, when we look at near objects, the lens moves forward toward the pupil ; when we look at remote objects, it moves back- ward toward the retina. This movement of the lens is apparently accomplished by the action of the ciliary muscle. This muscle (Fig, 155, s) arises, in front, from the conjunction of the sclerotic and the cornea, and run- ning backward and outward, is inserted into the anterior part of the choroid, about the situation at which, the hyaloid membrane passes off, to become the suspeusory ligament of the lens. As already mentioned, this muscle is supplied with nervous filaments from the ophthalmic ganglion. Its action is to draw the lens for- ward, by means of its attachment to the hyaloid' membrane and choroid coat ; and, in the human subject, the retreat or retrogres- sion of the lens toward the retina, after the ciliary muscle is relaxed, seems to be due to the islastic resiliency of the remaining tissues of the eyeball. But in order to allow of such a backward and forward movement of the lens, since the liquids of the eyeball are incompressible, there must be a corresponding displacement of other parts, both before and behind. This is undoubtedly provided for by the vascularity of the choroid coat. This membrane is supplied with an exceedingly abundant vascular plexus over its whole posterior portion ; and in front it is thrown into a circle of prominent converging folds, or processes, the ciliary processes, which are nothing more than erectile congeries of bloodvessels, covered with the pigment of the choroid. A portion of the ciliary processes projects in front of the lens, and their vascular network is continued over a great part of the pos- terior surface of the iris. Thus there is, both behind and in front of the lens, an erectile system of bloodvessels ; and as these blood- vessels become alternately empty or turgid, they will allow of the displacement of the lens in an anterior or posterior direction. Accordingly there is a certain accocamodation of the eye neces- sary to the distinct sight of objects at different distances. But the rfinge of this accommodation is limited, and the same eye cannot he made to see distinctly at all distances. For all ordinary eyes, the accommodation fails, and vision becomes imperfect, when the object . SIGHT, 499 is placed at less than six inches distance from the eye. But from that point outward, the eye can adapt itself to any distance at which light is perceptible, even to the immeasurable distances of the fixed stars. A much greater accommodating power, however, is re- quired for near distances than for remote, since the difference in divergence between rays, entering the pupil from a distance of one inch and from that of six inches, is greater than the difference be- tween six inches and a yard, or even distances which are immea- surably remote. Accordingly, near-sighted persons can see objects distinctly when placed very near the eye; since, as their lens con- verges the rays of light more powerfully than usual, they can be brought to a focus upon the retina, even when excessively diverg- ent at the time they enter the eye. But distant objects become indistinct, since, however far backward the lens is moved, the rays are still brought to a focus and cross each other, before reaching the retina, as in Fig. 161. Near-sighted persons, therefore, have a limited range of accommodation, like all others, only it is confined within short distances, owing to the excessive refracting power of the lens. On the other hand, long-sighted persons can see remote objects without trouble, since a very little movement of the lens will be sufiicient to adapt it for long distances ; but within short distances, the divergence of the rays becomes too great, and they cannot be brought to a focus. Circle of Vision. — Since the opening of the pupil will admit rays of light coming from various directions, there is in front of the eye a circle, or space, within which luminous objects are perceived, and beyopd which nothing can be seen, because the rays, coming from the side or from behind, cannot enter the pupil. This space, within which external objects can. be perceived, is called the " circle of vision." But, for short distances, there is only a single point, in the centre of the circle of vision, at which objects can be seen distinctly. Thus, if we place ourselves in front of a row of vertical stakes or palisades, we can see those directly in front of the eye with perfect distinctness, but those at a little distance on each side are only per- ceived in a confused and uncertain manner. On looking at the middle of a printed page, in the direct range of vision, we see the distinct outlines of the letters ; while at successive distances from this point, the eye remaining fixed, we can distinguish first only the separate letters with confused outlines, then only the words, and lastly only the lines and spaces. 500 a HE SPECIAL SENSES. This is because rays of light coming into the eye very obliquely, in a lateral or vertical direction, are not brought to their proper focus. Thus, in Fig. 162, the rays diverging from the point a, directly in front of the eye, fall upon the lens in such a way that they are all brought together at x, at the surface of the retina ; but those coming from h fall upon the lens so obliquely that, for rays having an equal divergence with those coming from a, there is more difference in their angles of incidence, and of course more difference in the amount of their refraction. They are consequently brought together more rapidly, and on reaching the retina are dispersed over the space y, z. Fig. Iii2. The perfection of the eye, as a visual apparatus, is very much increased by the action of the iris. This organ, as we have already mentioned, is a nearly vertical muscular curtain, placed in front of the lens, attached by its external margin to the junction, of the cornea and sclerotic^ and pierced about its centre by the circular opening of the pupil. It consists, according to most anatomis'ts,;of two sets of muscular fibres — viz., the circular and the radiating. The circular fibres, which are much the most abundant, are arranged in concentric lines about, the inner edge of the irjs, near the pupil; the others are said to radiate in a scattered manner, from its central parts to its outer margin. The action of these two sets of fibres is to contract and enlarge the orifice of the pupil. The circular fibres, in contracting, draw together the edges of the pupil, and so diminish its opening; and when these are relaxed, the radiating fibres come into play, and by drawing apart the edges of 'the orifice, enlarge SIGHT. 501 the pupillary opening. The action of the ciroular fibres, at tlie same time, is much the most marked and important of the two. For when the whole muscular apparatus of the eye is paralyzed by the action of belladonna, or by the division of the third pair of nerves, or in the general relaxation of the muscular system at the moment of death, the pupil is invariably dilated, probably by the passive elasticity of its tissues. During life, however, these different conditions of the pupil cor- respond with the different degrees of light to which the eye is ex- posed. In a strong light, the pupil contracts and shuts out the superfluous rays; in a feeble light, it dilates, in order to collect into the eye all the light which can be received from the object. This contractile and expansive movement of the pupil is a reflex action. It is not produced by the direct impression of the light upon the iris itself, but upon the retina; since, if the retina be affected with complete amaurosis, or if the light be entirely shut out from it by an opacity of the lens, no such effect is produced, though the iris itself be exposed to the direct glare of day. From the retina the impression is transmitted, through. the optic nerve, to the optic tubercles and the brain, 'thence reflected outward by the ooulo- motorius nerve to the ophthalmic ganglion, and so through the ciliary nerves to the iris. The pupil is subject, however, to various other nervous influences beside the impressions of light received by the retina. Thus in poisoning by opium, it is contracted ; in coma from compression of the brain, it is dilated ; in natural sleep it is contracted, and the eye- ball rolled upward and inward. In various mental conditions, the pupil is also enlarged or diminished, and thus modifies the expres- sion of the eye; and in viewing remote objects, it is generally enlarged, while, in looking at near objects, it is comparatively con- tracted. But still, the most constant and important function be- longing to the iris is the admission or exclusion of the rays, accord- ing to the intensity of the light. Our impressions of distance and solidity, in viewing external objects, are produced mainly Tjy the combined action of the two eyes. For, as the eyes are seated a certain distance apart from each other in the head, when they are both directed toward the same object, their axes meet at the point of sight, and form a certain angle with each other ; and this angle varies with the distance of the object^ Thus, when the object is within a short distance, the axes of the two eyes will necessarily be very convergent, and the angl^ which 502 THE SPECIAL SENSES. they form with each other a large one ; but for remote objects, the visual axes will become more nearly parallel, and their angle con- sequently smaller. It is on this account that we can always dis- tinguish whether any person at a short distance is looking at m, or at some other olg'ect in our direction; since we instinctively appreciate, from the appearance of the eyes, whether their visual axes meet at the level of our own face. In looking at a landscape, accordingly, we do not see the whole of it distinctly at the same moment, but only those parts to which out attention is immediately directed. This is because, in the first place, the focus of distinct vision varies, in each eye, for different distances, as we have seen in a former paragraph, and secondly, because both eyes can only be directed together, at one time, to objects at a certain distance. Thus, when we see the foreground or the middle ground distinctly, the distance is vague and uncer- tain, and when we direct our eyes more particularly to the horizon, objects in the foreground become indistinct. In this way we ap- preciate the difterence in distance between the various portions of the landscape, as a whole. In the case of particular objects, we are assisted also by the alteration in their individual characters ; for distance produces a diminution, both in apparent size and in in- tensity of color. The combined action of the two eyes is also very valuable, for near objects, in giving us an idea of solidity or projection^} For within a certain distance, the visual axes when directed together Fig. ltJ3 As SEEK BT THR LEFT Ete. As SEEN BY THE RIGHT EVE. at a solid object, are so convergent that -the two- eyes do not receive the same image. As in Figs. 163 and 164, which represent a skull SIGHT. 503 as seen by the two eyes, when placed exactly in front of the ob- server at the distance of eighteen , inches or two feet, the right eye will see the object partly on one side, and the left eye partly on the other. And by the union or combination of these two images by the visual organs, the impression of solidity is produced. By the employment of double pictures, so drawn as to represent the appearances presented to the two eyes by the same object, and so arranged that each shall be seen only by the corresponding eye, a deceptive resemblance may be produced to the actual appearance of. solid objects. This is accomplished in the contrivance known as the Stereoscope. Thus, if two pictures similar to those in Figs. 163 and 164 be so placed that one shall be seen only with the right eye and the other only with the left, the combination of the two figures will take place as if they came from the real object, and all the natural projections will come out in relief. But this effect is produced only in the case of objects situated within a moderately short distance. For very remote objects, we lose the impression of solidity, since the difference in the images on the two eyes becomes so slight as to be inappreciable, and we see only a plane expanse of surface, with sharp outlines and various shades of color, but no actual projections or depressions. The sensibility of the retina is such that it cannot distinguish luminous points which are received upon its surface at a very minute distance from each other. In this particular, the sensibility of the retina resembles that of the skin, since we have already found that the integument cannot distinguish the impressions made by the points of two needles placed a very short distance apart. The delicacy of this discriminating power, in the retina, is immeasurably superior to that of the skin; and yet it has its limits, even in the nervous expansion of the eye. For if we look at an object which is excessively minute, or which is so remote that its apparent size is very much diminished, we lose the power of distinguishing its difierent parts, and can no longer perceive its real outline. This is a very different condition from that in which the confusion of vision arises from defect of focusing in the eye, as, for example, in long or short-sightedness, or where the object is placed too near the eye or too much on one side. For when the difSculty depends simply on its minute size or its remote- ness, the rays coming from the top of the object and those coming from the bottom, are all brought to their proper focus at distinct points on the retina — only these points are too near each other fw the 504r lUK SPECIAL SEXSES. retina to distinguish them apart. Consequently we can no longer appreciate the form of the object. For the same reason, when we mix together minute grains of a diiferent hue, we produce an intermediate color. If yellow and blue be mingled in this way, we no longer perceive the separate blue and yellow grains, but only a uniform tinge of green ; and white and black granules, mixed together, produce, at a short dis- tance, the appearance of a continuous shade of gray. Impressions, once produced upon the retina, remain for a short time afterward. Usually these impressions are so evanescent after the removal of their immediate cause, and are so soon followed by others which are more vivid, that we do not notice their existence. They may very readily be demonstrated, however, by swinging rapidly in a circle before the eyes, in a dark room, a stick lighted at one end. As soon as the motion has attained a certain degree of velocity, the impression produced on the retina, when the lighted end of the stick arrives at any particular spot, remains until it has completed its revolution and has again reached the same point; so that the effect thus produced upon the eye is that of a continu- ous circle of light. The same fact has been illustrated by the optical contrivance, known as the Thaumatrope, in which successive pictures of similar figures in different positions are made to revolve rapidly before the eye, and thus to produce the apparent eflect of a single figure in rapid motion ; — since the eye fails to percei^ the intervals between the different pictures. The sense of vision, therefore, through the impressions of lighf^ gives us ideas of form, size, color, position, distance, and movement. But these ideas may also be excited by impressions derived from an internal source, as well as those produced by rays coming from without. And it is one of the most striking peculiarities of the sense of sight that these ideal or internal impressions which are excited in it by various causes, are much more vivid and powerful than those of any other of the senses. Thus, in a dream, we often see external objects, with all their visible peculiarities of liglit, color form, &c., nearly or quite as distinctly as when we are awake ; but the imaginary impressions of sound, in this condition, are always comparatively faint, and those of taste, smell, and touch, almost entirely imperceptible. Even in a reverie, in the waking condi- tion, when the absorption of the mind in its own thoughts is com- plete, and we are withdrawn altogether from outward influences, we see objects which have no present existence as if they were HEARING. 505 actually before us. It is this sense also whioli becomes most easily and thoroughly excited in certain nervous disorders ; as, for exam- ple, in delirium tremens, where the patient often sees passing before his eyes extensive and magnificent landscapes, crowds of human faces and figures, and series of towns and cities, which seem to be depicted upon the imagination with a force and distinctness, much superior to that of other delirious impressions. Since the sense of sight, therefore, depends less directly than the other senses upon the actual contact of material objects, it is also more easily thrown into activity when withdrawn from their influence. Hearing. — The sense of hearing depends upon the vibratioos excited in the atmosphere by sonorous bodies, which are themselves first thrown into vibration by various causes, and which then com- municate similar undulations to the surrounding air. These sono- rous vibrations are of such a character that they cannot be directly appreciated by ordinary sensibility, but the result of numerous and well-directed physical experiments on this subject leaves no doubt whatever of their existence ; and when such vibrations are commu- nicated to the auditory apparatus, they produce in it the sensation of sound. In the case of the aquatic animals, which pass their entire exist- ence beneath the surface of the water, the water itself, which is capaWe of vibrating in the same- way, communicates the sonorous impressions to the organ of hearing; but in terrestrial animals, and particularly in man, it is the atmosphere which always serves as the medium of transmission. The auditory apparatus, in man and in the quadrupeds, consists, first, of a somewhat expanded and trumpet-shaped mouth, or ex- ternal ear, destined to receive and collect the sonorous impulses coming from various quarters. This external ear is constructed of a cartilaginous framework, covered with integument, loosely attached to the bones of the head, and more or less movable by means of various muscles, which by their contraction turn its expanded orifice in different directions. In man, the movements of the external ear are almost always inappreciable, though the mus- cles may be easily demonstrated ; but in many of the loWer animals these movements are exceedingly varied and extensive, and play a very important part in the working of the auditory apparatus. At the bottom of the external ear, its orifice is prolonged into a tube or canSil, the external auditory meatus, partly cartilaginous and 606 THE SPECIAL SENSES. partly bony, wMch penetrates the lateral part of the temporal bone in a nearly horizontal and transverse direction. In the human subject, this canal is a little over one inch in length, and is lined by a continuation of the external integument. The integument toward its outer portion is beset with small hairs, and provided with ceruminous glands which supply a secretion of a waxy or resinous consistency. By these means the passage is protected from the accidental ingress of various foreign bodies. Secondly, at the bottom of the external meatus the auditory pas- sage is closed by a thin fibrous membrane, stretched across its. cavity, called the membrana tympani. Upon this membrane are received the sonorous vibrations which have been collected by the external ear and conducted inward by the external auditory meatus. Behind the membrana tympani is the cavity of the middle ear, or the cavity of the tympanum. This cavity communicates posteriorly with the mastoid cells, and anteriorly with the pharynx, by a narrow passage, lined with ciliated epithelium, and running downward, forward and inward, called the Eustachian tube. A chain of small bones, the malleus, incus, and stapes, is stretched across the cavity of the tympanum, and forms a communication between the membrana tympani on the outside, and the membrane closing the. foramen ovale in the petrous portion of the temporal bone. All the vibra- tions, accordingly, which are received by the membrana tympani, are transmitted by the chain of bones to the membrane of the foramen ovale. The tension of the membranes is regulated by two small muscles, the tensor tympani and stapedius muscles, which arise from the bony parts in the neighborhood, and are inserted respect- ively into the neck of the malleus and the head of the stapes, and which draw these bones forward and backward upon their articu- lations. Thirdly, behind the membrane of the foramen ovale lies the labyrinth, or internal ear. This consists of a complicated, cavity, excavated in the petrous portion of the temporal bone, and com- prising an ovoid central portion, the vestibule, a double spiral canal, the cochlea, and three semicircular canals, all communicating by means of the common vestibule. All parts of this cavity contain a watery fluid termed the perilymph. The vestibule and semi- circular canals also contain closed membranous sacs, suspended in the fluid .of the perilymph, which reproduce exactly the form of the bony cavities themselves, and communicate with each other in a similar way. These sacs are filled with another iv-atery fluid. HEARING. 507 the endolymph ; and the terminal filaments of the auditory nerve are distributed upon the membranous sac of the vestibule and upon the ampullffi, or membranous dilatations, at the commencement of the three semicircular canals. The remaining portion of the audi- tory nerve is distributed upon the septum' between the two spiral canals of the cochlea. Thus, the. essential or fundamental portion of the auditory appa- ratus is evidently the internal ear, a cavity, partly membranous and partly bony, in which is distributed a nerve of special sense, the auditory nerve, capable of appreciating sonorous impressions. The accessory parts, on the other hand, are the chain of bones and the membrane of the tympanum, which communicate the sonorous vibrations directly to the internal ear ; and the meatus and external ear, which collect them from the atmosphere. The reception of Fig. 165. Human Auditort ApPABATrs, showiug external auditory meatus, tympanuni, aad laby- ri^h. sonorous impulses is therefore accomplished in a very indirect way. For the sonorous body first communicates its vibrations to the atmosphere. By the atmosphere these vibrations are communicated tci the membrana tympani. From the membrana tympani, they are transmitted, through the chain of bones, to the membrane of the foramen ovale ; thence to the perilymph, or fluid, of the labyrinthic cavity, and from the perilymph to the membranous parts of the labyrinth and the nerves which are distributed upon them. The arrangement of the different parts composing the tympanum is of the greatest importance for the perfect enjoyment of the sense 508 THE SPKCIAL SENSES. of hearing. For the air on the two sides of the membrane of, the tympanum should be in the same condition of elasticity in order to allow of the proper vibration of the membrane; and this equilibrium would be liable to disturbance if the air within the tympanum were completely confined, while that outside is subjected to variations of barometric pressure. By means of the Eustachian tube, how- ever, a communication is established between the cavity of the tympanum and the exterior, and the free vibration of the membrane is thus secured. The exact tension of the membrana tympani itself is also provided for, as we have already observed, by the action of the two muscles inserted into the malleus and the stapes. By the contraction of the internal muscle of the malleus, or tensor tympani, the membrane of the tympanum is drawn inward and rendered more tense than usual. The action of the stapedius muscle is by some thought to relax the membrana tympani, by others to assist in the tension both of this membrane and that of the foramen ovale, to which the stapes is attached. But there is no doubt that both these mus- cles, by their, combined or alternate action, can regulate the tension of the tympanic membrane, to an extraordinary degree of nicety, and thus increase the ease and delicacy with which various sounds are distinguished. For if the membrane be so put upon the stretch that its fundamental note shall be the same with that of the sound which is to be heard, it will vibrate more readily in consonance with the undulations of the atmosphere, and the sound will be more distinctly heard. On the contrary, if the membrane be too highly stretched, very grave sounds may not be heard at all, until its tension is diminished to the requisite degree. -. Contrary to what is sometimes asserted, the communication of sonorous impulses to the internal ear is accomplished altogether by means of the tympanum and chain of bones. It has been thought tHfet sounds were transmitted, in many instances, directly to the internal ear by the medium of the cranial bones. This was inferred from such facts as the following. If a tuning-fork, in vibration, be taken between the teeth, its sound will appear very much louder than if it were simply held near the external ear ; and if, while it js so held, one of the ears be closed, the sound will appear very much louder on that side than on the other. The sound will also be heard if the tuning-fork be applied to the upper part of the cranium or the mastoid process, with a similar increase of resonance on closing the ears. Finally our own voices are heard, though the ears b»" HEARING. 509 both closed, and the sound is much louder with the ears closed than open. These are the facts which have led to the belief that, in such instances, the sound was communicated directly through the bones of the head, vibrating in consonance with the sounding body. But a little examination will show that such is not the case. When we hold the end of a vibrating tuning-fork between the teeth, we no longer hear the sound in the vibrating extremity of the instrument or its neighborhood, but in the mouth and the nasal fossce. It is the vibration of the air in these passages which produces the sound ; and this vibration is communicated to the cavity of the tympanum through the Eustachian tube. The apparent increase of sound, also, on closing the ears, which could not be explained on the supposition that it was conducted directly through the bones of the cranium, is due to the same cause. For it can easily be seen, on trying the experiment, either with a tuning-fork held between the teeth or simply with our own voices, that this apparent increase of sound takes place only when the ears are closed by genth pressure. If the pressure be excessive, so that the integument is forced inward into the meatus and the air in the meatus subjected to undue compres- sion, the sound no longer appears louder in the corresponding ear, and may even be lost altogether. The apparent increase of sound, therefore, in such cases, when the ear is gently closed, is due to the fact that the meatus is thus converted into a reverberatory cavity, by which the vibrations of the tympanum are increased in intensity. But if the air in the meatus be too much compressed by forcible closure, the vibrations of the tympanum are then interfered with and the sound is dimi- nished or destroyed. In all cases, then, it is the sonorous vibrations of the air which produce the sound, and these vibrations are received invariably by the membrape of the tympanum, and thence transmitted to the internal ear by the chain of bones. The cranial bones are incapable of communicating these vibrations to the labyrinth and its contents, except very faintly and imperfectly. For common experience shows that even the loudest and sharpest sounds, coming from without, are almost entirely lost on closing the external ears ; and our own respiratory and cardiac sounds, which are so easily heard as soon as the chest is connected with the ear by a flexible stethoscope, are entirely inaudible to us in the usual condition. The exact function of the different parts of the internal ear is 510 THE SPECIAL SENSES. not well understood. It has been thought to be the office of the semicircular canals to determine the direction from which the sono- rous impulses are propagated. This opinion was based upon the curious fact that these canals, always three in number, are placed in such positions as to correspond with the three different directions of vertical height, lateral extension, and longitudinal extension ; for one of them is nearly vertical and transverse, another vertical and longitudinal, and the third horizontal in position. The sono- rous impulses, therefore, coming in either of these directions, would be received by only one of the semicircular canals (by direct con- duction through the bones of the head) perpendicularly to its own plane; and an intermediate direction, it was thought, might be appreciated by the combined effect of the impulse upon two adja- cent canals. ' Enough has already been said, however, in regard to the com- munication of sound directly through the bones of the head to the internal ear, to show that this cannot be the way in which the diree- tion of sound is ascertained. Indeed, when we hear any loud and well-marked sound coming from a particular region, such as the music of a military band or the whistle of a locomotive, we have only to close the external ears to lose our perception both of the sound and its direction. The direction of sonorous impressions is appre- ciated in a different way. In the first place, we feel that the sound comes from one side or the other, by its making a more distinct impression on one ear than the opposite; and by inclining the head slightly in various directions, we easily ascertain whether the sound becomes more or less acute, and so judge of its actual source. Many of the lower animals, whose ears are very large and movable, use this method to great extent. A horse, for example, when upon the road, often keeps his ears in constant moixon, feeling, as it were, in the distance, for the origin of the various sounds which excite his attention. .Beside the above, we are further assisted in our judgment of the direction of sounds by our previous knowledge of the localities, the direction of the wind, and the manner in which the sound is reflected by surrounding objects. When these sources of informa- tion fail us, we are often at a loss. It is notoriously difficult, for example, to judge of the place of the chirping of a cricket in a perfectly closed room, or of the direction of a bell heard on the water in a thick fog. The sense of hearing has a much closer analogy with ordinary ON THE SENSES IN GENERAL. 511 i sensibility than that of sight. Thus, in the first place, hearing is accomplished by the direct intervention and contact of a materia) body — the atmosphere ; for sonorous impulses cannot be, produced in a vacuum, and we hear no sound from a bell rung under an exhausted receiver. Secondly, the nature of the impressions pro- duced by sound is such that we can often describe them by the same terms which are applied to ordinary sensations. Thus, we speak of sounds, as sharp and dull, piercing, smooth, or rough ; and we feel the impulse of a sudden and violent explosive sound, like that of a blow upon the tympanum. By {his sense, therefore, we distinguish the quality, intensity, pitch, duration, and direction of sonorous impulses. The delicacy with -which these distinctions are appreciated varies considerably in different individuals ; and in different kinds of animals there is reason tO believe that the diversity is much greater, some of them being almost insensible to sounds which are readily perceived by others. In man, the number and variety of tones which can usually be discriminated is very great ; and this sense, accordingly, in the complication and finish of its apparatus, and the perfection and deli- cacy of its action, must be regarded as second only to that of vision. On the Senses in Geneeal. — There are several facts connected with the operation of the ^nses, both general and special, which are common to all of them, and which still remain to be considered. In the first place, an impression of any kind, made upon a sensi- tive organ, remains for a time after the removal of its eaxiting cause. We have already noticed this in regard to the senses of taste, smell, and sight, but it is equally true of the hearing and the touch. Thus, if the skin be touched with a piece of ice, the acute sensa- tion remains for a few seconds, whether the ice be removed or not. For the higher order of the special senses, the time during which this secondary impression remains is a shorter one. In the case of hearing, however, it has been measured with a tolerable approach to accuracy ; for it has been found that, if the sonorous undulations follow each other with a greater rapidity than sixteen times per second, they become fused together into a continuous sound, pro- ducing upon the ear the impression of a musical note. The varying pitch of the note depends upon the rapidity with which the vibra- tions succeed each other. When the succession of yibrations ia very rapid, a high note is the result, and when comparatively slow, a low note is produced ; but when the number of impulses falls 512 THK SPECIAL SKNSE3. k below sixteen per second, we then begin to perceive the distinct vibrations, and so lose the impression of a continuous note. All the senses, in the second place, become accustomed to a con- tinued impression, so that they no longer perceive its existence. Thus, if a perfectly uniform pressure be exerted upon any part of the body, the compressing substance after a time fails to excite any sensation in the skin, and we remain unconscious of its existence. In order to attract our notice, it is then necessary to increase or diminish the pressure ; while, so long as this remains uniform, no effect is perceived. But if, after the skin has thus become accus- tomed to its presence, the foreign body be suddenly remo^^d, our attention is then immediately excited, and we notice the absence of an impression, in the same way as if it were a positive sensation. "We all know how rapidly we become habituated to odors, whether agreeable or disagreeable in their nature, in the confined air of a close apartment; although, on first entering from without our attention may have been attracted by them in a very decided manner. A continuous and uniform sound, also, like the steady rumbling of carriages, or the monotonous hissing of boiling water, becomes after a time inaudible to us; but as soon as the sound ceases, we notice the alteration, and our attention is at once excited. The senses, accordingly, receive their stimulus more from the varia- tions and contrasts of external impressions, than from these impres- sions themselves. Another important particular, in regard to the senses, is their capacity for education. The proofs of this are too common and too apparent to need more than a simple allusion. The touch may be so trained that the blind may read words and sentences by its aid, in raised letters, where an ordinary observer would hardly detect anything more than a barely distinguishable inequality of surface. The educated eye of the artist, or the naturalist, will distinguish variations of color, size, and outline, altogether inappreciable to ordinary vision ; and the senses of taste and stoell, in those who are in thfe habit of examining wines and perfumes, acquire a similar superiority of discriminating power. In these instances, however, it is not probable that the organ of sense itself becomes any more perfect in organization, or more susceptible to sensitive impressions. The increased functional power, developed by cultivation, depends rather upon the greater delicacy of the perceptive and discriminative faculties. It is a mental and not a physical superiority which gives the painter or the ON THE SKNSiJS IN GENERAL. 513 naturalist a greater power of distinguishing colors and outlines, and which enables the physician to detect nice variations of quality in the sounds of the heart or the respiratory murmur of the lungs. The impressions of external objects, therefore, in order to produce their complete effect, must first be received by a sensitive appa- ratus, which is perfect in organization and functional activity; and, secondly, these impressions must be subjected to the action of an intelligent perception, by which their nature, source and rela- tions may be fully appreciated. 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, taste, 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 by the same portion of the nerv- ous system. The introduction of food into the mouth, and its 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." 33 514 SYSTEM OF THE GR^AT SYMPATHETIC. CHAPTER VII. SYSTEM OP 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 reiilforced 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, &o. The first sympathetic ganglion in the head is ihQ 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 fifth 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 spheno-palatirie 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 fifth 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 suhmaxillary, situated upon the submaxillary gland. It communicates with the superipr cervical ganglion of the sympathetic by filaments which accompany the facial and external carotid arteries. It derives its sensitive filaments from the lingual braach of the fifth pair, and its SYSTEM OJT THE GREAT SYMPATHETIC. 515 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 Kg- 166. side of the third division of the fifth pair. 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 division of the fifth 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 three ganglia, the su- perior, 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 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. Course and distribatiou of the Great Sthpa- THBTIC. 516 SYSTEM OF THE GREAT SYMPATHETIC. In the chest, the ganglia of the sympathetic nerve are situated on 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 cceliac 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 distri- buted to the stomach, small and large intestine, spleen, pancreas, liver, kidneys, supra-renal capsules, and internal organs of gene- ration; Beside the above ganglia there are in the abdomen four other pairsy situated in front of the luihbar 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 of this system, and to appreciate the disturbances of sensation and motion conse- quent upon local lesions or irritations. The phenomena, further- more, which result from experiments upon this part of the nervous apparatus, are promptly produced, are well-marked in character. SYSTEM OF THE GBEAT SYMPATHETIC. 517 and are, as a general rule, readily understood by the experimenter. On the other hand, the 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 disturbance 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 unsatis- factory. Furthermore, the connections of the sympathetic ganglia with each other and with the cerebro-spinal axis are so numerous and so scattered, that these ganglia cannot be completely isolated without resorting to an operation still more mutilating and injuri- ous in its character. And finally, the sensible phenomena which are obtained by experimenting on the great sympathetic are, in the majority of cases, slow in making their appearance, and not particularly striking or pharacteristic 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 much less acutely, and only after somewhat prolonged application 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 sym- pathetic; as the liver, intestine, kidneys, &o. These organs are insensible, or nearly so, to ordinary impressions. "VVe 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- iSammation. There is the same peculiar character in the action of the motor 518 SYSTEM OF THE GREAT SYMPATHETIC. nerves belonging to the sympathetic system. If the facial or hypo- glossal, or the anterior root of a spinal nerve be irritated, the con- vulsive movement which follows is instantaneous, violent, and only momentary 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 day. 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, wbich 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 fifth pair, and a motor root from the oculo-motorius. The reflex action by which the pupil contracts under a strong light foiling 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 SYSTEM OS" THE GEEAT SYMPATHETIC. 519 we pass suddenly from a brilliantly lighted apartment into a dark 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 after- ward continues to diminish in size during several seconds, until the proper equilibrium is fairly established. Furthermore, if we pass suddenly from a dark room into the bright sunshine, we are imme- diately conscious of a painful sensation in the eye, which lasts for a considerable time ; and which results from .the inability of the 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 ooulo-motorius nerve. For it has been found that if the oculo- motorius 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 reaches 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 qua,ntity of stimu- lus admitted to the organ, and the mode in which it is received. The superficial set of these muscles is animated by branches of the facial nerve ; the deep-seated or internal set, by filaments from a sympathetic ganglion. T]hus, 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 520 SYSTEM OP THE GREAT SYMPATHETIC. facial nerves, and are for the most part voluntary in their action. The iris, on the other hand, is a more deeply-seated muscular curtain, which regulates the quantity of light admitted through the pupil. There is also the ciliary muscle, which regulates the position of the crystalline lens, and secures a correct focusing of the light, at different distances. Both these 'muscles are supplied, as we have seen, by filaments from the ophthalmic ganglion, and their move- ments are 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 tlie muscles of the middle ear. In order to understand their action, we must recollect that sounds are trans- mitted 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. SYSTEM OF THE GREAT SYMPATHETIC. 521 The membrane of the tympanum, accordingly, -which is an elastic sheet, stretched across the passage to the internal ear, may be made more or less sensitive to sonorous impressions by varying its con- dition of tension or relaxation. This condition is regulated, as we have already seen, by the combined action of the two muscles of the middle ear, viz., the tensor tympani and the stapedius. The first named muscle, the action of which is perfectly well understood, is supplied with nervous filaments from the otic ganglion of the sympathetic. By its contraction, the handle of the malleus is drawn inward, bringing the membrana tympani with it,, and putting this membrane upon the stretch. On the relaxation of the muscle, the chain of bones returns to its ordinary position, by the elasticity of the neighboring parts, and the previous condition of the tympanic membrane is restored. This action, so far as we can judge, is purely involuntary. But the stapedius muscle is separately supplied by a minute branch of the facial nerve. It is probable that this arrange- ment enables us to make also a certain degree of voluntary exer- tion, in listening intently for faint or distant sounds. 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 temperature in the affected 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 temperature between the two sides at the end of ten minutes; and in the rabbit, at the end of half an htjur. 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 also be measured by the thermo- meter. Bernard' has found it to reach 8° or 9° F. The elevation of temperature and congested state of the parts are sometimes found to be diminished by the next day, and afterward disappear rapidly. ' Rechcrches Gxpfrimentales sur le Grand Sympathique. Paris, 1854. 522 SYSTEM OF THE GREAT SYMPATHETIC. Occasionally, however, they last for a long time. Bernard (op, cit.) has seen the unnatural temperature of the affected parts remain, in the rabbit, from fifteen to eighteen days, and in the dog for two months. Where the superior cervical ganglion has been extirpated, he has even found the above appearances to continue, in the dog, for a year and a half. They may also, according to the same authority, be reproduced several times in the same animal, by repeated divi- sions of the sympathetic nerve. The above effect is due to a peculiar modification in the nutri- tion of the affected parts, which has some analogy with inflamma- tion. The unnatural heat, the congestion, and the increased sensi- bility which are present, all serve to indicate a certain resemblance between the two conditions. None of the more serious consequences of inflammation, however, such as cedema, exudation, sloughing or ulceration, have ever been known to follow from this operation; and the term inflammation, accordingly, cannot properly be applied to its results. Division of the sympathetic nerve in the middle of the neck has also a very singular and instantaneous effect on the muscular apparatus of the eye. Within a very few seconds after the above operation has been performed upon the cat, the pupil, of the cor- responding eye becomes strongly contracted, and remains in that condition. At the same time the third eyelid, or " nictitating mem- brane," with which these animals 5''g- i67. are provided, is drawn partially over the cornea, and the upper and lower eyelids also approxi- mate very considerably to each other; so that all the apertures guarding the eyeball are very perceptibly narrowed, and the ex- pression of the face on that side is altered in a corresponding degree. This effect upon the pupil has been explained by supposing^ the Cat, aftei aection of the right eympathetic. circular fibres of the. iris, Or the constrictors of the pupil, to be animated exclusively by nervous filaments derived from the oculo- motorius ; and the radiating fibres, or the dilators, to be supplied by the sympathetic. Accordingly, while division of the ooulo- raotorius would produce dilatation of the pupil, by paralysis of SYSTEM OF THE GREAT SYMPATHETIC. 523 the circular fibres only, division of the sympathetic would be followed by exclusive paralysis of the dilators, and a permanent contraction of the pupil would consequently take place. The above explanation, however, is not a satisfactory one ; since, in the first place, division of the oculo-motorius, as the experiments of Bernard have shown,^ does not by itself produce complete dilata- tion of the pupil ; and, secondly, after division of the sympathetic nerve in the cat, as we have already shown, not only is the pupil contracted, but both the upper and lower eyelids and the nictitating membrane are also partially drawn over the cornea, and assist in excluding the light. The last-named effect cannot be owing to any direct paralysis, from division of the fibres of the sympathetic. It is more probable that the section of this nerVe operates simply by exaggerating for a time the sensibility of the retina, as it does that of the integument ; and that the partial closure of the eyelids and pupil is a secondary consequence of that condition. It will be remembered that in describing the inflammation of the eyeball, consequent upon section of the fifth pair of nerves, we found that there were reasons for believing this effect to be due to injury of certain sympathetic fibres which accompany the fifth pair. If the fifth pair in fact be divided at the level of the Cas- serian ganglion, where it is joined by sympathetic fibres from the carotid plexus, or between this ganglion and the eyeball, a destruc- tive infiammation of the organ follows. But if the section be made behind the ganglion, so as to avoid the filaments of communication with the sympathetic, no inflammatory change takes place. If this fact be really owing to the presence of sympathetic fibres which accompany the fifth pair, it indicates a remarkable difference in the effects of dividing the sympathetic near the eyeball and at a dis- tance from it; since no real inflammation of the eyeball or its appendages is ever produced by division of this nerve in the middle of the neck, but only the elevation of temperature and increase of sensibility, which have been already described. The influence of the sympathetic nerve and the consequences of its division upon the thoracic and abdominal viscera have been only very imperfectly investigated by experimental methods. It undoubtedly serves as a medium of reflex action between the sensi- tive and motor portions of the digestive, excretory, and generative ' Le(;on9 sur la Pliysiologie et la PatUologie du Systfeme Nervenx, Paris, 1858, Tol. ii. p. 203. 524 SYSTEM OF THE GREAT SYMPATHETIC. apparatuses ; and it is certain that it also takes part in reflex actions in which the cerebro-spinal system is at the same time interested. There are accordingly three different kinds of reflex action, taking place wholly or partially through the sympathetic system, which may be observed to occur in the living body. 1st. Reflex, actions takings place from the internal organs, through the sympathetic and cerebro-spinal systems, to the voluntary miiscles and sensitive surfaces. — The convulsions of young children are often owing to the irritation of undigested food in the intestinal canal. Attacks of indigestion are also known to produce temporary amau- rosis, double vision, strabismus, and even hemiplegia. Nausea, and a diminished or capricious appetite, are often prominent symptoms of early pregnancy, in'duced by the peculiar condition of the uterine mucous membrane. 2d. Reflex actions taking place from the sensitive surfaces, through the cerebrospinal and sympathetic systems, to the involuntar'y muscles and secreting org'ans.-r— Imprudent exposure of the integument to cold and wet, will often bring on a diarrhoea. Mental and moral impressions, conveyed through the special senses, will affect the motions of the heart, and disturb the processes of digestion and secretion. Terror, or an absorbing interest of any kind, will pro- duce a dilatation of the pupil, and communicate in this way a pecu- ' liarly wild and unusual expression to the eye. Disagreeable sights or odors, or even unpleasant occurrences, are capable of hastening or arresting the menstrual discharge, or of inducing premature delivery. 3d. Reflex actions taking place through the sympathetic system from one part of the internal organs to another. — The contact of food with the mucous membrane of the small intestine excites a peristaltic movement in the muscular coat. The mutual action of the diges- tive, urinary and internal generative organs upon each other takes place through the medium of the sympathetic ganglia and their nerves. The variations of the capillary circulation in different abdominal viscera, corresponding with the state of activity or re- pose of their associated organs, are to be referred to a similar nerv- ous influence. These phenomena are not accompanied by any consciousness on the part of the individual, nor by any apparent intervention of the cerebro-spinal system; SECTION III. REPRODUCTION. CHAPTER I. ON THE NATURE OP REPRODUCTION, AND THE ORIGIN OP PLANTS AND ANIMALS. The process of reproduction is the most characteristic, and in many respects the most interesting, of all the phenomena presented by organized bodies. It includes the whole history of the changes taking place in the organs and functions of the individual at suc- cessive periods of life, as well as the production, growth, and de- velopment of the new germs which make their appearance by generation. For all organized bodies pass through certain well-defined epochs or phases of development, by which their structure and functions undergo successive alterations. We have already seen that the living animal or plant is distinguished from inanimate substances by the incessant changes of nutrition and growth which take place in its tissues. The muscles and the mucous membranes, the osse- ous and cartilaginous tissues, the secreting and circulatory organs, all incessantly absorb oxygen and nutritious material from with- out, and assimilate their molecules; while new substances, produced by a retrogressive alteration and decomposition, are at the same time excreted and discharged.* These nutritive changes correspond in rapidity with the activity of the other vital phenomena ; since the production of these phenomena, and the very existence of the vital functions, depend upon the regular and normal continuance of the nutritive process. Thus the organs and tissues, which are always the seat of this double change of renovation and decay, ( 525 ) 526 NATURE OP REPEOUUCTION. retain nevertlieless their original constitution, and continue to be capable of exhibiting the vital phenomena. The above changes, however, are not in reality the only ones which take place. For although the structure of the body and the composition of its constituent parts appear to be maintained in an unaltered condition,^ by the nutritive pi'ocess, from one moment to another, or from day to day, yet a comparative examination of them at greater intervals of time will show that this is not pre- cisely the case ; but. that the changes of nutrition are, in point of fact, progressive as well as momentary. The composition and pro- perties of the skeleton, for example, are not the same at the age of twenty-five that they were at fifteen. At the later period it con- tains more calcareous and less organic matter than before ; and its solidity is accordingly increased, while its elasticity is diminished. Even the anatomy of the bones alters in an equally gradual manner; the medullary cavities enlarging with the progress of growth, and the cancellated tissue becoming more open and spongy in texture. We have already noticed the difference in the quantity of oxygen and carbonic acid inspired and exhaled at difterent ages. The muscles, also, if examined after the lapse of some years, are found to-be less irritable than formerly, owing to a slow, but steady and permanent deviation in their intimate constitution. The vital properties- of the organs, therefore, change with their varying structure ; and a time comes at last when they are per- ceptibly less capable of performing their original functions than before. This alteration, being dependent on the varying activity ' of the nutritive process, continues necessarily to increase. The very exercise of the vital powers is inseparably connected with the sub- sequent alteration of the organs employed in them ; and the func- tions of life, therefore, instead of remaining indefinitely the same, pass through a series of successive changes, which finally terminate in their complete cessation. The histpry of a living animal or plant is, therefore, a history of successive epochs or phases of existence, in each of which the struc- ture and functions of the body differ more or less from those in every other. Every living being hai a definite terra of life, through which it passes by the operation of an invariable law, and which, at some regularly appointed time, comes to an end. The plant germinates, grows, blossoms, bears fruit, withers, and decays. The animal is born, nourished, and brought to maturity, after which he retrogrades and dies. The very commencement of existence, by NATUBB OF EEPBODUCTIOIT. 527 leading through its successive intermediate stages, conducts at last necessarilj to its own termination. But while individual organisms are thus constantly perishing and disappearing from the stage, the particular kind, or species, remains in existence, apparently without any important change in the cha- racter or appearance of the organized forms belonging to it. The horse and the ox, the oak and the pine, the different kinds of wild and domesticated animals, even the different races of man himself, have remained without any essential alteration ever since the earliest historical epochs. Yet during this period innumerable individuals, belonging to each species or race, must have lived through their natural term and successively passed out of existence. A species may therefore be regarded as a type or class of organized beings, in which the particdlar forms or structures composing it die oflF con- stantly and disappear, but which nevertheless repeats itself from year to year, and maintains its ranks constantly full by the regular accession of new individuals. This process, by which new organ- isms make their appearance, to take the place of those which are destroyed, is known as the process of reproduction or generation. Let us now see in what manner it is accomplished. It has always been known that, as a general rule in the process of generation, the young animals or plants are produced directly from the bodies of the elder. The relation between the two is that of parents and progeny ; and the new organisms, thus generated, become in turn the parents of o^ers which succeed them. For this reason wherever such plants or animals exist, they indicate the previous existence of others belonging to the same species ; and if by any accident the whole species should be destroyed in any par- ticular locality, no new individuals could be produced there, unless by the previous importation of others of the same kind, , The commonest observation shows this to be true in regard to those animals and plants with whose history we are more familiarly acquainted. An opinion, however, has sometimes been maintained that there are exceptions to this rule ; and that living beings may, under certain circumstances, be produced from inanimate substances, without any similar plants or animals having preceded them ; pre- senting, accordingly, the singular phenomenon of a progeny without parents. Such a production of organized bodies is known'by the name of spontaneous generation. It is believed by the large majority of physiologists at the present day that no such spontaneous gene- ration ever takes place; but that plants and animals are always 528 NATURE OF REPRODUCTION. derived, by direct reproduction, from previously existing parents of the same species. As this, however, is a question of some im- portance, and one which has been frequently discussed in works on physiology, we shall proceed to pass in review the facts which have been adduced in favor of the occurrence of spontaneous generation, as well as those which would lead to its disproval and rejection. It is evident, in the first place, that many apparent instances of spontaneous generation are found to be of a very different character as soon as they are subjected to a critical examination. Thus grass- hoppers and beetles, earthworms and crayfish, the swarms of minute insects that fill the air over the surface of stagnant pools, and even frogs, moles, and lizards, have been supposed in former times to be generated directly from the earth or the atmosphere ; and it was only by investigating carefully the natural history of these animals that they were ascertained to be produced in the ordinary manner by generation from parents, and were found to continue the repro- duction of their species in the same way. A still more striking instance is furnished by the production of maggots in putrefying meat, vegetables, flour paste, fermenting dung, &c. If a piece of meat be exposed, for example, and allowed to undergo the process of putrefaction, at the end of a few days it will be found to contain a multitude of living maggots which feed upon the decomposing flesh, l^ow these maggots are always produced under the same conditions of warmth, moisture and exposure, and at the same stage of the putrefactive process. They are never to be found in fresh meat, nor, in fact, in any other situation than the one just mentioned. They appear, consequently, without any similar individuals having existed in the same locality ; and considering the regularity of their appearance under the given conditions,, and their absence elsewhere, it has been believed that they were spontaneously generated, under the influence of warmth, moistnre, and the atmosphere, from the decaying organic substances. A little examination, however, discovers a very simple solution of the foregoing difficulty. On watching the exposed animal or vegetable substances during the earlier periods of their decompo- sition, it is found that certain species of flies, attracted by the odor of the decaying material, hover round it and deposit their eggs upon its surface • or in its interior. These eggs, hatched by the warmth to which they are exposed, produce the maggots ; which are simply the young of the winged insects, and which after a time become transfornied, by the natural progress of development, into INFUSORIAL ANIMALCULES. 529 perfect insects similar to their parents. The difficulty of account- ing for the presence of the maggots by generation, therefore, de- pends simply on the fact that they are different in appearance from the parents that produce them. This difference, however, is merely a temporary one, corresponding with the difference in age, and dis- appears when the development of the animal is complete ; just as the young chicken, when recently hatched, has a different form and plumag^ from those which it presents in its adult condition. Nearly all the causes of error, in fact, which have suggested at various times the doctrine of spontaneous generation, have been derived from these two sources. First, the ready transportation of eggs or germs, and their rapid hatching under favorable circum- stances ; and secondly, the different appearances presented by the same animal at different ages, in consequence of which the youthful animal may be mistaken, by an ignorant observer, for an entirely different species. These sources of error are, however, so readily detected, as a general rule, by scientific investigation, that it is hardly necessary to point out the particular instances in which they exist. In fact, whenever a rare or comparatively unknown animal or plant has been at any time supposed to be produced by sponta- neous generation, it has only been necessary, for the most part, to investigate thoroughly its habits and functions, to discover its secret methods of propagation, and to show that they correspond, in all essential particulars, with the ordinary laws of reproduction. The limits, therefore, within which the doctrine of spontaneous genera- tion can be applied, have been narrowed in precisely the same degree that the study of natural history and comparative physiology has advanced. At present, indeed, there remain but two classes of phenomena which are ever supposed to lend any support to the above doctrine ; viz., tlie existence and production, 1st, of infuso- rial animalcules, and 2d, of animal and vegetable parasites. We shall now proceed to examine these two parts of the subject in succession. Infusorial Animalcules.— If water, holding in solution or. ganic substances, be exposed to the contact of the atmosphere at- ordinary temperatures, it is found after a short time to be filled with swarms of minute living organisms, which are visible only by the microscope. The forms of these microscopic animalcules are exceedingly varied ; owing either to the great number of species in existence, or to their rapid alteration during the successive pe- 34 530 NATURE OP EEPKODUOTIOIf. Different kinds of Infusoria. riods of their growth. Ehrenberg has described more than 300 different varieties of them. They are generally provided with cilia attached to the exterior of their bodies, and are, for the most part, in constant and rapid motion in the fluid which they inhabit. Owing to their presence in '^' ■ ' animal aod vegetable watery infusions, they have received the name of "infusoria," or " infusorial animalcules,". Now these infusoria are always produced under the conditions which we have de- scribed above. The animal or vegetable substance used for the infusion may be pre- viously baked or boiled, so as to destroy all living germs' which it might accidentally contain; the water in which it is infused may be carefully distilled, and thus freed from all similar contamination; and yet the infusorial animalcules will make their appearance at the usual time and in the usual abundance. It is only requisite that the infusion be exposed to a moderately elevated temperature, and to the access of atmospheric air; conditions which are equally necessary for maintaining the life of all animal and vegetable organisms, what- ever be the source from which they are derived. Under the above circumstances, therefore, either the animalcules must have been produced by spontaneous generation in the -water j infusion, or their germs must have been introduced into it through the medium of the atmosphere. No such introduction has ever been directly de- monstrated, nor have even any eggs or germs belonging to the infusoria ever been detected. Nevertheless, there is every probability that the infusoria are produced from germs, and not by spontaneous genei-ation. Since ■ the infusoria themselves are microscopic in size, it is not surprising that their eggs, which must be smaller still, should have escaped observation. We know, too, that in many instances the minute germs of animals or plants may be wafted about in a dry state by the atmosphere, until, by accidentally cowling in contact with warmth and moisture, they become developed and bring forth living organ- INPgSOBIAL .ANIMALCOLKS. 531 isms. The eggs of the infusoria, accordingly, may be easily raised, and held suspended in the atmosphere, under the form of minute dust-like particles, ready to germinate and become developed when- ever they are caught by the surface of a stagnant pool, or of any artificially prepared infusion. . In point of fact, the atmosphere does really contain an abundance of such dust-like particles, even when it appears to be most transparent and free from impurities. This may be readily demonstrated by admitting a single beam of sunshine into a darkened apartment, when the shining particles sus- pended in the atmosphere become immediately visible in the track of the sunbeam. Again, if a perfectly clean and polished mirjor be placed with its face upward in a securely closed room, and left undisturbed for several days, its surface at the end of that time will be found to be dimmed by the settling upon it of minute dust, deposited from the atmosphere. There is no reason, therefore, for disbelieving that the air may always contain a sufl&cient number of organic germs for the production of infusorial animalcules. There is some difficulty, however, in obtaining direct proof that it is through the medium of the atmosphere that organic germs pene- trate into the watery infusions. It is true that if such an infusion be prepared from baked meat or vegetables and distilled water, and afterward hermetically sealed, no infusoria are developed in it ; but this only shows, as we have already intimated, that the free access of air is necessary to the development of all organic life, just as it is to the support of animals and plants under ordinary conditions of growth and reproduction. Such a result, therefore, proves nothing with regard to the external origin of the infusoria. In order to be conclusive, such an experiment should be so contrived that the watery infusion, previously freed from all foreign contamination, should be supplied with a free access of atmospheric air, while the introduction of living germs by this channel should at the same time be rendered impossible. An experiment of this kind has in reality been contrived and successfully carried out by Schultze, of Berlin.' This observer prepared an infusion containing organic substances in solution, and inclosed it in a glass flask (Fig. 169, a) of such a size, that the infusion filled about one-half the entire capacity of the vessel. The mouth of the flask was fitted with an air-tight stopper provided with two holes, through which were passed narrow glass tubes bent at right angles. To each of these tubes was attached a ■ Edinburgh New PhiloEophical Jonrnal, Oct. 1837. 532 NATURE OF KEPRODUCTION. Fig. 169. potash-apparatus {b, c), similar to those used for condensing carbonic acid in organic analyses. One of these (6) was filled with concen- trated sulphuric acid, the other (c) with a solution of caustic potassa. The flask with the organic infusion having been subjected to a boiling temperature, in order to destroy any living germs which it might con- tain, the stopper was inserted, and the whole apparatus exposed to the light, at the ordinary summer tempera- ture. The connections of the apparatus being perfectly tight, no air could pene- trate into the flask, except by passing through either the sulphuric acid or the potassa; either of which would retain and destroy any organic germs which might be suspended in it. Every day a fresh supply of air was introduced into the flask by drawing it through the tubes 6, c ; and in this way the atmospheric air above the infu^ sion was constantly renewed, while at the same time the introduction of living germs from without was effectually prevented. Schultze kept this apparatus under his observation, as above, from the last of May till the first of August; frequently examining the edges of- the fluid with a lens, through the sides of the glass jar, but without ever detecting in it any traces of living organisms. At the end of that period the flask was opened, and the fluid which it contained subjected to direct examination, equally without result. It was then exposed, in the same vessel and in the same situation as before, to the free access of the atmosphere, and at the end of two or three days it was found to be swarming with infusoria. It is plain, therefore, that the infusoria cannot be regarded as produced by spontaneous generation, but must be considered as originating in the usual manner from germs; since they do not make their appearance in the watery infusion, when the accidental introduction of germs from without has been effectually prevented. Schnltze's experiment oa SpoWTA- XEOcs Genebatioh.— a. Flask con- taining watery infasion. b. Potash-ap- paratus containing sulphuric acid. c. Potash-apparatuH containing caustic po- tassa. Animal- and Vegetable Parasites. — This very remarkable group of organized bodies is distinguished by the fact that they, live either upon the surface or in the interior of other animal or vegetable organisihs. Thus, the mist letoe fixes itself on the branches ANIMAL AND VEGETABLE PARASITES. 636 of aged trees ; the Oidium albicans vegetates upon the mucous sur- faces of the mouth and pharynx ; the Botrytis Bassiana attacks the body of the silkworm, and plants itself in its tissues ; while many species of trematoid worms live attached to the gills of fish and of ■water-lizards. These plarasites are usually nourished by the fluids of the animal whose body they inhabit. Bach particular species of parasite is found to inhabit the body of a particular species of animal, and is not found elsewhere. They are met with, moreover, as a general, rule, only in particular organs, or even in particular parts of a single organ. Thus the Tricocephalus dispar is found only in the CEBCum; the Strongylus gigas in the kidney; the Distoma hepati- cum in the biliary passages. The Distoma variegatum is found only in the lungs of the green frog, the Distoma cylindraceum in those of the brown. The Taenia solium is found in the intestine of the human subject in certain parts of Europe, while the Taenia lata occurs exclusively in others. It appears, therefore, as though some local combination of/ conditions were necessary to the production of these parasites; and they have been supposed, accordingly, to originate by spontaneous generation in the localities where they are exclusively known to exist. A little consideration will show, however, that the above condi- tions are not, 'properly speaking, necessary or sufficient for the production, but only for the development of these parasites. All the parasites mentioned above reproduce their species by generation. They have male and female organs, and produce fertile eggs, often in great abundance. The eggs coutained in a single female Ascaris are to be counted by thousands ; and in a tapeworm, it is said, even by millions. Now these eggs, in order that they may be hatched and produce new individuals, require certain special conditions which are favorable for their development; in the same manner' as the seeds of plants require, for their germination and growth, a certain kind of soil and a certain supply of warmth and moisture. It is accordingly no more surprising that the Ascaris vermicularis should inhabit the rectum, and the Ascaris lumbricoides the ileum, than that the Lobelia inflata should grow only in dry pastures, and the Lobelia cardinalis by the side of running brooks. The lichens flourish on the exposed surfaces of rocks and stone walls ; while the fungi vegetate in darkness and moisture, on the decaying trunks of dead trees. Yet no one imagines these vegetables to be spon- taneously generated from the soil which they inhabit. The truth 534 NATUBK OF REPRODUCTION. is simply this, that if the animal or vegetable germ be deposited in a locality which affords the requisite conditions for its development, it becomes developed ; otherwise not. Each female Ascaris pro- duces, as we have stated above, many thousands of ova. Now, though the chances are very great against any particular one of these ova being accidentally transported into the intestinal canal of another individual, it is easy to see that there are many causes in operation by which some of them might.be so transported. By far the greater number undoubtedly, perish, from not meeting with the conditions necessary for their development. One in a thousand, or perhaps one in a million, is accidentally introduced into the body of another individual, and consequently becomes developed there into a perfect Ascaris. The circumstance, therefore, that particular parasites are confined to particular localities, presents no greater difficulty as to their mode of reproduction, than the same fact regarding other animal and vegetable organisms. Neither is there any difficulty in accounting for the introduction of .parasitic germs into the interior of the body. The air and the food offer a ready means of entrance into the respiratory and digestive passages ; and, a parasite once introduced • into the intes- tine, there is no difficulty in accounting for its presence in any of the ducts leading from or opening into the alimentai-y canal. Some parasites are known to insinuate themselves directly underneath the surface of the skin; as the Pulex penetrans or ''chiggo" of South America, and the Ixodes Americanus or "tick." Others, like the (Estrus bovis, penetrate the integument for the purpose of depositing their eggs in the subcutaneous areolar tissue. Some may even gain an entrance into the bloodvessels, and circulate in this way all over the body. Thus the Filaria rubella is found alive in the bloodvessels of the frog, the Distoma haematobium in those of the human subject, and a species of Spiroptera in those of the dog. It is easy to see, therefore, how, by such means, parasitic germs may be conveyed to any part of the body ; and may even be deposited, by accidental arrest of the circulation, in the substance of the solid organs. The most serious difficulty, however, in the way of accounting for the production of parasitic organisms, was that presented by the existence of a.class known as the encysted or sexless entozoa. These parasites for the most part occupy the interior of the solid organs and tissues, into which they could not have gained access by the ANIMAL ANJJ VEGETABLE PARASITES. 535 mucous canals. Thus the Ccenurus cerebralis is found imbedded in the substance of the brain, the Trichina spiralis between the ^ fibres of the voluptary muscles, and the Cysticercus cellulosse in the areolar tissue of various parts of the body. They are also distin- guished from all other parasites by two peculiar characters. First, they are inclosed in a distinct cyst, with which they have no organic connection and from which they may be readily separated; and se- condly, they have no genera- tive organs, nor is there any Fig. 170. apparent difference between the sexes. The Trichina spi- ralis, for example (Fig. 170), is inclosed in an ovoid or spindle-shaped cyst, swollen in the middle and tapering at each extremity, with a round- „ „ , , \. Trichina Spiralis: from r«tun femoriN mus- ed cavity in its central por- cle br human subject. Magumed SI diameters. tion, in which the worm lies '^ coiled up in a spiral form. The worm itself has neither testicles nor ovaries, nor does it present any trace of a sexual organization. Now we have seen that it is easy to account for the conveyance of these or any other parasites into the interior of vascular organs and tissues ; the eggs from which they are produced being trans- ported by the bloodvessels to any part of the body, and there retained by a local arrest of the capillary circulation. In the case of the encysted entozoa, however, we have a much greater diffi- culty; since these parasites, are entirely without sexual organs or generative apparatus of any sort, nor have they ever been dis- covered in the act of producing eggs, or of developing in any manner a progeny similar to 'themselves. It appears, accordingly, difficult to understand how animals, which are without a sexual apparatus, should have been produced by .sexual generation. As it is certain that they can have no progeny, it would seem equally evident that they must have been produced without a parentage. This difficulty, however, serious as it at first appears, is susceptible of a very simple explanation. The case is in many respects analogous to that of the maggots, hatched from the eggs of flies in putrefying meat. These maggots are also without sexual organs; for they are still imperfectly developed, and in a kind of embryonic cpndi- tion. It is only after their metamorphosis into perfect insects, that generative organs are developed and a distinction between the 536 NATUKB OF BEPBODUCTIOK. Pig. 171. sexes manifests itself. This is, indeed, more or less the case with all animals and with all vegetables. The blossom, which is the sexual apparatus of the plant, does not appear, as a general rule, until the, growth of the vegetable has continued for a certain time, and it has acquired a certain age- and strength. Even in the human subject the sexual organs, though present at birth, are still very imperfectly developed as to size, and altogether inactive in func- tion. It is only later that these organs acquire their full growth, and the sexual characters become complete. In very many of the lower animals the sexual organs are entirely absent at birth, and appear only at a later period of development. Now the encysted or sexless entozoa are simply the undeveloped young of other para- sites which propagate by sexual generation; the membrane in which they are inclosed being either an embryonic envelope, or else an adventitious cyst formed round the para- sitic embryo. These embryos have come, in the natural course of their migrations, into a situation which is not suitable for their com- plete development. Their development is accordingly arrested before it arrives at matu- 'rity ; and the parasite never reaches the adult condition, until removed from the situation in which it has been placed, and transported to a more favorable locality. The above explanation has been demon- strated to be the true one, more particularly with regard to the Taenia, or tapeworm, and several varieties of Cysticercus. The Txnia (Fig. 171) is a parasite of -vyhich different species are found in thes intestine of the human swbject, the dog, cat, fox, and other of the lower animals. Its upper extremity, termed the " head," con- sists of a nearly globular mass, presenting upon its lateral surfaces a set of four muscular disks, or " suckers,* and terminating anteriorly in a conical projection which is provided with a crown of curved processes or hooks, by wh^ph the parasite attaches itself to the intestinal mucous membrane. ' To thfs " head" succeeds a slender ribbon-shaped neck, which is at first smooth, but AKIMAL AND VEGETABLE PARASITES. 537 which soon becomes transversely wrinkled, and afterward divided into distinct rectangular pieces or " articulations." These articula- tions multiply by a process of successive growth or budding, from the wrinkled portion of the neck ; and are constantly removed farther and farther from their point of origin by new ones formed behind them. As they gradually descend, by the process of growth, farther, down the body of the tapeworm, they become larger and begin to exhibit a sexual apparatus, developed in their interior. In each fully formed articulation there are contained both male and female organs of generation ; and the mature eggs, which are produced in great numbers, are thrown off together with the articu- lation itself from the lower extremity of the tapeworm. Since the articulations are successively produced, as we have mentioned above, by budding from the neck and the tack part of the head, the para- site cannot be effectually dislodged by taking away any portion of the body, however large ; since it is subsequently reproduced from the head, and continues its growth as before. But if the head itself be removed from the intestine, no further reproduction of the articu- lations can take place. The Gysticercus is an encysted parasite, different varieties of which are found in the liver, the peritoneum, and the meshes of the areolar tissue in various parts of the body. It ccmsists (Fig. 172) first, of a globular sac, or cyst (a), which is not adherent to the tissues of the organ in which the parasite is found, but may be easily sepa- Fig. 172. Kg. 173. Gtsticbrcub. — a. External cyst, b In- ternal Bac, coDtaininf^ fluid, c. Narrow canal, formed by involution of walls'of sac, at the bottom of wliicli is tlie head of the tsnia. Cyi4Ticebcitb, unfolded. rated from them. In its interior is found another sac (ft), lying loose in the cavity of the former, and filled with a serous fluid. This second sac presents, at one point upon its surface, a puckered depression, leading into a long, narrow canal (c). This canal, which 538 NATURE OF REPRODUCTION. is formed by an iuvolution of the walls of the second sac, presents at its bottom a small globular mass, like the head of the Tania, provided with suckers and hooks, and supported upon a short slender neck. If the oiiter investing sac be removed, the narrow canal just described may be everted by careful manipulation, and the parasite will then appear as in Fig. 173, with the head and neck resembling those of a Taenia, but terniinating behind in a dropsical sac-like swelling, instead of the chain of articulations which are characteristic of the fully formed tapeworm. Now it has been shown, by the experiments of Kiichenmeister, Siebold, and- others, that the Oysticercus is only the imperfectly developed embryo, or young, 'of the Taenia. "When the mature articulation of the tapeworm is thrown off) as already mentioned, from its posterior extremity, the eggs which it incloses have already passed through a certain period of development, so that each one contains an imperfectly formed embryo. The articulation, contain- ing the eggs and embryos, is then taken, with the food, into the stomach of another animal; the substance of the articulation, to- gether with the external covering of the eggs, is destroyed by di- gestion, and the embryos are thus set free. They then penetrate through the walls of the stomach, into the neighboring organs or the areolar tissue, and becoming encysted in these situations, are there developed into cysticerci, as represented in Fig. 172. After- ward, the tissues in which they are contained being devoured by a third animal, the cysticercus passes into the intestine, fixes itself to the mucous membrane, and, by a process of budding, produces the long tape-like series of ■articulations, by which, it is finally con- verted into the full-grown Taenia. Prof. Siebold found the head of the Cysticercus fasciolaris, met with in the liver of rats and mice, presenting so close a resem- blance to the Taenia crassicOllis, inhabiting the intestine of the cat, that he was led to believe the two parasites to be identical. This identity was, in fact, proved by the experiments of Kiichenmeister ; and Siebold afterward demonstrated' the same relation to exist between the Cysticercus pisiformis, found in the peritoneum of rab- bits, and the Taenia serrata, from the intestine of the dog. This experimenter succeeded in administering to dogs a quantity of the cysticerci, fresh from the body of the rabbit, mixed with milk ; and ' In Baffalo Medical Journal, Feb. 1853 ; also in Siebold on Tape and Cystic •Torms, Sydenham translation : London, 1857, p. 59. ANIMAL AND VEGETABLE PARASITES. 539 on killing the dogs, at various periods after the meal, from three hours to eight weeks, he found the cysticerci in various stages of development in the intestine, and finally converted into the full grown Taenia, with complete articulations and mature eggs. Dr. Kiichenmeister' has also performed the same experiment, with success, on the human subject. A number of cystiperci were ad- ministered to a criminal, at different periods before his executiorf, varying from 12 to 72 hours ; and upon post-mortem examination of the body, no less than ten young tseniae were found in the intestine, four of whjoh could be distinctly recognized as specimens of Taenia solium. Finally, both Leuckart and Kiichenmeister' have shown, on the other hand, that the eggs of Taenia solium, introduced into the body of the pig, will give rise to the development of Cysticercus cellulosae ; thus demonstrating that the two kinds of parasites are identical in their nature, and differ only in the manner and degree of their development. There remains, accordingly, no good reason for believing that even the encysted parasites are produced by spontaneous genera- tion. Whatever obscurity may hang round the origin or reproduc- tion of any class or species of animals, the direct investigations of the physiologist always tend to show that they do not, in reality, form any exception to the general law in this respect ; and the only opinion which is admissible, from the facts at present within our knowledge, is that organized beings, animal and vegetable, wherever they may be found, are always the progeny of ^eviotisly existing parents. ' On Animal and Vegetable Parasites, Sydenham translation : London, 1857, p. 115. 2 Op. cit., p. 120. HO SSXUAL GEXEBATIOK. CHAPTER II. ON SEXUAL GENERATION, AND THE MODE OP ITS , ACCOMPLISHMENT. Fig. 174. The function of generation is performed by means of two sets of organs, each of which gives origin to a, peculiar product, capable of uniting with the other so as to produce a new individual. These two sets of organs, belonging to the two different sexes, are called the male and female organs of generation. The female organs produce a globular body called the germ, or egg, which is capable of being developed into the body of the young animal or plant ; the male organs produce a substance which is necessary to fecundate the germ, and enable it to go through with its natural growth and development. Such af6 the only essential and uni- versal characters of the organs of gene- ration. These organs, however, exhibit various additions and modifications in different classes of organized beings, while they show throughout the same fundamental and essential characters. In the flowering plants, for example, the blossom, which is the generative ?,pparatus (Fig. 174), consists first of a female organ containing the germ (a), situated usually upon the highest part of the leaf-bearing stalk. This is surmounted by a nearly straight column, termed the pistil (6), dilated at its summit into a globular expansion, and occupying the centre of the flower. Around it are arranged several slender filaments, or stamens, bear- ing upon their extremities the male organs, or anthers (c, c). The Blossom op CoNVOLVCLns PuBPliBEtTS. (MorDing glory. ) — a. Germ. b. Pistil, c. c. Stamens, with aachers. d. Corolla, e. Calyx. SEXUAL GENERATION. 541 Fig. 175. tvhole is surrounded by a circle or crown of delicate and brilliantly colored leaves, termed the corolla (c^, which is frequently provided with a smaller sheath of green leaves outside, called the calyx (e). The anthers, when arrived at maturity, discharge a fine organic dust, called the pollen, the granules of which are caught upon the extremity of the pistil, and then penetrate downward through its tissues, until they reach its lower extremity and come in contact with the germ. The germ thus fecundated, the process of genera- tion is accomplished. The pistil, anthers, and corolla wither and fall off, while the germ increases rapidly in size, and changes in form and texture, until it ripens into the mature fruit or seed. It is then ready to be separated from the parent stem ; and, if placed in the proper soil, will germinate and at last produce a new plant similar to the old. In the above instance, the male and female organs are both, situated upon the same flower ; as in the lily, the violet, the con- volvulus, &c. In other cases, there are separate male and female flowers upon the same plant, of which the male flowers produce only the pollen, the female, the germ and fruit. In others still, the male and female flowers are situated upon different plants, which otherwise resemble each other, as in the willow, poplar, and hemp. In animals, the female organs of generation are called ovaries, since it is in them that the egg, or "ovum," is produced.* The male organs are the testicles, which give origin to the fecun- dating product, or "seminal fluid," by which the egg is fer- tilized. "We have already men- tioned above that in the articula- tions of the tapeworm the ovaries and testicles are developed to- gether. (Fig. 175.) The ovary {a, a, a) is a series of branching follicles terminating in rounded extremities, and communicating with each other by a central canal. The testicle (6) is a narrow, convoluted tube, very much folded SiNQLB Articulation of Tj&ni^a CKA8S1C0LI.I8, from hmall iuteBtiue of cat.-* a, a, a. Ovary filled with egga. b. Testicle, v. Geoital orifice. 542 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 are 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 soinetimes said to be " hermaphrodite," or of double sex. In reality, 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 qpimals, 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 the anatomy of the two sexes. Such are the uterus and mammary glands of the female, the vesicul^ seminales and prostate gland of the male. The femal^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 getieral structure of the body, which affect the whole externa;! 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 GESEBATION. 5i3 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. 5U EGG AND FEMALE OKGANS OF GENEHATIOK. CHAPTER III. ON THE EGG, AND THE FEMALE ORGANS OP GENERATION. Fig. 176. 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 y'j of an inch in diameter, in the lamprey 5^5, in quadrupeds and in the human species t^j. 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 viiellus. The vitelline membrane in birds and reptiles is very thin, measur- ing often not more than 75^5^ of an inch in thickness, and is at the same time of a somewhat fibrous texture. In man and the higher animals, on the contrary, it is perfectly smooth, structure- less and transparent, and is about f ^V^r of an inch in thickness. Notwithstanding its delicate and transparent appearance, it has a considerable degree of resistance and elasticity. ♦The egg of the humau fibject, for example, may be perceptibly attened 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. 176) 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 HnHAS OrcB, masrnifled 85 diameters, a. ViteLllnenieinbraDe. &. Vitellns. c. Germinative vesicle. d. Germinatlve spot. ^ KGO AND FEMALE OEGAN3 OF GKNE.BATIOK. 545 vitelline membrane, however, is the one more generally adopted, and is also the more appropriate of the two. The vitellus (6) 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 gj^ to 5^^ of an inch in diameter. It presents upon its surface a dark spot, like a nucleus {d), which is known ^^^' ^ ' ^* by the name of the germinative spfit. The germinative vesicle, with its nupleus-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 carfeful examination. If the egg be ruptured by excessive pres- humah otuh, mptnred by sure under the microscope, the vitellus is ?"»"'••«; '"owiDg the vueimn , ' partially expelled, the germina- seen to have a gelatinous consistency. It twe Tesicie at a, and the «mooth is gradually expelled from the vitelline ™' "' "" '"""""' """"'• cavity, but still retains the granules and oil globules entangled in its substaifoe. (Fig. 177.) 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*gg 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 inclosufe, intended to secure the vitellus from injury, and enable it to retain its figure during the early 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 animals. In nearly all thf se species, in fact, where the fecundated 35 • 54r6 EGG ANU FEMALE ORGANS OF GENEBATION. 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- ment of the young is completed, such an egg as above described is sufficient for the formation of the embryo ; since during its develop- raenj 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 deve- lopment 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 thg egg with asecretion, or to regard the ovary as in any respect resembling a glandular organ. The egg is simply an organized bo^, 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 spe8,king, 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 EGG AND FEMALE ORGANS OF GBNEKATION. 547 Fig. 178. oviducts, which are destined to receive the eggs at their internal extremity and convey them to the external generative orifice. The 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 corftmences 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. 178), 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 eggs which they contain, undergo at particular sea- sons a periodical development or increase in growth™ If we examine the female frog in the latter part of summer or the fall, we shall find the ovaries presAting the appearance of small clusters of minute and nearly colorless eggs, the smaller of which are perfectly (transparent and not over i\js 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 thei~r former size, and forming large lobulated masses, crowded with dark-colored opaque eggs, measur- ing jij of an inch 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 increase in size and become somewhat altered in structure. The vitellus 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, Ffmale Gekebatitr Or- GAKB OP Froq. — a. n. Ovaries. &, b. Oviducts, c, c. Their internal orifices, d. Cloaca, showing exter- naL orifices of oviducta. 548 EOG AND 7EMAL£ OBGAN3 OF OENEBATION. brown, yellow, or orange color. In the human subject, however, the change consists only in an increase of size and granulatioH, 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 fly 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 expelted 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 diiferent crops may be recongized 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 inafctive 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, bi-monthly,, or even monthly; hut 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 riiost of the other vital phenomena. Action of the Oviducts and Fernale Generative Passages.-^It frogs and lizards, the ripening and discharge of the eggs take place, as ^bove mentioned in the early spring. At the time of leaving the ""■ary, the eggs consist simply of the '^" dark-colored and granular vitellus, ^ps^l^l^ inclosed in the vitelline membrane. ^^P«^ They are then received by the inner • ••• y®^F^ extremity of the oviducts, and carried ® 'imm^'lM downward by the peristaltic move- -(, ment of these canals, aided by the mattre froqb' Eaaa.— /I. While more powcrful Contraction of the t« tt ;:;::. *• ''" "^""^ abdominal muscles. During the pas- sage of the eggs, moreover, the mucous membrane of the oviduct secretes a colorless, viscid, albuminoid substance, which is deposited in successive layers round each egg, forming a thick and tenacious coating or finvelope. (Fig. 179,) When the eggs are finally discharged, this albuminoid matter EGG AND FEMALE OBGANS OF GENERATION. 549 absorbs the water in wliicli the spawn is deposited, and swells np into a transparent 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 develdpment and early growth of the embryo. • In the terrestrial reptiles and in birds, the oviducts perform a still more important secretory function, la the common fowl, the ovary consists, as in the frog, of a large number of follicles, loosely connected by areolariissue, in which the eggs can be seen in different stages of development. (Fig. 180, 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 occur^, 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 the vitellus, is a round white spot, consisting of ^ layer of minute granules, termed the " cicatricula." It is in the central part of the cicatricula that the germinative vesicle is fount} 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, d), where the mucous mem- brane is smooth and transparent, the yolk merely absorbs a certain quantity of fluid, so as to become more flexible and yielding in con- sistency. It then passes into a second division of the generative 550 EGG AND FEMALE OB6AN3 OF GENERATION. canal, in which the mucous membrane is thick and glandular in texture, and is also thrown into numerous longitudinal folds, which project into the cavity of the oviduct. This portion of the oviduct {d, e), extends over about nine inches of its entire length. In its upper part, the mucous membrane secretes a viscid material, by which the yolk is encased, and which soon consolidates into a gela- tinous, membranous deposit ; thus forming a second homogeneous layer, outside the vitelline membrane. Now the peristaltic movements of this part of the oviduct are such as to give a rotary, as well as a progressive motion to the egg ; and the two extremities of the membranous layer described above become, accordingly, twisted in opposite directions into two fine cords, which run backward and forward from the opposite poles of the egg. These cords are termed the '^chalazse," and the mem- brane with which they are connected, the " chalaziferous membrane." Throughout the remainder of the second division of the oviduct, the mucous membrane exudes an abundant, gelatinous, albuminoid substance, which is deposited in successive layers round the yolk, inclosing at the same time the chalaziferous membrane and the chalazse. This substance, which forms the so-called albumen, or " white of egg," is semi-solid in consistency, nearly transparent, and of a faint amber color. It is deposited in greater abundance in front of the advancing egg than behind it, and forms accordingly a pointed or conical projection in front, while behind, its outline is rounded oil, parallel with the spherical surface of the yolk. In this way, the egg acquires, when covered with its albumen, an ovoid form, of which One end is round, the other pointed ; the pointed extremity being always directed downward, as the egg descends along the oviduct. In the third division of the oviduct (J), which is about three and a half inches in length, the mucous membrane is arranged in longi- tudinal folds, which are narrower and more closely packed than in the preceding portion. The material secreted in this part, and de- posited upon the egg, condenses into a firm fibrous covering, com- posed of three different layers which closely embrace the surface of the albuminous mass, forming a tough, flexible, semi-opaque envelope for the whole. These layers are known as the external, middle, and internal fibrous membranes of the egg. Finally the egg passes into the fourth division of the oviduct {g), which is wider than the rest of the canal, but only a little over two inches in length. Here the mucous membrane, which is arranged EGG AND FEMALE ORGANS OP GENERATION. 551 Fig. 180. in abundant, projecting, leaf-like villosities> exudes a fluid very rich in calcareous salts. The most external of the three membranes just described is permeated by this fluid, and very soon the calcare- ous matter begins to crystallize in the interstices of its fibres. This deposit of calcareous matter goes on, growing constantly thicker and more condensed, until the entire external membrane is converted into a white, opaque, brittle, calcareous shell, which incloses the remaining portions and protects them from ex- ternal injury. The egg is then driven outward by the contraction of the muscular coat through a narrow por- tion of the oviduct (A), and, gradually dilating the passages by its conical extremity^ is finally discharged from the external orifice. The egg of the fowl, after it has been discharged from the body, con- sists, accordingly, of various parts; some of which, as the yolk and the vitelline membrane, entered into its original formation, while the remain^ der have been deposited round it dur- ing its passage through the oviduct. On examining such an egg (Fig. 181), we find externally the calcareous shell (A), while immediately beneath it are situated the middle and internal fibrous shell-membranes (e,/). Soon after the expulsion of the egg there is a partial evaporation of its watery ingredients, which are replaced by air penetrating through the pores Femalk Oexeratite Oroahs of Fowl..— n. Ovarf. b. GmaSan follicle, from vBich th* egg has jnst been discharged, c. Volk, entering upper extremity ol ovidaci. d, e. Second diVUiou of OTiduct, in which ohalazlfsroas membraa.i, chalazia, and albumen are formed. /. Third p.trtion. In which the flbroas shell membranes are produced, ff. Fourth portion laid open, showing egg completely formed, with calcareous shell, h. Narrow canal throui^b. which the egg is discharged. 55.2 EGG AND FKMALK ORGANS OF GENEBATION. « Hi the shell at its rounded extremity. The air thus introduced' accumulates between the middle and internal fibrous membranes at this spat, separating them from each other, and forming a cavity or air-chamber (g), which is always found between the. two fibrous metabranes at the rounded end of the egg. Next we come to the albumen or " white" of the egg (d) ; next to the chalaziferous mem- brane and chalazae (c) ; and finally to the vitelline membrane (J) Fig. 181. Diairram of Fowl's Egg. -^ffl. Tolk. &. Titelline membrane, c. Chalaziferoas membrane, d. Albumea. d, /. Middle and internal shell membranes, g. Air-chamber. A. Calcareous sbelL inclosing the yolk (a). After the expulsion of the egg, the external layers of the albumen liquefy; and the vitellus, being specifically lighter than the albumen, owing to the large proportion of oleagin- ous matter which it contains, rises toward the surface of the egg, with the cicatricula uppermost. This part, therefore, presents itself almost immediately on breaking open the egg upon its lateral surface, and is placed in the most favorable position for the action of warmth and atmospheric air in the development of the chick. The vitellus, therefore, is still the essential and constituent portion of the egg ; while all the other parts consist either of nutritious mate- rial, like the albumen, provided for the support of the embryo, or of protective envelopes, like the shell and the fibrous membranes. In the quadrupeds, another and still more important modification of the oviducts takes place. In these animals, the egg, which is originally very minute in size, is destined to be retained within the generative passages of the female during the development of the embryo. "While the upper part of the'bviduct, therefore, is quite narrow, and intended merely to transmit the egg from the ovary, EGG AND FEMALE ORGANS OF GENERATION. 553 and to supply it with a little albuminous secretion, its lower por- tions are very much increased in size, and are lined, moreover, with a mucous membrane, so constructed as to provide for the protection and nourishment of the embryo, during the entire period of gesta- tion. The upper and narrower portions of the oviduct are known as the "Fallopian tubes" (Fig. 182); while the lower and more Fig. 182. Uterus and Ovaries op the Sow.— «,(?. OTarieK, &,&. Fallopian tubes. .ff,c. Horns uf nieiuH. d. Body of uterus, c. Vagina. highly developed portions constitute the uterus. These lower por- tions unite with eaclu'Other upon the median line near their infe- rior termination, so as to form a central organ, termed the " body" of the uterus ; while the remaining ununited parts are known as its " cornua" or " horns." In the human subject, the female generative apparatus presents the following peculiarities. The ovaries consist of Graafian follicles, which are imbedded in a somewhat dense areolar tissue, supplied with an abundance of bloodvessels. The entire mass is covered with a thick, opaque, yellowish' white layer of fibrous tissue called the " albugineous tunic." Over the whole is a layer of peritoneum, which is reflected upon the vessels which supply the ovarj', and is continuous with the broad ligaments of the uterus. The oviducts commence by a wide expansion, provided with fringed edges, called the " fimbriated extremity of the Fallopian tube." The Fallopian tubes themselves are very narrow and con- voluted, and terminate on each side in the upper part of the body of the uterus. In the human subject, the body of the uterus is so much developed at the expense of the cornua, that the latter hardly appear to have an existence ; and in fact no trace of them is visible externally. But on opening the body of the uterus its cavity, is 554 EGG AND FEMALE OBGANS OF 6ENEBATION. seen to be nearly triangular in shape, its two superior angles run- ning out on each side to join the lower extremities of the Fallopian tubes. This portion evidently consists of the comua, which have been consolidated with the body of the uterus, and enveloped in its thickened layer, of muscular fibres. Fig. 183. Grneratite OunAKB OF HiTMAM Fehale. — a^ a. Ovaries. &, b. Fallbpiaa tabes. c. Body of uterus, d. Cervix, e. Vagiua. The cavity of the body of the uterus terminates below by a con- stricted portion termed the os internum, by which it is separated from the cavity of the cervix. These two cavities are not only different from each other in shape, but differ also in the structure of their mucous membrane and the functions which it is destined to perform. The mucous membrane of the body of the uterus in its usual condition is smooth and rosy in color, and closely adherent to the subjacent muscular tissue. It consists of minute tubular follicles somawhat similar to those of the gastric mucous membrane, ranged side by side, and opening by distinct orifices upon its free surface. The secretion of these follicles is destined for the nutrition of the embryo during the earlier periods of its formation. The internal surface of the neck of the uterus, on the other hand, is raised in prominent ridges which are arranged usually in two lateral sets, diverging from a central longitudina;! ridge ; presenting the appearance known as the " arbor vitas uterina." The follicles of this part of the uterine mucous membrane are different in struc- ture from those of the foregoing. They are of a globular- or sac- EGG AND FEMALE ORGANS OF GENERATION. 555 like form, and secrete a very firm, adhesive, transparent mucus, which is destined to block up the cavity of the cervix during ges- tation, and guard against the accidental displacement of the egg. Some of these follicles are frequently distended with their secretion, and project, as small, hard, rounded eminences, from the surface of the mucous membrane. In this condition they are sometimes designated by the name of " ovula Nabothi," owing to their having been formerly mistaken for eggs, or ovules. The cavity of the cervix uteri is terminated below by a second constriction, the "os externum." Below this comes the vagina, which constitutes the last division of the female generative pas- The accessory female organs of generation consist therefore of ducts or tubes, by means of which the egg is conveyed from within outward. These ducts vary in the degree and complication of their development, according to the importance of the task assigned to them. In the lower orders, they serve merely to convey the egg rapidly to the exterior, and to supply it more or less abundantly with an albuminous secretion. In the higher classes and in the human subject, they are adapted to the more important function of retaining the egg during the period of gestation, and of providing during the same time for the nourishment of the young embryo. 556 MALE ORGANS OF GENEBATION. CHAPTER IV. ON THE SPERMATIC FLUID, AND THE MALE ORGANS OF GENERATION. The mature egg is not by itself capable of being developed into the 'embryo. If simply discharged from the ovary and carried through the oviducts toward the exterior, it soon dies and is de- composed, like any other portion of the body separated from its natural connections. It is only when fecundated by the spermatic fluid of the male, that it is stimulated to continued development, and becomes capable of a more complete organization. The product of the male generative organs consists of a colorless, somewhat viscid, and albuminous fluid, containing an innumerable quantity of minute filamentous bodies, termed spermatozoa. The name spermatozoa has been given to these bodies, on account of their exhibiting under the microscope a very active and continu- ous movemetit, bearing some resemblance to that of certain animal- cules. The spermatozoa of the human subject (Fig. 184, a) are about gjj of an inch in length, according to the 'measurements of Kol liker. Their anterior extremity presents a somewhat flattened, triangular-shaped enlargement, termed the " head." The head con stitutes about one-tenth part the entire length of the spermatO' zoon. The remaining portion is a very slender filamentous prO' longation, termed the "tail," which tapers gradually backward, becoming so exceedingly delicate towards its extremity, that it is difficult to be seen except when in motion.- There is no further organization or internal structure to be detected in any part of the spermatozoon ; and the whole appears to consist, so far as can be seen by the microscope, of a completely homogeneous, tolerably firm, albuminoid substance. The terms head and tail, therefore, as justly remarked by Bergmann and Leuckart,' are not used, when describing the different parts of the spermatozoon, in the same sense as that in which they would be applied to the corre- ' Vergleichende Physiologie. Stuttgart, 1852. MALE ORGANS OP GENERATION. 557 sponding parts of. an animal, but simply for tte sake of conveni- ence ; just as one might speak of the head of ap arrow, or the tail of a comet. In the lower animals, the spermatozoa have usually the same general form as in the human subject ; that is, they are slender filamentous bodies, with the anterior extremity more or less en- larged. In the rabbit they have a head which is roundish and flattened in shape, somewhat resembling thei globules of the blood. In the rat (Fig. 184, b) they are much larger than in man, measur- ing nearly xiu of an inch in length. The head is conical in shape, Fig. 184. Spkbmatozo*.— a. Human, ft. Of Rat. c. Of Menobranclins. MagniJed 4S0 times. about one-twentieth the whole length of the filamgnt, and often slightly curved at its anterior extremity. In the frog and in rep- tiles generally, the spermatozoa are longer than in quadrupeds. In the Menobranchus, or great American water-lizard, they are of very unusual size (Fig. 184, c), measuring not less than ,'j of an inch in length, abo»t one-third of which is occupied by the head, or enlarged portion of the filament. The most remarkably peculiarity of the spermatozoa is their 558 MALE ORGANS OP GENERATION. very singular and active movement, to whicli we have already alluded. If a drop of fresli 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, &o.^ 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 move- ment, by which the spermatozoon is driven from place to place in the spermatic fluid, just as the fish or the tadpole is propelled through the water. In other instances, as for example in the water- lizard, and in some parasitic animals, the spermatozoa have a con- tinuous writhing or spiral-like movement, which presents a very peculiar and elegant appearance when lairge 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 iaidestined 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 spermatozoa is a characteristic property belonging to them, arhich continues for a certain time, even after they have been separated from the rest of the body. MALE OKGANS OF GENEBATIOK. 559 In order to preserve tbeir vitality, the spermatozoa must be tept 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'the case of birds and quadrupeds, or if it be subjeofied 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, which are characteristic of the male, as the qva- ries 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 tha 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 sper- matozoa 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 ^ the liquefac- tion of the vesicle, and then fill nearly the entire cavity of the seminiferous ducts, mingled only with a very minute quantity of transparent fluid. In the seminiferous tubes themselves, the spermatozoa are al- ways inclosed in the interior of their parent vesicles ; they are libe- rated, and mingled promiscuously together, only after entering the rete testis and the head of the epididymis. MALE OBGANS OF GENERATION. 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 various secre- tions which assist in the accomplishment of its functions. As the gperm 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 efierentia, 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 reaches 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 depo- sited in the vesiculae seminales, where it accumulates, as fresh quan- tities are produced in the testicle and conveyed downward by the spermatic duct. It is probable that a second secretion is supplied also by the inj;ernal surface of the vesiculse 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, apd 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 deferejis,_the prostate, Cowper's glands, and the mucous fol- licles of the urethra. Of all these ingredients, it is the spermatozoa 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 suc- cession of filters) so as to separate the spermatozoa from the liquid portions, the filtered fluid is destitute of any fecundating properties; MALM ORGANS OF GENBKATION. 561 ■vrhile tte 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,testi- cles 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 accomplishment 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 enlarge 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 Ijecome increased in vascularity and ready to perform their part in the reproductive function. In the fish, for example, where the testicles occupy the same position in the abdomen as the ovaries in the opposite sex, these bodies enlarge, become distended with their contdhts, 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 " erethism." The female, distended with eggs, feels the' impulse which leads to their expulsion; while the male, bearing the weight of the enlarged testicles and the accumu- lation 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 protection and development of the young ; after which the male, apparently 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 be altogether dependent on the occurrence of fortuitous circum- stances, and their impregnation, therefore, often liable to fail. In point of fact, however, the simultaneous functional excitement of 36 562 MALE ORGANS OF GENKBATION. • 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- pregnated 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 t]je 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 congress, 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 and 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. PEBIODICAL OVULATION. 563 CHAPTER V. ON PERIODICAL OVULATION, AND THE FUNCTION OF MBNSTBUATIO I. PERIODICAL OVULATION. We Lave 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. Sggs exist originally in the ovaries, of all animals, as part of their natural structure. In describing the ovaries of fish and reptiles we have said that they consist of nothing more than Graafian vesi- cles, each vesicle containing an egg, and united with the others 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 egg ; and it is from this egg that the young offspring is afterward produced. The process of reproduction was formerly regarded as essentially different in the.oviparous and the 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 bring forth their young alive, such as the quadrupeds and the human species, the embryo was supposed tQ 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, 564 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 is 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 OAdparous, the young is produced equally from an egg ; and in all classes the egg, some- times 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 accordingly, 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 bom 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. These eggs hecome more fully developed at a certain age, when, 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 young 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 ^;eadily 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. 565 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 the 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 fa-ct 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 566 OVULATION AND FUNCTION OF MENSTKUATION. chicks. In oviparous animals, therefore, the discharge of the egg, as well as its forniation, is independent of sexual intercourse. Continued observation shows this to be the case, also, in the viviparous quadrupeds. The researches of Bisohoff, Pouchet, and Coste have demonstated 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 ovaries will show that Grraafian 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- stances 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 plape 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. 180), we see at a glance how the eggs grow and ripen, one after the 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 tihie, eggs which are almost microscopic in size, colorless, and transparent; those which are larger, firtner, ' 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 the 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 discharged. ' Memoire sut la cTiute pgriodiqne de I'oeuf, &o., Annales des Sciences Natnrelles, Aoiit— Septembre, 1844. PERIODICAL OVULATION. 567 In the human species and the quadrupeds, on the other hand, the microscopic egg never becomes "large enough to distend the follicle by its own size. The rupture of the follicle and the libeua- 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 Grraafian 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 proligerus. 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 568 OVULATION AND FUNCTION OF MENSTRUATION, accordingly always situated at the most superficial portion of the follicle, and advances in this way toward the surface of the ovary. • 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. 185. Graaftan Follicle, near the period of rupture —a. Membraneof the vesicle. 6. MembraD* graaulosa. c. Cavity of foLlicle. d. Egg. e. Peritoneum. /. Tunica albugLuca. g, g. TisEue uf the ovary. to project from the surface of the ovary, still covered by the albu- gineous tunic and the peritoneum. (Fig. 185.) 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 pf ^^' ■'^^" fluctuation can be readily per- ceived on applying the fingers to its surface. Finally, the pro- cess of effusion and distension still. going on, the wall of the vesicle yields at its most promi- nent portion, the contained fluid is driven out with a gush, by the reaction and elasticity of the neighboring ovarian tissues, car- rying with, it the egg, still en- tangled in the cells of the pro- ligerous disk. The rupture of the Graafian vesicle is accompanied, in some instances, by an abundant hemor^ OvAET wtTH Graafian Follicle RnpTHRED: atrt, egg just discharged with a portion of raembi'aua granulosa. PERIODICAL OVULATION. 569 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 no 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., by 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 yoiing ones, a similar number of eggs ripen and are discharged at each period. In the mare, in the cow, and in the human female, where tbere is usually but one foetus at a birth, the eggs are matured singly, and the Graafian vesicles ruptured, one after another, at successive periods of ovulation. 4:th. The ripening and discharge of the egg are accompanied by a peculiar condition of the entire .system, known as the " rutting" condi- tion, or " oestruation." 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 degree of excitement, according to the particular species of ani- mal. 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 neighborilig parts are more particularly. affected, being usually increased in quan- tity and at the same time altered in quality. In the bitch, the vaginal mucous membrane becomes red and tumefied, and pours out an abundant secretion which is often more or less tinged with blood. The secretions acquire also at this time a peculiar odor, which appears 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 bf apes it has been observed that these periods are accompanied not only by a bloody discharge from the vulva, but also by an en- gorgement and infiltration of the neighboring parts, extending even 570 OVULATION AND FUNCTION OP MENSTRUATION. to the skin of the buttocks, the thighs, and the under part of the tail.i The system ai> large is also visibly affected by the process going on in the ovary. In the cow, for example, the approach of an 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 rung 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. MENSTEUATION. In the human female, the return of the period 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, ' Poaohet, Theorie positive de I'ovulation, &c. Paris, 1847, p. 230. MENSTBUATION. 571 accompanied by some disturbance of the general system, and tbe female is then known to have arrived at the period of puberty. Afterwarfi, the bloody discharge just spoken of returns at regular 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 .''lightly 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 572 OVULATION ASD FUNCTION OF MENSTRUATION. rusty tinge, until it finally disappears altogether, and the female returns to her ordinary condition. The menstrual epochs of the human female correspqud with the periods of cestruation 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 begm to take place only at the period of puberty, when the aptitude for impregnation commences. Like them, they recur during the child- bearing period at regular intervals; and are liable to the same interruption by pregnancy and lactation. Finally, their disappear- ance corresponds with the cessation of fertility. The periods of cestruation, 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 difiering only in degree from that which shows itself in many species of animals. , The most complete evidence, hovjever, that the period of: men- struation is in reality 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 ruptured, and in the fourth distended, prominent, and ready to burst. Goste* has met with several of the same kind; Dr. Michel' found a vesicle ruptured and filled with blood in a woman who was executed fpr 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. ' London Philosophical Transactions, 1797, p. 135. 2 Reoherohes sur les Ovaires, Paris, 1840, p. 78. ^ Aunales des Sciences Naturelles, August, 1844. * Histoire da Developpement des Corps Organises, Paris, 1847, vol. i. p. 221. * Am. Journ. Med. Sci., July, 1848. " Corpus Luteum of Menstruation and Pregnancy, in Transactions of American Medical Association, Philadelphia, 1851. MENSTKUATION. 573 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 ■whole of the generatiye 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 Bischoff, Pouchet, and Eaciborski, 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 apparently take place 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 c6nveyed downward by the peristaltic action of the muscular coat of this canal.. In the higher classes, on the contrary, the egg is micto- scopio in size, and would be liable to be lost, were there not some further provision for its safety. The wide extremity of the Fallo- pian 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 converging stream, or vortex, by which the egg is neces- sarily drawn toward the narrow portion of the tube, and subse-, qUently conducted to the cavity of the uterus. ' Log oit. 574 OVULATION AND FUNCTION OF MENSTEOATION. Accidental causes, however, someijiimes disturb this regular course or passage of the egg. The egg may be arrested, for example, at the surface of the ovary, and so fail to enter the tube at aU. 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 coii\cidence 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 different individuals. The above facts indicate-also the true explanation of certain , exceptional oases, which have sometimes been observed, in which fertility exists without menstruation. Various authors (Churchill, Eeid, Velpeau, &c.) have related instances of fruitful women in whom MENSTRUATION. 575 the menses were very scanty and irregular, or even entirely absent. The menstrual flow is, in fact, only the external sign and accompa- niment of a more important process taking place within. It is habitually 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 maj 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 haemoptysis, 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. 576 MEXSTRUATIOIf AST) PBEGNANCT. CHAPTEE VI. ON THE CORPUS LUTEUM OF MENSTBUATION AND PREGNANCY. After the rupture of the Graafian follicle at the menstrual period, a bloody cavity is left in the ovary, which is subsequently oblijierated by a kind of granulating process, somewhat similar in character to the healing of an abscess. For the Graafian follicle is intended simply for the formation and growth of^ the egg. Aft^r 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 follicle 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 follicle, at each menstrual epoch, swells, protrudes front the surface of the ovary, and at last ruptures apd 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 extra vasated in any other part of the body, and the coagulum is retained in the interior of the Graafian follicle. COKPUS LUTEUM OF MENSTRUATION. 57? The opening by which the egg 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 Fig- 187. be opened at this time by a longitudinal inci- sion (Fig. 187), 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 edges of the lacerated \ " opening, just as the coagulum in a recently 2i:,;'lZ,Tlll" ligatured artery is entangled by the divided menstroation, and aued edges of the internal and middle coats ^ but rhot^"i°^Jit°3'rc: there is no continuity of substance between "on.— «. Tissue of the . 111 1 1 i>i ovary. ■&. Kembrane of the them, and the clot may be everywhere readily veaicie. o-. pomt of rupture, separated by careful manipulation. The mem- brane 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 firsl; 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 psCrtially 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 be'comes thickened and convoluted, and tends partially to fill up 37 678 MENSTRUATION AND PKITGNANCy. Fig. 188. the cavity of the follicle. This hypertrophy and convolution of the membrane just named commences and proceeds most rapidly at the deeper part of the follicle, directly opposite the situation of the superficial rupture. From this point the membrane 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 on 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. 188), 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- otaky cut open, «ho,viuif corpus volutcd Wall is about oue-cighth of luteum divided i^ngiindinaiiy ; tiiree an iuch thick at its dccpest Dart, and veeka after menstrnatioa. From a girl .,„. n-i aead of haemoptysis. 01 an mdetinite yellowish or rosy hue, not very different in tinge from 'the rest of the ovarian tissue. The convoluted wall aiid the con- 'tained clot lie simply in contact with each other, as at firsty without any intervening membrane or other organic connection; and they may still be readily separated from each other by the hamdle 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 COBPUS LUTEUM OF MENSTRUATION. 579 Otart, sbowlDg corpas luteum four weekH after men- straation ; from a woman de&d of apoplexy. ( the areolar tissue, by whichit was previously connected with the substance of the ovary. After the third week from the close of menstruation, the corpus luteum passes into a retrogr'ade condition. It diminishes.percep- tibly in size, and the central coagulum con- tinues 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. 189.) The external cicatrix may still usually be seen, as well as. the point where the central coagulum comes in contact with the peritoneuni. There is still no organic connection between the central coagulum and the convoluted wall ; biit 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 effected 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 th# 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 as- sumes a brighter and more decided yellow. This change of color in the convoluted wall is produced in consequence a^ 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 „„.., .v„„,„„ , •' ' M/ OvABT, sDowiav corpus 111- which it is readily distinguished from alk tenm, Binn weeks atjer meiBlma. the neighboring tissues. After this period, the process of atrophy and degeneration goes on rapidly. The clot becomes constantly Fig. 190. tinn ; frnm a girl dead of tuber- cular ineniugitia. 580 MENSTEQATION AND PREGNANCY. more dense and shrivelled, and is soon converted into a minute, titellate, white, or reddish-white cicatrix. The yellow wall becomes softer and more friable, as is the case with all tissues undergoing fatty degeneration, and shows less distinctly the marking of its convolutions. At the same time its surfaces become confounded with the central coagulum on the one hand, and with the neigh- boring tissues on the other, so that it is no longer possible to separate thetn fairly from each other. At ther end of eight or nine weeks (Fig. 190) the whole body is reduced to the condition of an insignifi- cant; 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 peculiar 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 some- times elapses before its final and complete disappearance. 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 recurred 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 ^sappear 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 PEEGNANCY. 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 takes place, the appearance of the corpus luteum becomes so much altered as to be readily dis- CORPUS LUTEUM OP PREGNANCY. 581 tinguished from that which simply follows the ordinary menstrual 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 menstruation passes rapidly through its different stages, and is very soon reduced to a condition of- atrophy, that of pregnancy continues its develop- ment for a long time, attains a larger size and firmer organization, 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 is capricious, and even the mental faculties and the moral disposition are frequently more or less affected. The ovaries, however, feel the disturbing influence 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 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 increased in size to such an extent as to measure seven-eighths of an inch in length by half an inch in depth. (Fig. 191.) The central coagulum has by this time become almost entirely decolorized, so as 582 MENSTRUATION AND PREGNANCY, Corpus Lptbum of pregnancy, at end of pecond month ; from a woman dead from induced abortion. 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. 191, forming a little cavity containing" a few Fig- 191- drops of clear fluid and in- closed by ar whitish, fibrinous layer, the remains of the solid portion of the clot. It is this fibrinous layer which has sometimes been mistaken for a distinct organized mem- brane, lining the internal sur- face of the convoluted wall, and which has thus led to the belief that the yellow matter of the corpus luteum is normally deposited outside the mem- brane of the Graafian follicle. Such, however, is not its real struc- ture. The convoluted wall of the corpus luteum is the mem])rane of the follicle itself, partially altered by hypertrophy, 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 oc- casional, not a constant pheno- menon. More frequently, the fibrinous clot is solid through- out, the serum being gradually absorbed, as it separates spon- taneously from the coagulum. During the third and fourth 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. 192.) The con- voluted wall is still thicker and more highly developed than before, having a thickness, at its deepest part, of three-sixteenths of an inch. Its color, however, has already begun to fade, and is now of a dull yellow, instead of the bright, clear tinge which it previously ex- Fig. 192. Corpus Lutbcm of pfegnaiicy, at end of fourth month ; from a woman dead hf polBou. CORPUS LUTEUM OF PBEGNANCY. 583 Fig. 193. Hbited. The central coagulum, perfectly colorless and fibrinous in appearance, is often so much flattened by the lateral compres- sion 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 convolutions. The central coagulum is reduced, by this time to the condition of a whitish, radiated cicatrix. The atrophy of the organ continues dur- ing the ninth month. At the termination of pregnancy, it is redticed to the size of half an inch in length and three-eighths of an inch in depth. (Fig. 193.) It is then of a faint indefinite hue, but little contrasted with the remaining tissues of the ovary. The central cicatrix has become very small, and appears only as a thin whitish lamina, with radiating processes which run in be- tween the interstices 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 tex ture, as a prominent feature in the ovarian tissue, and a reliable indication of pregnancy. The convoluted 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 traces may remain, however, for a long time afterward, more or less GoRPUB Luteum of preg- nancy, at term ; from a woman dead in delivery from rupture of the uterus. 584: MENSTRUATION AND PREGNANCY. concealed in tlie 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 are 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 corpora 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 forrtier passes through all the im- portant phases of its growth and decline in the period of two nionths, the latter lasts 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 import- ant a characteristic, is only temporary in its duration ; not making its appearance till about the end of the fourth week, and again 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. COEPUS LUTEUM OF rREGNANOY. 585 At the end of three weeks One month Two months Six months Nine months CoRins LnTECM of Menstedatiox. Coefus Luteum of Peerxascy. Three-qnarters of an inch in diameter ; central clot reddish ; con- voluted wall pale. Smaller ; convoluted wall bright ' Larger ; convoluted wall bright yellow ; clot still reddish. yv How ; clot still reddish. Reduced to the condition of an "Seven-eighths of an inch in dia- insignificant oicatilz. Absent. Absent. 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 con vo luted, but without any bright yellow color. 586 DEVELOPMENT OF THE IMPREGNATED EGG. CHAPTEE VII. ON THE DEVELOPMENT OF THE IMPREGNATED EGG— SEGMENTATION OP. THE VITELLUS— BLAS- TODERMIC MEMBRANE— FORMATION OF ORGANS IN THE FROG., We have seen, in the foregoing chapters, how the egg, produced in the ovarian folliclei, 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 that which marks its complete maturity, is the disappearance 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 is thereby fecundated. By the influence of fecundation, 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 SEGMEKTATIOX OF THE VlTEl.LUS. 587' Fig. 194. 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 segmentation, 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. 194, a.) Almost at the same time another furrow, run- ning 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. 194, h), the edges and aingles of which are rounded off, and which are still con- tained in the cavity of the vitelline membrane. The spaces between them and the internal surface of the vitelline membrane are occupied 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. 194, c.) These vitelline spheres have a somewhat firmer consistency than the original substance of the vitellus ; SEQMB.vTATioir of the vitblujb. 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 polygonal forms, and flattened 588 DEVELOPMEXT OF THE IMPKEGNATED EGG. against the internal surface of the vitelline membrane. (Fig 194, d.) They have by this time been converted 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 membrane. 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 kre rather larger arid looser in texture; The egg then 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 tjike 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 blastoderpiic membrane is, in fact, the body of the foetus. It is at this time, it is true, exceedingly simple BLASTODEKMIC MEMBRANE. 589 in texture ; but we shall see hereafter that all the future organs! 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 classes of animals. It is always in this way that 'the egg commences its development, whether it be destined to form a;fterward a fish or a reptile, a bird, a quadruped, or a man. The peculiarities belonging to difierent species show themselves afterward, by variations in the manner and extent of the develop- ment of different parts. In the higher animals and in the human subject the development of the egg becomes an exceedingly com- plicated 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 appro- priate nutritious material. We find accordingly that under these circumstances the development of the egg is distinguished by a character of great simplicity; since the wfiote of the vitellus is diivctly converted into the body of the embryo. There are no acces- sory 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 : I7ie 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. . 590 DEVELOPMENT OP THE IMPREGNATED EGG. The first sign of advancing organization in the external layer of 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. 195), the wide edges of which are somewhat Fig. 195. more opaque than the rest of the blasto- dermic niembrahe. 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, IMPREGNATED E«a. ,,uh Com- 1° t^^ atca pellucida, the substance of mencement of formation of embryo:, the blastodermic membrane piscs UD in showing embryonic Rpot, area pella- ., . n , i oi This exhalation is not a simple effect 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 incubated, though they may be freely exposed to the air. The exhalation 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 jJrevent 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 Dgvelopperaent du Foetus. .Paris, 1850, p. 143. 606 AMMONAND ALLANTOIS. teen days' incubation, the egg absorbs 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 amounts 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 jnembranes 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 br.eak 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 furnishes the Materials which are used to accomplish its own demo- lition, and at the same time to eflect 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- t » Pp. cjt., pp. 138 and 149. DKVELOPMENT OF THE CHICK, 607 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 vsrhen the chick is completely formed, and has become capable of carrying on an in- dependent fexistence, 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. 608 DEVELOPMENT OF THE EGG IN HUMAN SPECIES. CHAPTEE X. DEVELOPMENT OP THE EGG IN THE HUMAN SPECIES.— POBMATION. OP THE CHORION. We have already described, in a preceding chapter, the manner in Tvhich 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 siurface of ^^" 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- eS into a simple vascuh^r memhra'ne. (Fig. 213.) This membrane, moreover, being, after a time, thoroughly fused and united with the two Tyhich have preceded it, takes the place which was previously Hdman Ovitm, about the end of the first month ; showing formation of chorion. — 1. Umbilical vesicle, 2, Amnion, 3, Chorion, FORMATION OF THB CHOKION. 609 occupied by them. It is then termed the chorion, and thus becomes the sole external investing membrane 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 membranes, as follows: first, the Original vitelline niembrane; secondly, the outer lamina of the amniotic fold ; and, thirdly, the allantois; the last predominating over the two formet 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, jn respect to its mode of formation, is the same thing with the allantois of the lower animals ; its chief peculiarity consisting in the fact that 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. 213), 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 repeate* budding and sprouting of lateral ofishoots from every part. After this process of growth 39 610 DEVELOPMENT OF THE EGG IN HUMAN SPECIES. Fig. 214. has gone on for some time, the external surface of the chorion pre- sents a uniformly velvety or shaggy appearance, owing to its heing covered everywhere with these tufted and compound villosities. The villosities themselves, when examined by the microscope; have an exceedingly well-marked' and characteristic appearance. (Fig. 214.) They originate from the surface of the chorion by a somewhat narrow stem, and divide into a multitude of secondary and tertiary branches, of varjdng size and figure; some of them slender and filamentous, others club-shaped, many of them irfegularly 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 the villosity are seen to contain nu- 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. Whenever we find, in the uterus, any portion of a inembrane 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 he 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 Compound villosity of Human Cho- BIOK, ramified extremity. From a three mouths' foetas. Magnified SO diameters. FOBMATION OF THE CHORION, 611 Fig. 215. by vascular absorption, instead of the slow process of imbibition, whicb has heretofore taken place through the comparatively incom- plete 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. 215), like the vessels in the papillae of the skin, and retrace their course, to unite finally with the venous trunks of the chorion. The vUli of the chorion are, therefore. Extremity VILLOSITY nr very analogous in structure t8 those of chokion, more lighiy magni- .■,*,,' -Ill' n 1 fled; showiug the arrangement of the intestine ; and their power of absorp- bioodyes-eis m its interior. tion, as in other similar instances, corre- sponds 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 bloodv'essels 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 lamina? 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 612 DEVELOPMENT OF THE EGG IN HnjIAN SPECIES. completely separated from the inner, by the disappearance of the partition which for a. time connected the two with each other (Fig. 209, c), this source of vascular supply is cut off; and the second chorion 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 than 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. Th^ villosities of 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 he- 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 adja- ^'8" ^^^' cent 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 corresponds with the insertion of the foetal vessels, the villosi- ties, instead of becoming atro- phied, continue to grow; and this portion of the chorion be- comes even more shaggy and thickly set than before. The consequence ip that the chorion afterward presents a very different appearance at different portions of its surface. (Fig. 216.) The greater part of it is smooth-; but a certain portion, constituting about one-third of the whole, is covered with a soft and spongy mass of long, thickly-set, compound villosities. It is'this thickened HrMAH Ovum lit end of third month placental portion of the chorion fully formed. showing .FOBMATION OF THE CHORION. 613 and staggy portion, whicli 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 ^n extent corresponding 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 o'f the membrane they are merely distributed as a few single and scatteted vessels. The chorion, accordingly, is the external investing membrane of the egg, 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. 614 DEVELOPMENT OF UTERINE MUCOUS MEMBRANE. CHAPTER XI. DEVELOPMENT OF UTEEINB MUCOUS MEMBRANE- FORMATION OF THE DECIDUA. In fish, reptiles, and birds, the 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 sufl&cient 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 hypertrophjed, 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 h'tere,'^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 welt marked and readily distinguishable FOKMATIOX OP THK DECIDUA. 615 Fig. 217. iiii Uterine Muoors Mehbeane, ag >ieen in vertical Kection, — a. Free surface. b. Attached tmrftux. from that of other parts. It consists, throughout, of minute tubular follicles, ranged side by si3e, and running perpendicularly to the free surface of the mucous membrane. (Fig. 217.) Near this free surface, they are nearly straight; but toward the deeper sur- face of the mucous membrane, where they terminate in blind extremities, they become more or less wavy or spiral in their course. The tubules are about yjj of an inch in diameter, tad are lined throughout with co- lumnar epithelium. (Fig. 218.) 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 '^' 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 increas- ed activity of growth and an unusual development. It becomes tumefied and vascular ; and, as it increases in thickness, it projects, in rounded eminences or convolutions, into the uterine cavity. (Fig. 219.) 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 eye upon the uterine surface, as numerous minute perforations. The bloodvessels of the mucous membrane also enlarge and multiply, and inosculate freely with Utkrtne Tdbut.rs, from mucous membrane of nuimpre^nated hnmau uterus. 616 DEVKLOPJIENT OF UTKBINE MUCOUS MEMBBANB, eact other ; so that the vascular network encircliug 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 fexudation 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 mucousf 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 Decidva 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, meanwhile, begins to be filled with an abundant secretion of its peculiarly visqid mucus, which blocks Tip, 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. 219.) 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. 219.) It is at this situation that an adhesion subsequently takes 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 FOBMATION 0¥ THE DECIDUA. 617 mucous membrane. Its projecting folds begin to grow up around the egg in such a manner as to partially inclose if in a kind of circumvallation of the decidua, and to shut it off, more or less com- Fig. 219. Fig. 220. Ikprbgnated Uterus; nhowingr formation of decidua, Tlie decidua Is represented in black ; and the egg is seen, at tlie fundus of the uterus, eu- gAged between two of its projecting convolutions. IsipaKaNATKD Utkbur, with pro- Jpctiug folds of decidua growing up around the egg. The narrow opening where the edges of the folds approach each otherf^ls seen over the must pi^omi- neut portion of the egig. Fig. 221. pletely, from the general cavity of the uterus. (Fig. 220.) 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 pro- cess of growth goes on, this opening be- comes narrower and narrower, w.hile the projecting folds of decidua approach each other over the surface 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 (Eig. 221), 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 for- mation, which has thus gradually enveloped it, and by which it is concealed from view when the uterine cavity is laid open. This iMPRIiUMATKiJ UtKRI.'s; — showing egg completely inclosed hj decidua reflexa. 618 DEVELOPMENT OF UTEBIKE MUCOUS MEMBKANE, newly-formed layer of decidua, enveloping, as above described, the projecting portion of the egg, is called the Decidua reflexn; because it is reflected over the egg, by a continuous growth from the general surface of the uteri ne mucous membrane. The orifices of the uterine tubules, accordingly, in consequence of the manner ia which the decidua reflexa is formed,- will be seen not only on its external sur- face, 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 origihal 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 eontinued to increase in thickness and vascularity. The remainder of the decidua vera, however, ceases to grow as rapidly as before, and no longer keeps pace with the increasing size of the egg and of the uterus. It is still very thick and vascu- lar 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 concentrated in the egg, and in that portion of the uterine mucous membrane 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 beien taking, place in the walls of the uterus, the FORMATION OP THB DECIBUA. 619 formation of the embryo in the egg, and the development of the amnion and chorion have been going on simultaneously. Soon after the entrance of the egg into the uterine cavity, its external investing membrane becomes coviered with projecting filaments, or villosities, as previously described. (Ghap. 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 foUiclea of the uterine mucous membrane, formed either from the cavities of the tubules themselves, or by the adjacent surfaces of minute projecting folds. When the formation of the decidua reflexa is accomplished, the chorion has already become uniformlv 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 surfece of the decidua reflexa with which it is covered. (Fig. 222.) In this way the egg becomes entangled with the decidua, apd cannot then be read- ily 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- ..J^r/clVer. Jrervn! tritious fluids, exuded from the soft and '"*'"«' "' "'""-ion ana decidna'. 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 necessairy ; 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. Bach villosity, then, as it lies imbedded in its uterine follicle, contains a vascular loop through wtich 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, 620 DEVELOPMENT OF UTEBIKB MUCOUS MEMBRANE. Fig. 223. disappear over a portion of its surface, while they at the same time become concentrated and still further developed at a particular spot, the situation of the future placenta. (Pig. 223.) 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 vascjularity ; while else- where, over the prominent portion of the dgg, 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 takipg 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. PBEavANT Utbrus; showing formation of placenta, hj ttie united deveiopment of a portion of the de- cidua and the villosities of' the cho- THE PLACENTA. 621 CHAPTER XII. THE PLACENTA. Wk 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 w:ay 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 arid 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 valvular 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. 224.) 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 parts. In other instances, however, the development of the foetus requires a more elaborate arrangement of the vascular membranes. 622 THE PfiACENTA. In the cow, for example, the external membrane of the egg, beside being everywhere supplied with branching vessels, presents upon Fig. 224. F(ETAL Fig, with \ta membraaes, contained in cavi^t/ of atera8.-*a, a, h^ b. Walls of oterns. c, c. Cavity of uterus, d. Amnion, e, e. AUantoia. 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 in thick. Velvety, vascular masses. 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 adhierent to it, is known by the Fig. 225. CoTTLRDON OF C w' 8 Ut E B I' 9. — a, a. Surface of fcetal cliorion. ft, &. Blood venaela of foetal chorion, d^ d. Bloodvessels of uterine macons membraae. c, c. Surface of aterine macoas mem- braae. name of the Cotyledon, Each cotyledon forms/ therefore, a little placenta. (Fig. 225.) In its substance tbe tufted vascular loops coming from the uterine mucous membrane {d, d) are entangled TUB PLACENTA. 623 with those coming from the membranes of the foetus (6, b). There is no absolute adhesion between the two sets of vessels, but only an interlacement pf their ramified extremities; and, with a little care in manipulation,. the fcetal 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 fcetal 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 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 tl* 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 fcetal 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 rapidly' elongate 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 multi- plied 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, also become unusually developed. They enlarge and inosculate freely with each other; so that every uterine follicle is soon covered 624 THE PLACENTA. 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 &l\ 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 fcetal vessel 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 up^ 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 ^'g- 226. sinuses, which communicate freely with the enlarged vessels in the muscular walls of the uterus. As the original capil- lary plexus occupied the entire thickness of the hypertrophied decidua, the vascular sinuses, into which it is thus converted, are equally extensive. They commence at the inferior sur- face 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 fcetal 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 blood- vessels, both in the foetal and maternal portions of the placenta, is so excessive that all the other tissues, which originally co-existed Extremity of F inereases in abundanceji and the molecules continue to enlarge until they become converted into fully formed oil-globules, which fill the interior of the fibre or less completely. Fig. 234. more and njask, to a certain ex- tent, its anatomical cha- racters. (Fig. 234.) The universal fatty degenera- tion, thus induced, .gives to •• Jihe uterus a softer consist- ency, and a pale yellpwish color which is characteristic .of it at this period. The muscular fibres which have become altered by the fatty deposit are afterward gra- dually absorbed and disap- pear;, their place being subsequently taken by other fibres of new forma- tion, which already begin 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. MUBCDLXB riBBES OF HuMAH Utebdb, three weeks after parti^ltion ; from a woman dead of peri- tonitis. Op. cit. DEVELOPMENT OS THE EUBSYO. 637 CHAPTER XIV;. DEVELOPMENT OP THE BMBBYO— NEEV OUS SYSTEM. ORGANS OF SENSE, SKELETON, AND LIMBS. Fig.' 235. 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 aboye described, on each side the median furrow. The two laminae of which this is com- posed, on the right and left sides (Fig. 235, 3,i>), unite with each other in front, forming a rounded dilatation (c), the cephalic extremity, and behind at rf, forming a pointed or caudal extremity. Near the pos- terior extremity, there is a smaller dilatation, which marks the future situation of the lumbar enlargement of the spinal cord. .A.S the laminse 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 cwm, 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 anteripr bulbous enlargement into three secondary compartments or vesicles (Fig. 236), which are partially separated from each other by transverse constrictions. These vesicles are known as the three cerebral vesi- cles, from which all the d liferent 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 tuberoula quadrigemina ; and the third, or posterior, the medulla •oblongata. All three vesicles are at this time hollow, and their Formajtion of Crbe- bho-Spisal Axis.— n, b. Spinal cord. e. Ce- phalic extremity. d. Caudal extremity. 638 DEVELOPMENT OP THE EMBBTO. cavities communicate freely witli eacli other, through the interven- ing constrictions. Very soon the anterior and the posterior cerebral vesicles suflfer 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 intottwo portions, of which the ante- rior becomes the cerebellum, and the pos- terior the mediiia 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. 237); 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 ipry 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 of the parts begin to alter. Tiie hemispheres and tubercula quadrigemina grow faster than the poste- rior portions of the encephalon ; and the cerebellum becomes doubled backward over the n^dulla oblon- gata. (Fig. 238.) Subsequently, the hemispheres rapidly enlarge^ growing upward and backward, so as to cover in and conceal both the optic tha- lami and the tubercula quadrigemina ^Fig. 289); the cerebellum tending in the same way to grow backward, and projecting farther Formation of the Gsbebbo- SpiwAL Axi^. — 1. Vesicle of tiie liemi-pherpR. 2. Vesicle of tlie tubercula quadrii^emiDa. 3. Vesicle oflhe medulla oblonjfata. Fig. 237. ^ -^ 4 FfflTAL Pia, five- eighths of an inch long, showing brain and spioal cord. — 1. Hemiapheres. 2. Tu- bercala quadrigemi- na. 3. CerebeUnm. 4. Medulla oblongata. NEKVOUS SYSTEM. 639 and farther over the medulla oblongata. The subsequent higtory of the development of the encephalon is little more than a con- Fig. 239. F(ETAi. Pig, one and a. quarter inch long. — 1. Heniisiilieres. 2. Tubercula quadrigemina. 3. Cerebellum. 4. Me- dulla oblongata. Head of F(etal Fig, three and a half Inches long. — x. Hemlsptieres. 8. Cerebellam. 4. Medulla oblongaia. tinuation of the same process ; the relative dimensions of the parts constantly changing, so that the hemispheres become, in the adult condition (Fig. 240), the largest of all the divisions of the ence- Fig. 2i0. Brain op Adult Fro. — 1. HemisphereE. 3. Cerebellam. 4. Medu'ua oblongata, phalon, while the cerebellum 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 theii* nervous matter. In the human foetus, these con- volutions begin to appear about the beginning of the fifth month (Longet), and grow constantly 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 r and. by folding over 6i0 DEVKLOPMENT OF THE EMBBYO. agaipst eacli other in the median line, form the right and left hemi- spheres, separated from each other by the longitudinal fissure. A similar process of growth taking place in the 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 t^e-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 vitreovs 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 avery 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 tne 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, subset 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 tlje direction of its receding bloodvessels, viz., from SKELETON AND LIMBS, 641 tlie centre toward the circumference. It ias completely disappeared by the end of the seventh month. (Cruveilhier.) 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 contac^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 offshoot. 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. VII.), 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 in 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 may even close spontaneously ; or it may be complicated with an imperfect deve- 41 642 DEVELOPMKNT OF THE EMBRYO. lopment or complete absence of the spinal cord at the same spot, 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 latter 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 articular 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 fcetal 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, projects at this time considerably beyond the pelvis, forming a tail, like that of the lower animals^ which is curled forward toward the SKELETON AND LIMBS. 643 abdomen, and terminates in a pointed extremity. Subsequently the 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 accord- ingly disappears. The integument of 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 ii to pass easily through the generative passages. 6J:J: DEVE1.0PMENT OF THE ALIMENTABY CANAL CHAPTER XV. DETBLOPMBNT OP 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 formation of the intestinal canal ; so that, during' a certain period, the abdomen of the embryo is widely open in front, presenting a long oval excayj^tiOn, in which the nearly straight, intestinal tube is' to be seen, Tunpjng from its anterior to its posterior extremity. The formaticjn ofthe 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 becomes 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 oil 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. 241. The abdominal walls are AND ITS APPENDAGES. 645 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, Tb^ intestine makes at Fig. 241. FormatioQ of Alihrn T AR Y Canal. — a, b. Commencemeiit of amnion, c. e. Intestine, d. Pharynx, e. Urinary bladder. / Allantois, g. Umbilical vesicle, m. Dotted line, showing the place of formation of the (esophagus. 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, (g.) 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, become widely different from each other. The upper portion, which is the small intestine, grows mostly in the direction of its length, and becomes a very long, narrow, and convoluted 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 olsliquely into the side of the colon. This diverticulum of the colon is at first uniformly 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 appendix vermiformis ; while the remaining portion, which is con- tinuous with the intestine, becomes exceedingly enlarged, and' forms the caput cqli. The caput coli and the appendix are at first situated near the 646 DEVELOPMENT OP THE ALIMENTARY CANAL Fig. 242. umbilicus ; but between the fourth and fifth .moilths (Cruveilhier) their position is altered, and they then become fixed in the right iliac re^on. 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 sacculi 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 in- testinal loop, or hernia, from the umbili- cal opening. (Fig. 242.) In the human embryo, this protrusion of the intestine can be readily seen during the latter part of the second month. At a subsequent period, however, the walls of the abdo- men grow more rapidly than the intes- tine. They accordingly gradually' en- velop the hernial protrusion, and at last inclose it again in the cavity of the ab- domen. 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 sucli 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 allaniois. (Fig. 241, /.) It is at FtETAL Fro, showing lobp of intestine, forming umbilical hernia ; from n specimen in the anthor'a, possession. From the coDTexity of the loop athinfilament is seen passing to the umbilical vesicle, which is here flattened into a leaf-like form. AND ITS APPENDAGES. 6i7 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 cAoriow. 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 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. 241, e), becomes the wn- nary hladder ; 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 above 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 between 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- 6J:8 DEVELOPMENT OF TUK ALIMENTAET CAITAL 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 the biliary substances (tauro-cholates and glyko-cholates) when care- fully 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 young foetuses, about •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 foetal 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. According to Kdl- liker,^ the soft downy hairs of the foetus, exfoliated from the skin, ' Physiological Chemistry, Philadelphia edition, vol. i. p. 532. = Gewebelehre. Leipzig, 1°52, p. 139. AND ITS APPENDAGES. 64:9 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 foetal pig we have found the relative size of the liver greatest within the first month, when it amounts to very nearly 12 per cent, 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 hile 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. 241, 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- ' Lemons de Physiologie Expgrimentale, Paris, 185.5, p. 398. 650 DEVELOPMENT OP THE ALIMBNTAEY CANAL lish a communication with the intestinal tube. The formation of the anterior perforation, and its appendages, takes place in the fol- lowing mianner : — 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 cephalic mass there is now formed a large cavity (Fig. 241, dj, 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 situatipn 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 extremityof the abdo- men, 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 x, in Fig. 241. 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- ing, 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 epithelium of the oesophagus may be seen to terminate abruptly, by a well- defined line of demarkation, at the cardiac orifice of the stomach ; beyond which, throughout the remainder of the alimentary canal, the epithelium is of the columnar variety, and easily distinguish- able 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 AND ITS APPENDAGES. 651 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 shuts 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 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 abdoniinal organs then partially protrude into the cavity of the chest on that side, forming congenital diaphragmatic hernia. The lung on the afiected 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 seri- ous inconvenience. 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 condition known as ectopia cordis. This malformation is necessarily latal. 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 mentioned. (Fig. 243.) One of them grows directly downward from the frontal region (i), and is called the frontal or intermaxillary process, because it afterward contains in 652 DEVELOPMENT OP THE ALIMENTARY CANAL Pig. 243. Ukad of Hitman Embrto, at about the twentieth day. After Longet ; from' a specimen in the collection of M. Coste. — 1. Frontal or intermaxillary process. 2. Pro- cetis of superior maxilla. 3. Pro- cess of inferior maxilla. its lower extremity the intermaxillary bones, iri which the incisor teeth of the upper jaw are inserted. The next process (2) ori^n- ates from the side of the opening, and, advancing toward the median line, forms, with its fellow of the opposite side, the superior maxilla. The processes of the remaining 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, and at the same time two oif- shoots show themselves upon its sides, which curl round and inclose two circular orifices, the opening of the anterior nares ; the ofishoots themselves becoming the aire nasi. (Fig. 244.) The mouth at this period is very widely open, owing to the imperfect development of the upper and lower jaw, and the incomplete formation of the lips and cheeks. The processes of the superior max- illa 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 intermaxillary pro- cesses which grow down between them. They then unite with the intermaxillary processes, which have at the same time united with each other, and the upper jaw and lip are thus completed. (Fig. 245.) The external edge of the ala nasi also adheres to the superior maxillary process and unites with it, leaving only a curved crease Fin. 244. H IS A D F H [J M A N £ M B K V o at about the sixth week. From a specimen iu the author's possession. AND ITS APPENDAGES. 653 or farrow, as a sort of cicatrix, to mark the line, of union between them. Sometimes the superior maxillary and the intermaxillary pro- cesses fail to unite with each other ; and we then have the mal- formation known, as hare-lip. The fissure of hare-lip, consequently, as a *''S- 245. general rule, is not situated exactly in the median line, but a little to one side of it, on the external edge of the inter- maxillary 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 de- tached from the remainder, contains head or hdmam embrto, abc.nt ,1 n • • J. ii 1 the end of the aecotid month. — From a the four upper mClSOr teeth, and cor- specimen m the a.thor-, poa.e.»loa. responds with the intermaxillary bone of the lower animals. In some rare instances the fissure of hare- lip is situated in the median line, the two intermaxillary bones never having united with each other. A case of this kind has been observed by Prof. Jeffries Wyman.' 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. 243.) As development proceeds, they come t9 be situated farther forward (Fig. 244), 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. 245), 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 Ml 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- ' Transactions Boston Society for Medical Improvement, March 9th, 1863. 654 DEVELOPMENT OP THE ALIMENTARY CANAL, ETC. low. Fissure of tlie palate is caused by a deficiency, more or less complete, of one of the horizontal maxillary plates. It is accord- ingly situated a little on 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 of 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 division^, have been successively formed ; and the development of the upper part of the alimentary canal is then complete, DEVJfiLOPMENT OF THE KIDNEYS. 655 CHAPTER XVI. DETELOPMBNT OP THE KIDNEYS, WOLFFIAN BODIES, AND INTEENAL ORGANS OF GENE- RATION. Fig. 246. 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. These 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. 246), 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 -j'j 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 "Wolffian bodies, in their intimate structure, closely resemble 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 ex- F— About the same time that the kidneys are formed behind the Wolffian bodies, two oval-shaped organs make their appearance in front, on the inner side of the "WnlflWi bodies and between them inches long. From a specimen in , - ." , . rni T T anther's possession, — 1,1. Kidni 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 precisely the same situation and present precisely the same appear ance, whether the foetus is afterward to belong to the male or the female sex. (Fig. 248.) 42 Fig. 248. Internal Organs of Genr- ATION, &c. ; in a fcetal pig three the neys, 2, 2. Wolffian bodies. 3, 3. Internal organs of generation ; testicles or oraries. 4. Urinary bladder turned over In front. 6. Intestine. 658 DEVELOPMENT OP THE KIDNEYS. . A short distance above the internal organs of generation there commences, on each side, a narrow tube or duct, which runs from above downward along the anterior border of the Wolffian body, immediately in front of and parallel with the excretory duct of this (i^rgan. 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, or what will afterward be the base of the urinary bladder. These tubes serve as the excretory ducts of the internal organs of generation ; and will afterward become the vasa deferenUa in the male, and the Fallopian tubes in the female. According to Coste, the vasa defe- rentia 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 tjie same author, that the vasa deferentia become adherent to the testicles, and a communication is established between them and the tubuli semi- 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 O^ans of Gfeneration ; 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 t]ie 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 organs becomes altered. The descent of the testicles is accompanied by certain other alterations in the organs themselves and their | appendages, 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 the^e organs; (Fig. 249.) 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 MALE ORGANS OF GENERATION'. 659 Ijitersai. Oboaks or Gekekatiox, &c., in a foetal pig Dearly four inches long^ Prom a specimen in the author's possessioD. — 1, 1. Kidneys. 2, 2. Wolffian bodies. 3, 3. Testicles. ' 4. Urinary bladder. &. Intestine. direction, crossing the corresponding vas deferens a short distance above its union with its fellow of the opposite side. Below this point, the cord spoken of continues to run obliquely outward and downward; and, passing through the abdominal walls at the situa- ^'8' ^^• tion of the inguinal canal, is in- serted into the subcutaneous tis- sues near the symphysis pubis. The lower part of this cord be- comes the gubernaculum testis ; and muscular 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 become afterward convoluted, and converted into the epididymis. 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 lastj when the testicles have arrived at the internal inguinal ring, the AVolffian 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 arriveat 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 m.uscle. DEVELOPMENT OF THE KIDNEYS. At last, the testicles descend fairly to the bottom of the scrotum the gubemaculum constantly shortening, ' and the vas deferens 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 downwardi, and joins its fellow of the opposite side ; after which they both open into the prostatic portion of the urethra by distinct orifices, situated on each side the median .line. At the satae 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 vesiculse 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 communis cation with the peritoneal cavity. In niany 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 Fig. 250. the abdomen, by the action of the guber- naculum 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 pro- cess, the serous surfaces come actually Formation of Tunica 'va- in coutact with cach Other, and, adhering GinAi,i6 Testis. — 1. Testicle ^ ,, . ,i ■ ., .. ,,-,. nir, \ ■ neaily at the bottom of the scro- togethci at thlS SltUatlOU (Fig. 250, 4), turn. 2 Cavity of tunica Ta,;in» lis. form a kind of cicatrix, or umbilicus, by 3. Cavity of peritoneum. 4. Obliter- ' , t ■ o ated neck of peritoneal sac. the Complete closurc and consolidatiou: of which the cavity of the tunica vaginalis (2) is finally shut off altogether from the general • cavity of the peritoneum (3). The tunica vaginalis testis is, therefore, originally a part of the peritoneum, from which it is subsequently separated by the process just described. FEMALE OBGANS OP GENERATION. 661 The separation of the tunica vaginalis from the peritoneum is usually completed in the human subject before birth. But some- times it fails to take place at the proper time, and the intestine is then apt to protrude into the scrotum, in front of the spermaitic cord, giving rise, in this way, to a congenital. inguinal hernia. (Fig. 251.) The parts implicated, however, in this malformation, have still, as in the case of congenital umbili- cal hernia, a tendency to unite with each Fig. 251. other and obliterate the unnatural open- ing; 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 gubernaculum, as has been described, by certain writers, but by a simple process cosQEiriTAii»aDi»AiHivR- „ .1 . T . 1 . T,v. KIA. — 1. Testicle. 2,2,2. Inteo- 01 growth takmg place in difierent parts, tine, in different directions, at successive periods of foetal life. The gubernaculum, accordingly, has no proper function as a muscular organ, in the human subject, but is merely the anatomical vestige, or analogue, of a corresponding muscle in certain of the lower animals, where it has really an important function to perform. For in them, as we have already mentioned, both the gubernaculum and the cremaster remain fully developed in the adult condition, and are then employed to elevate ■ and depress the testicle, by the alternate contraction of their mus- cular fibres. Female Organs of Generation. — At an early period, as we have mentioned above, the ovaries have the same external appearance, ani occupy the same position in the abdomen, as the testicles in the opposite sex. The descent of the ovaries also takes place, to a great extent, in the same manner with the descent of the testicles. When, in the early part of this descent, they have reached the level of the lower edge of the kidneys, a cord, analogous to the gubernaculum, may be seen proceeding from their lower extremity, crossing the efferent duct on each side, and passing downward, to be attached to the subcutaneous tissues at the situation of the inguinal ring. That part of the duct situated outside the crossing of this cord, becomes afterward convoluted, and is converted into the Fallopian 662 DEVELOPMENT OP THE KIDNEYS, ETC. tube ; while that point which is inside the same point, becomes con- verted into the uterus. The upper portion of the cord itself becomes the ligament of the ovary ; its lower portion, the round UgaTuent of the uterus. As the ovaries continue their descent, they pass below and be- hind the Fallopian tubes, which necessarily perfdrm at the same time a movement of rotation, from before backward and from above downward: j the whole, together with the ligaments of the ovaries and the round ligaments, being enveloped in double folds of peritoneum, which enlarge with the growth of the parts them- selves, and constitute finally the broad ligaments of the uterus. It will be seen from what has been said above, that the situation occupied by the "Wolffian bodies in the female is always the space between the ovaries and the Fallopian tubes; for the Wolffian bodies accompany the ovaries in their descent, just as, in the male, they accompany the testicles. As these bodies now become grad- ually atrophied,' their glandular structure disappears altogether; but their bloodvessels, in many instances, remain -as a convoluted vascular plexus, occupying the situation above mentioned. The Wolffian bodies may therefore be said, in these instances, to un- dergo a kind of vascular degeneration. This peculiar degeneration is quite evident in the Wolffian bodies of the foetal pig, some time before the organs have entirely lost their original form. In the cow, a collection of convoluted bloodvessels may be seen, even in the adult condition, near the edge of the ovary, and between the two folds of peritoneum forming the broad ligament. These are undoubtedly vestiges of the Wolffian bodies, which have undergone the vascular degeneration above described. While the above changes are taking place in the adjacent organs, the two lateral halves of the uterus fuse with each other more and more upon the median line, and become covered with an exces- sively developed layer of muscular fibres. In the lower animfils, the uterus remains divided at its upper portion, running out into two long conical tubes or cornua (Fig. 182), presenting the form known as the uterus bicornis. In the human subject, however, the fusion of the two lateral halves of the organ is nearly complete; so that the uterus presents externally a rounded, but somewhat flattened and triangular figure (Fig. 183), with the ligaments of the ovary and the round ligaments passing off from its superior angles. But, internally, the cavity of the organ still presents a strongly ' marked triangular form, the vestige of its original division. FEMALE OEGANS OF GENEKATION. 663 Occasionally the human uterus, even in the adult condition, re- mains divided into two lateral portions by a vertical septum, which runs from the middle of its fundus downward toward the os in- ternum. This septum may even be accompanied by a partial external division of the organ, corresponding with it iu direction and producing the malformation known as "uterus bicornis." or " double uterus." The OS internum and os externum are produced by partial con- strictions of the original generative passage ; and the anatomical distinctions between the body of the uterus, the cervix and the vagina, are produced by the different development of the mucous membrane and muscular tunic in its corresponding portions. During foetal life, however, Be neck of the uterus grows much faster than its body; so that, at the period of birth, the entire organ is very far from presenting the form which it exhibits in the adult condition. In the human fcetus at term, the cervix uteri constitutes nearly two-thirds of the entire length of the organ; while the body forms but little over one-third. The cervix, at this time, is also much larger in diameter than the body ; so that the whole organ presents a tapering form from below upward. The arbor vitse uterina of the cervix is at birth very fully de- veloped, and the mucous membrane of the body is also thrown into three or four folds which radiate upward from the os intetnum. The cavity of the cervix is filled with a transparent semi-solid mucus. The position of the uterus at birth is also different from that which it assumes in adult life ; nearly the entire length of the organ being above the level of the symphysis pubis, and its inferior extremity passing below that point only by about a quarter of an inch. It is also slightly anteflexed at the junction of the body and cervix. After birth, the uterus, together with its appendages, con- tinues to descend ; until, at the period of puberty, its fundus is situated just below the level of the symphysis pubis. The ovaries at birth are narrow and elongated in form. They contain at this time an abundance of eggs; each inclosed in a Graafian follicle, and averaging gj^ of an inch in diameter. The vitellus, however, is imperfectly formed in most of them, and in some is hardly to be distinguished. The Graafian follicle at this period envelopes each egg closely, there being nothing between its internal surface anJ the exterior of the egg, excepting the thin layer of cells forming the "membrana granulosa." Inside this 664 DEVELOPMENT OF THE KIDNEYS, ^TC. layer is to be seen tlie germinative vesicle, with the germinative spot, surrounded by a faintly granular vitellus, more or less abundant in different parts. Some of the Graafian follicles con- taining eggs are asiarge as ^^^ of an inch; others as small as Ttjss. In the very smallest; the cells of the membrana granulosa appear to fill entirely the cavity of the follicle, and no vitellus or germina-, tive vesiqle is to be seen. DEVELOPMENT OF THB CIBO0LATORY APPARATUS. 665 CHAPTER XVII. DEVELOPMENT OP THE CIRCULATORY . APPARATUS. Theee are three distinct forms or phases of development assumed by the circulatory system duftng different periods of life. These different forms of the circulation are intimately connected with the manner iawMch nutrition and respiration, or the renovation of the blood, are accomplished at different epochs ; and they follow each other in the progress of development, as different organs are em- ployed in turn to accomplish the above functions. The first form is that of the vitelline circulation, which exists at a period when the vitellus, or the umbilical vesicle, is the sole source of nutrition for the foetus. The second is the placental circulation, which lasts through the greater part of foetal life, and is characterized by the existence of the placenta ; and the third is the complete or adult circulation, in which the renovation and nutrition of the blood are provided for by the lungs and the intestinal canal. First, or Vitelline Circulation. — It has already been shown, in a previous chapter, that when the body of the embryo has begun to be formed in the centre of the blastodermic membrane, a number of bloodvessels shoot out from its sides, and ramify ov«r the remainder of the vitelline sac, forming, by their inosculation, an abundant vascular plexus. The area occupied by this plexus in the blastodermic membrane around the foetus is, as we have seen, the " area vasculosa." In the egg of the fowl (Fig. 252), the plexus is limited, on its external border, by a terminal vein or "sinus — the "sinus terminalis ;" and the blood of the embryo, after circulating through the capillaries of the plexus, returns by several venous branches, the two largest of which enter the body near its anterior and posterior extremities. The area vasculosa is, accordingly, a vascular appendage to the circulatory apparatus of the embryo, spread out over the surface of the vitellus for the purpose of absorb- ing from it the nutritious material requisite for the gtowth of the 666 DEVELOPMENT OF THE CIBCULATORT APPARATUS. newly-formed tissues. In the egg of the fish (Fig. 253), the princi- pal'vein is seen passing up in front underneath the head ; while the arteries emerge all along the lateral edges of the body. The entire Fig. 252, Eoo OF PowL ia process of development, shOTring area vaticulosa, with vitelline circalation, terminal sinna, &c. Fig. 253. vitellus, in this way, becomes covered with an abundant vascular network, connected with the internal circulation of the fcetus by arteries and veius, Very soon, as the embryo and the entire egg increase in size, there are two arteries and two veins which become larger than the others, and which subsequently do the whole work of conveying the blood of the fcetus to and from the area vasculosa. These t\^o arteries emerge from the lateral edges of the foetus, on the right and left sides ; while the two veins re-enter at about the same point, and nearly parallel with them. These four vessels are then termed the omphalo-mesentenc arteries and veins. The arrangement of the circulatory apparatus in the interior of the body of the foetus, at this time, is as follows : The heart is situated at the median line, just beneath the head and in front of the oesophagus. It receives at _its lower extremity the trunks of the two omphalo-mesenteric veins, and at its upper extremity divides into two vessels, which, arching over backward, attain the anterior surface of the vertebral column, and then run from above downward along the spine, quite to the posterior Egg of Pish (Jar- rabacca), showing vitel- line circulation. PLACENTAL CIRCULATION. 667 extremity of the foetus. These arteries are called the vertebral arteries, on account of their course and situation, running parallel with the vertebral column. They give ofi^ throughout their course, many small lateral branches, -which supply the body of the foetus, and also two larger branches-^the omphalo-mesenterio arteries — which pass out, as above described, into the area vasculosa. The two vertebral arteries remain separate in the upper ^art of the body, but sOon fuse with each other a little below the level of the heart ; so that, below this point, there remains afterward but one large artery, the abdominal aprta, running from above downward along the median line, giving off the omphalo-mesenteric arteries to the area vasculosa, and supplying smaller branches to the body, the walls of the intestine, and the other organs of the foetus. The above description shows the origin and formation of the first or vitelline circulation. A dbange, however, now begins to take place, by which the vitellus is superceded, as an organ of nutrition, by the placenta, which takes its place ; and the second or placeMal circulation becomes established in the following manner : — Second Circulation. — After the umbilical'vesicle has been formed by the process already described, a part of the vitellus remains in- cluded in it, while the rest is retained in the abdomen and inclosed in the intestinal canal. As these two organs (umbilical vesicle and ^'8- 2.')4. intestine) are originally parts of the same vitelline sac, they remain supplied by the same vascular system, viz : the omphalo-mesen- teric vessels. Those which remain within the abdomen of the foetus supply the mesentery and intes- tine ; but the larger trunks pass outward, and ramify upon the walls of the umbilical vesicle. (Fig. 254.) At first, there are, as we have mentioned above, two omphalo-mesenteric arteries emerging from the body, and two omphalo-mesenteric veins return- ing to it ; but soon afterward, the two arteries are replaced by a common trunk, while a similar change takes place in the two veins. Subsequently, therefore, there remains but a single artery and a Diagram of YoCKa Embbto akb its VrbbeIiS, Bbowing cfrcnlatioa of umbilical YOBicle. and also that of allantoiB, beginning to be formed. 66^ DEYKLOPMENT OF THE CIBCULATOET APPARATUS. single vein, connecting the internal and external portions of the vitelline circulation. The vessels belonging to this system are therefore called the omphalo-mesenteric vessels, because a part of them (omphalic ves- sels) pass outward, by the umbilicus, or " omphalos," to the umbili- cal vesicle, while the remainder (mesenteric vessels) ramify upon the mesentery and the intestine. At first, the circulation of the umbilical vesicle is more import- ant than that of the intestine ; and the omphalic artery and vein appear accordingly as large trunks, of which the mesenteric ves- sels are simply small branches. (Fig. 254.) Afterward, however, the intestine rapidly enlarges, while the umbilical vesicle dimi- nishes, and the proportions existing between the two sets of vessels are therefore reversed. (Fig. 255.) The mesenteric vessels then • Fig. 255. Diaitrain of Embeto and its Vessels; showing the second clrealation. The pharynx, ffidopbagus, and intesUual caaal, have become farther deyeloiled, and the mesenteric arteries have enlarged, while the umbilical voBicle apd its yascnlar branches are very much reduced in size. The large umbilical arteries are seen passing out to the placenta. come to be the principal trunks, while the omphalic vessels are simply minute branches, running out along the slender cord of the umbilical vesicle, and ramifying in a few scanty twigs upon its surface. DEVBLOPMKNT OF THE ARTERIAL SYSTEM. .669 In the injean time, the allantois is formed by a protrusion from the lower extremity of the intestine, which, carrying with it two arteries and two veins, passes out by the anterior opening of the body, .and comes in contact with the external membrane of the egg. The arteries of the allantois, which are termed the' umbilical arteries, are supplied by branches of the abdominal aorta ; the um- bilical veins, on the other hand, join the mesenteric veins, and empty with t'hem into the venous extremity of the heart. As the umbilical vesicle diminishes, the allantois enlarges ; and the latter soon becomes converted, in the human subject, into a vascular chorion, a part of which is devoted to the formation of the placenta. (Fig. 255.) As the placenta soon becomes the only source of nutri- tion for the foetus, its vessels are at the same time very much increased in size, and preponderate over all the other parts of the circulatory system. ' During the early periods of the formation of the placenta, there are, as we have stated above, two umbilical arteries and two umbilical veins. But subsequently one of the veins disappears, and the whole of the blood is returned to the body of the foetus by the other, which becomes enlarged in proportion. For a long time previous to birth, therefore, there are in the umbili- cal cord two umbilical arteries, and but a single umbilical vein. Such is the second, or placental circulation. It is exchanged, at the period of birth, for the third or adult circulation, in which the blood which had previously circulated through the placenta, is diverted to the lungs and the intestine. These are the organs upon which the whole system afterward depends for the nourish- ment and renovation of the blood. During the occurrence of the above changes, certain. other altera- tions take place in the arterial and venous systems, which will now require to be described by themselves. Development of the Arterial System.— At an early period of deve- lopment, as we have shown above, the principal arteries pass off from the anterior extremity of the heart in two arches, which curve backward on each side, from the front of the body toward the vertebral column, after which they again become longitudinal in direction, and receive the name of "vertel»ral arteries." Very soon these arches divide successively into two, three, four, and five secondary arches, placed one above the other, along the sides of the neck. (Fig. 256.) These are termed the cervical arches. In the fish, these cervical arches remain permanent, and give off from their convex borders the branchial arteries, in the form of vascular tufts, 670 DEVELOPMENT OF THE CIKO-ULATORT APPARATUS. to the gills on each side of the neck ; but in the humaa subject and the quadrupeds, the branchial tufts are never developed, and the cervical arches, as well as the trunks with which they are con- nected, become modified by the progress of development in the following manner : — Fig. 256. 2 Early coodition of Arterial Ststeh: showing the heart (1), with its two ascend- ing arterial trunks, giving off on each side five cervical arches, which terminate in the vertebral arteries (2, 2). The vertebral arte- ries unite below the heart to form the aorta (3). Adult condition of Arterial Sts- T EM. —1,1. Carotids. 2, 2. Vertebrals. 3, 3. Kight and left subclavians. 4, 4. Bight and left superior intercostals. 5. Left aortic arch, which remains perma- nent. 6. Bight aortic arch, which die- appears. The two ascending arterial trunks on the anterior part of the neck, from which the cervical arches are given off, become con- verted into the carotids. (Fig. 257, 1,1.) The fifth, or uppermost cervical arch, remains at the base of the brain as the inosculation, through the circle of Willis, between the internal carotids and the basilar artery, which is produced by the union of the two verte- brals. The next, or fourjh cervical arch, may be recognized in an inosculation which is' said to be very constant between the superior thyroid arteries, branches of the carotids, and the inferior thyroids, which come from the subclavians at nearly the same point from which the vertebrals are given off. The next, or third cervical arch, remains on each side, as the subclavian artery (3, s). This vessel, DEVELOPMENT OF THE ARTERIAL SYSTEM. 671 though at first a mere branch of communicatioa between the caro- tid and the vertebral, has now increased in size to such an extent that it has become the principal trunk, from which the vertebral itself is given off as a small branch. Immediatelj below this point of intersection, also, the vertebral artery diminishes very much in relative size, loses its connection with the abdominal aorta, and supplies only the first two intercostal spaces, under the name of the superior intercostal artery (4, 4). The second cervical arch becomes altered in a very different manner on the two opposite sides. On the left side, it becomes enormously enlarged, so as to give off, as • secondary branches, all the- other arterial trunks which have been described, and is converted in this manner into the arch of the aorta (a). On the right side, however, the coft-esppnding arch (e) becomes smaller and smaller, and at last altogether disappears ; so that, finally, we have only a single aortic arch, projecting to the left of the median line, and continuous with the thoracic and abdo- minal aorta. The first cervical arch remains during foetal life upon the left . side, as the " ductus arteriosus," presently to be described. In the adult condition, however, it has disappeared equally upon the right and left sides.- In this way the permanent condition of the arterial circulation is gradually established in the upper part of the body. Corresponding changes take place, however, during the same time, in the lower part of the body. Here the abdominal aorta runs undivided, upon the median line, quite to the end of the spinal column; giving off" on each side successive lateral branches, which supply the intestine and the parietes of the body. "When the allantois begins to be developed, two of these lateral ^branches accompany it, and become, consequently, the umbilical arteries. These two vessels increase so rapidly in size, that they soon appear as divisions of the aortic trunk ; while the original continuation of this trunk, running to the end of the spinal column, appears only as a small branch given off at the point of bifurcation. When the lower limbs begin to be developed, they are supplied by two small branches, given off from the umbilical arteries near their origin. Up to this time the pelvis and posterior extremities are but slightly developed. Subsequently, however, they grow more rapidly, in proportion to the rest of the body, and the arteries which supply them increase in a corresponding manner. That portion of the umbilical arteries, lying between the bifurcation of the aorta and the origin of the branches going to the lower ex- 672 DEVELOPMENT OF THE CIRCULATOBY APPARATUS. tremities,. becomes the common iliaiCS, which in their turn afterward divide into the umbilical arteries proper, and the femorals. Sub- sequently, by the continued growth of the pelvis and lower extremities, the relative size of their vessels is still further in- creased ; and at last the arterial system in this part of the body assumes the arrangement which belongs to the latter periods of gestation. The aorta divides, as before, into the two common iliacs. These also divide into the external iliacs, supplying the lower ex- tremities, and the internaL iliacs, supplying the pelvis; and this division is so placed that the umbilical or hypogastric arteries arise from the internal iliacs, of which they now appear to be secondary • branches. After the birth 9[ the foetus, and the separation of the placenta, the hypogastric arteries become partially atrophied, and are con- verted, in the adult condition, into solid, rounded cords, running upward toward the umbilicus. Their lower portion, however, remains pervious, and gives off arteries supplying the urinary .bladder. The obliterated hypogastric arteries, therefore, the rem- nants of the original umbilical or allantoic arteries, run upward from the internal iliacs along the sides of the Fig. 258. urinary bladder, which is the remnant of the ori- ginal allantois itself. The terminal continuation of the original abdoming,l aorta, is the arteria sacra media, which, in the adult, runs downward on the anterior surface of the sacrum, supplying branches to the rectum and the anterior sacral body; nerves. Development of the Venous System. — ^According to the observations of M. Coste, the venous system at first presents the same simplicity and symmetry with the arterial. The principal veins of the body consist of two long venous trunks, the ver- tebral veins (Fig. 258), which run along the sides of the spinal column, parallel with the vertebral arteries. They receive in succession all the inter- costal veins, and empty into the heart by two lateral trunks of equal size, the canals of Cumer. When the inferior extremities become developed, their two veins, returning from below, join the vertebral veins near the posterior portion of the and, crossing them, afterward unite with each other, thus Early condition of Ve- 11008 System; show- ing the vertebra;! veins emptying into the heart by two lateral trnnks, the "canals of Cuvier." DEVKLOPMENT OF THE VENOUS SYSTEM. 673 constituting another vein of new formation (Fig. 259, a), which runs upward a little to the right of the median line, and empties by itself into the lower extremity of the heart. The two branches, by means of which the veins of 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 &nd 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. 259), 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. 259, h), passing over from left to right, and emptying into the right verte- bral 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 (&), and is finally conveyed to the heart by the right descending vertebral. Soon afterward, this branch of com- munication enlarges so rapidly that it prepon- derates altogether over the left superior verte- bral vein, from which it originated (Fig. 260), 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. 43 'Fig. 259. VsNors System far- ther advanced, sliowing formation of iliac and aub- clarian veins. — a. Vein of new formation, wliicli be- comes the inferior veua cava. 6. Transverse branch of new formation, -which afterward becomes the left vena innominata. Fig. 260. Fnrthe- development of theVKNpuei System.— The vertebral veins are much diminished in size. and the caniil of Cnvier, im the left side, is gradual y disappearing, c. Trans- verse branch of new forma- tion, which is to become the vena azygos mlu^r 674 DEVELOPMENT OF THE CIRCULATORY APPARATUS. Fig. 261. On the left side, that portion of the superior vertebral vein, which is below the subclavian, remains as a small branch of the vena in- nominata, receiving the six or seven upper intercostal 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 cavd^is a totally distinct vein, of new formation, resulting frpm 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. 261, 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 communfcate 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 original direction. That upon the right side still receives all the right intercostal veins, and becomes the vena azygos major (s). It also receives a small branch of communication from its fellow of the left side (Fig. 260,';,c), and this branch soon enlarges to such an extent as to bring over to the veila 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 Adnlt Ciindition of Ve- nous System. — 1. Right auricle of heart. 2. Vena cava superior! 3, 3. Jugular veins. 4,4. Subclavian veins. 5. Vena cava inferior. 6, 6. niacTelns. 7. Lumbarveins. .S. Vena azygos major. 9. Vena azygos minor. 10. Su- perior intercostal vein. DEVELOPMENT OF THE HEPATIC CIBOULATIOS,. 675 upper intercostal veins on the left side still empty, as before, into their own vertebral vein (lo), which, joining the left vena innomi,. nata above, is known as the superior intercostal vein. The left canal of Cuvier has 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 ate still two parts of the circulatory apparatus, the deve- lopment of which presents peculiarities sufficiently important to be described separately. These are, first, the liver and the ductus venosus, and secondly, the heart, with the ductus arteriosus. Development of the Hepatic Circulation and the Ductus Yenosiis. — 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 omphalo-mesenterio« vein, just below its *''S- 262. termination in the heart. (Fig. 262.) As soon as the organ has attained a con- siderable 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. •-""-i..-.-. The omphalo-mesenteric vein below the Eariy form of hkpatio cir- hver then becomes the jJortaZmw; while ,eric vein. 2. He,.atic vein. 3. above the liver, and between that organ Hoait. The dotted iine shows tiio . ^ ^ . . , n Situation of the futare umbilical and the heart, it receives the name 01 vein, the hepatic vein (3). The liver, accord- ingly, is at this time supplied with blood entirely by the portal vein, coming from the umbilical vesicle and the intestine ; and all the blood derived from this source must pass through the hepatic cir- culation 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 coming 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 676 DEVELOPMENT OP THE CIBCULATORY APPARATUS. Fig. 263. Hepatic Circijlation farllieradvanceij.— 1. I'ortal veiu. 2. Umbilical vein. 3. Hepatic vein. the intestine also remains rnactiye, while the placenta increases in size and in functional importance, a time soon arrives when the liver receives more blood by the umbilical vein thari by the portal vein. (Fig. 263.) The umbilical vein then passes into the liver at the longitudinal fissure, and sup- plies the left lobe entirely with its own branches. To the right it sends off a large branch of communication, which opens in- to the portlal vein, and partially supplies the right lobe with umbilical blood. The liver is thus supplied with blood from two different sources, the most abundant of which is the umbilical vein; and aU the blood entering the liver circulates, as be- fore, through its capillary vessels. But we have already seen that the liver is much larger, in pro- portion to the entire body, at an early period of foetal life, than in the later months. In the foetal pig, when very young, it amounta to nearly twelve per cent, of the weight of the whole body ; but be- fore birth it diminishes to seven, six, and even three or four per dent. For some time, therefore, previous to birth, there is much more blood re- turned from the placenta than is re- quired 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 without 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 (5) (Fig. 264), which>, enlarging excessively, be- comes at last converted into a wide canal, or branch of communi- cation, passing directly from the umbilical vein below to the hepatic vein above. The circulation through the liver, thus established, is Fig. 264. Hepatic Circflatiok daring 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 venoHus. 6, Hepatic vein. Fig. 235. DEVELOPMENT OF THE HEPATIC CIKGULATION, 677 as follows : A certain quantity of venous blood still enters through the portal vein (,), and circulates in a part of the capillary system of the right lobe. The umbilical vein (a), bringing a much larger quantity of blood, enters the liver also, a little to the left, and the blood which it contains divides into three principal streams. One of them passes through the left branch (a) into the capillaries of the left lobe ; another turns off through the right branch (4), and, join- ing the blood of the portal vein, circulates through the capillaries of the right lobe; while the third passes directly onward through the venous duct (5), and reaches the hepatic vein without having passed through any part of the capillary plexus. This condition of the hepatic circulation continues until births At that time, two important changes take 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 intestinal canal, en- tering upon the active performance of its functions, becomes the sole source of supply for the hepatic venous blood. The following changes, there- fore, take place at birth in the ves- sels of the liver. (Fig. 265.) First, the umbilical vein shrivels and be- comes converted into a solid rounded cord (a). This cord may be seen, in the adult condition, running from the internal surface of the abdominal walls, at -the umbilicus, to the longi- tudinal fissure of the liver. It is then known under the name of the round ligament. Secondly, the ductus venosus also becomes obliterated, and converted into a fibrous cord. Thirdly, the blood entering the liver by the portal vein (1), passes off by its right branch, as before, to the right lobe. But in the branch (■»), 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 pf the portal vein ; and its blood passes Adult form of Hepatic CiHorLA- Tios.— 1. Portal v6io. 2. Obliterated umbilical vein, forming the round liga- mcDt; the contiuiiatiou of the dotted lines through the liver shows the situa- tion of the obliterated dnctus venosus. 3. Hepatic vein. 4. Left brauch of portal vein. 678 DEVELOPMENT OF THE CIBOULATORY APPAKATUS. from right to left, to be distributed to tbe capillaries of the left lobe. Accordinf to Dr. Guy, the umbilical vein is completely closed at the end of the fifth day after birth. Bevehpment of the Heart, and the Ductus Arteriosus. — When the embryonic circulation is first established, the heart is a simple tubu- lar sac (Fig. 266), receiving the veins at its lower extremity, arid 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. 267); 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. 266. ' Fig. 267. Fig. 268. 2 7 FffiTAL IIji'AKT, divided Earliest form of F (ETA L FffiTAL Heart, twisted into rightand left cavities,— Heart. — 1. Venous ex- -upon itself. — 1'. Venous ex- 1. Venous extremity. 2. tremity. 2. Arterial ex- tremity. 2. Arterial extre- Arterial extremity. 3, 3. tremity, tally. Pulmonary branches. Soon afterward, this single cardiac tube is divided into two paral- lel tubes, right and left, by a Ipngitudinal partition, which grows from the inner surface of its walls . and follows the twisted course of the organ itself. (Fig. 268.) 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. DEVJELOPMENT OF THE HEART. 679 Fig. 269. FffiTAi- Hr ART Still farther developed.— I Aorta. 2. Ph.- moDury artery. 3, 3. Pul- monary brauches 4. Ductus aitei-iosus. 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 eeptum; 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 the immediate neighborhood of the heart, joining again, however, a short distance be- yond the origin of the pulmonary branches. (Fig. 269.) 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 pulmonary 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; aqd 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 pulmpnary branches increase in proportion to the pulmpnary 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 con- siderable portion of the blood, there- fore, coming from the right ventricle still passes onward to the aorta with- out being distributed to the lungs. But at the period of birth, the lungs enter upon the acJtive performance of the function of respiration, and imme- diately require a much larger supply of blood. The right and left pul- monary branches then enlarge, so as to become the two principal divisions of the pulmonary trunk. (Fig. 270.) The ductus arteriosus at the Fig. 270. Hbabt of Infant, showing dis- appearance of arterial duct after birtli .— 1. Aorta. 2. Palmonary artery. 3, 3. Pulmonary branches. . 4. Ductus arteriosus becoming obliterated. 680 DEVELOPMENT OP THE CIECULATORY APPARATUS. same time becomes contracted and shrivelled to such an extent that its cavity is obliterated ; and it is finally converted into an im- pervious, rounded cord, which remains until adult life, running from the point of bifurcation of the pulmonary artery to the under side of the arch of the aorta. The obliteration of the arterial diict id complete, at latest, by the tenth week after birth. (Cruy.) 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 foramen 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 thpoTigh the heart, which is characteristic of foetal life, and which may be described as follows : — It will be found upon examination that the two vense cavse, 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 transversely 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 crescentio border, leaving at that part an oval opening, the foramen ovale. The stream of blood, coraing 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 inJ;o 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 DEVELOPMENT OF THE HEART. 681 Fig. 271. 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 valve. This valve, which is very thin and transparent (Fig. 271, /), is attached to the anterior border of the orifice of the inferior vena cava, and terminates by a crescentic edge, directed towai'd the left; the valve, in this way, standing 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. 271, 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 sometimgs described, in the cavity of the right auricle ; but, owing to the peculiar position and direction of the two veins at this period, with 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 inuominata, 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. 272.) For the blood of the superior vena cava Heart OF Human Fcetub, at the end of the Bixth month ; from a'tipecinietf in the author's pos- sesHiou. — a. Inferior vena cava. h. Superior vena cava. e. Cavity of rl^ht 'anxicle, laid open from tlie front, d. Appendix auricularis. e. Cavity of riifht 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 /, then crossing behind the cavity of the right auricle, and passing through the foramen ovale, to the left side of the heart. 682 DEVELOPMEXT OF THE CIRCULATORY APPARATUS, Fig 272. passes through the right auricle downward into the right ventricle, thence through the pulmonary artery and ductus arteriosus, into the thoracie aorta, while the blood of the inferior vena cava, enter- ino- 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, retilrning from the head and ripper 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 certalin period of foet&l life, is so complete that Dr. John Eeid,' 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 tho- ' racic aorta, with only a slight admixture of red at the posterior part of the right auricle. All the branches of the thoracic and abdominal aorta were filled with yellow, 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. Diiijfmm of' CittcufiATroN TnRriuoH Tnii F(ETAL Hkart. — «. Superior vena cavii. 6. Inferior veua cava, r, c, c, c. Arch or aorta and its brasicheB. d. Pulmuuury artery. Edinburgh Medical and Surgical Journal, vol. zliii. 183%. DEVBLOPMENT OF THE HEABT. 683 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 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-twen- tieths 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. Keid, there was always an ad- mixture 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 manipulation of these experiments, in which it is not always easy to imitate ex- actly the force and rapidity of the different currents of blood in the living foetus. The above results, howeyer, are such as to leave no doubt of the principal 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 heartj 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 (Pig. 271), 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 684 DEVELOPMENT OF THE CIRCULATORY APPARATUS. membranous ridge, ■wbich can exert no influence on the direction of the current of blood passing by it. Thus, the cavity of the infe- rior vena cava, at its upper extremity, ceases to be separated from 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 ovate. 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 jihrough 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 the 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 with the pulmonaiy blood, DEVELOPMENT OF THE HEART. 685 no, longer admits a free access from tte right auricle through the foramen ovale j and the valve of the foramen, pr|psed backward more closely against the edges of the septum, becomes after a time 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 alwiays 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. 272, is then replaced by the adult circulation, represented in Fig. 273. Fig. 273; Diagram of Adhi.t Ci^cclation tbrobohtht! Heart.—",". Snperior and inferior vens CBvaj. S. Eight ventricle. ,,-. Plllmonay artery, dividing into right and left branches, d. Palmo- nary vein. e. Left ventricle. /. A»rta. 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, which has become adhe- 686 DEVELOPMENT OF THE CIBCULATOET APPARATUS. rent to the edges of the septum. The auricular septum in the adult heart is, therefye, thinner at this spot than elsewhere ; and presents, on the side of the right auricle, an oval depression, termed the fossa 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 days after birth. It often, however, remains partially pervioils 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 aperture existing even in adult life. In these instances, however, although the adhesion and solidification of the auricular septum may not be complete, yet no disturbance (^ the circulation "results, and no ad- mixture 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. DEVELOPMENT OP THE BODY AFTEB BIRTH. CS7 CHAPTER XVIII. DEVELOPMENT OP THE BODY APTBE BIETH. The newly-born infant is still very far from having arrived at a state of complete development. The changes through which it has passed during intra-nterine 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 embrypnic 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 uppei* and lower extremities are habitua,lly 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 688 DEVKLOPMENT OF THE BODY AFTER BIETH. seems to be accomplished, daring this period, to a considerable 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 whea 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 difierent from that which they eiert 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 ^nd 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 DEVELOPMENT OF THE BODY AFTEE BIRTH. 689 the estimates of Cruveilheir, Solly, 'Wilson, &c. ; that of the organs in the foetus at term f][fcm our own. observations. Wei PfflTTTS AT TeRU. ADULT. ight of the entire body . 1000.00 1000.00 (( u enoephalon 148.00 23.00 ti ft liver . 37.00 29.00 (( (( heart 7.77* 4.17 tt u kidneys . 6.00 4.00 U 11 renal capsules . 1.63 0.13 It tt thyroid gland . 0.60 0.51 tt u thymus gland . 3.00 0.00 It will be observed that most of the internal organs diminish in relative size after birth, owing principally to the increased develop- ment of the osseous and muscular systems, 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 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 incisfrs 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 44 DEVELOPMENT OF THE BODY AFTER BIRTH. Ahe first set of teeth are thrown off 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 off 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"* sexes becomes more complete and well marked. The beard is developed in the male ; and ip 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 orgains of the body arrive at the adult condition, and the entire process of development is then complete. • INDEX. Absorbent glands, 168, 317 vessels, 167, 317 Absorption, 162 by bloodvessels, 165 by lacteals, 168 of fat, 171 of different lii^aids by animal sub- stances, 312 of oxygen in respira,tion, 243 by egg during incubation, 606 of calcareous matter by allantois, 606 Acid, carbonic, 242, 246 . lactic, in gastric juice, 139 . in souring milk, 99, 336 glyko-oholic, 180 tauro-cholio, 181 ■ pnenmic, 247 uric, 347, 354 oxalic in urine, 860 Acid fermentation of urine, 360 Acidity of gastric juice, cause of, 139 of urine, 353 Acini, of liver, 338 Adipose vpsicles, 90 digestion of, 158, 159 Adult pirculation, 670 establishment of, 685 Aerial respiration, 233 Age, influence of, on «xhalation of car- bonic acid, 250 on comparative weight of organs, 689 Air, quantity of, used in respiration, 238 alterations of, in respiration, 241 circulation of, in lungs, 239 Air-cells of lungs, 235 Air-chamber, in fowl's egg, 552 Albumen, 100 of the blood, 224 in milk, 335 of the egg, how produced, 550 its liquefaction and absorption dur- ing development of foetus, 602- 604 Albuminoid substances, 95 digestion of, 141 Albuminose, 142 interference with Trommer's test, 143 with action of iodine and starch, 144 Alimentary canal in different animals, 116, 119 development of, 644 Alkalies, effect of, on urine, 354 Alkaline chlorides, 71-74 phosphates, 77 ■ carbonates, 76, 77 Alkaline fermentation of urine, 360 AlKalescence of blood, due to carbonates, 76 Allantois, 599 formation of, 601 in fowl's egg, 604 function of, 605 in foetal pig, 622 Alligator, brain of, 382 Amnion, 509 formation of, 600 enlargement of, during latter part of pregnancy, 630 contact with chorion, 631 Amniotic folds, 600 Amniotic fluid, 630 its use, 631 contains sugar at a certain period, 648 'Amniotic umbilicus, 600 Analysis, of animal fluids, 64, 65 of milk, 112, 334 of wheat flour, 112 of oatmeal, 112 of eggs, 113 of meat, 113 of saliva, 124, 126 of gastric juice, 139 of pancreatic juice, 155 of bile, 176 of blood-globules, 218 of blood-plasma, 223 of mucus, 328 of sebaceous matter, 329 of perspiration, 331 of butter, 336 of urine, 352 of fluid of thoracic duct, 318 of chyle and lymph, 320 Ahdral and Gavabret, production of carbonic acid in respiration, 250 Animal functions, 59 Animal heat, 253-263 in different species, 255 mode of generation, 257 (691) 692 INDEX. Animal heat influeuced by local causes, 261 in different organs, 262 increase of, after section of sympa- thetic nerve, 521 Animal and vegetable parasites, 532 Animalcules, infusorial, 529 mode of production, 530 Annulus ovalis, 686 Anterior columns of spinal cord, 380 their excitability, 402 Aorta, development of, 670 Aplysia, nervous system of, 375 Appetite, disturbed by anxiety, &c., 149 necessary to digestion of food, 149 Aquatic respiration, 233 Arch of aorta, formation of, 671 Arches, cervical, 670 transformation of, 671 ^ Area pelluoida, 590 vasculosa, 003, 666 Arteries^ 281 motion of blood in, 282 pulsation of, 283-285 elasticity of, 281, 286 rapidity of circulation in, 289 omphalo-meseuteric, 666 vertebral, 669 umbilical, 669 Arterial pressure, 287 Arterial system, development of, 669-672 Articulata, nervous system of, 376 reflex action in, 377 Articulation of tapeworm, 541 Arytenoid cartilages, 240 movements of, 241 Assimilation, 324 destructive, 341 Auditory apparatus, 505 nerves, 447, 507 Auricle, single, of fish, 265 double, of reptites, birds, and mam- malians, 266, 267 contraction of, 279 -Auriculo-veutricular valves, action of, 269 Axis-cylinder, of nervous filaments, 370 Aztec children, 426 -Azygous veins, formation of, 673 Bbaumoht, Dr. , experiments on Alexis St. Martin, 135-14B Beknakd, on the different kinds of saliva, 125 on effect' of dividing Steno's duct, 131 on digestion of fat in intestine, 155 on formation of liver-sugar, 2O0, 202, '203 on decomposition of bicarbonates in lung, 247 on temperature of blood in different organs, 262 BiDDEE AND ScHMiDT, on daily quantity of bile, 188 on effect of excluding bile from in- testine, 195 on reabsorption of bile, 196 Bile, 17S composition of, 176 tests for, 184 daily quantity of, 188 functions of, 193 reaction with gastric juice, 193 reabsorption, 196 mode of secretion, 337 Biliary salts, 177 of human bile, 183 Biliverdine, 103, 176 tests for, 184 passage into the urine, 357 Blastodermic membrane, 688 Blood, 213 red globules of, 214 white globules, 220 plasma, 223 coagulation o'f, 225 buffy coat, 230 entire quantity of, 231 alterations of, in respiration, 243 temperature of, 254 in different organs, 262 circulation of, 264 through the heart, 270 through the arteries, 282 through the veins, 290 through the capillaries, 296 BoDSSiNOAULT, on chloride of sodium in food, 73 on internal production of fat, 93 Brain, 381, 417 of alligator, 382 of rabbit, 383 human, 386, 417 remarkable oases of injury to, 419, 420 size of, in different races, 423, 424 in idiots, 425 development of, 638, 639 Branchiae, 232 of meno-branohus, 233 Broad ligaments, formation of, 662 Bronchi, division of, 234, 235 ciliary, motion in, 239 Brunner's glands, 152 Buffy coat of the blood, 230 Butter, 335 composition of, 336 condition in milk, 91, 335 Butyrine, 336 Canals of Cuvier, 672 Capillaries, 295 their inosculation, 296 motion of hlood in, 297 Capillary circulation, 296 INDEX. 693 Capillary circulatiou, causes of, 298 rapidity of, 301, 302 peoiiliarities of, in different parts, 303 Caput coli, formation of, 645 Carbonic acid, in the breath,, 242 proportion of, to oxygen absorbed, 242, 243 in the blood, 243 origin of, in lungs, 247 in the blood, 348 In the tissues, 248 mode of production, 248 daily quantity of, 250 variations of, 250 exhaled by skin, 252 by egg, during incubation, 606 absorbed by vegetable^ 260 Carbonate of lime, 76 .of soda, 76 of potassa, 7^ of ammonia, in putrefying urine, 360 Cardiac circulation, in fcetus, 682 in adult, 685 Carnivorous animals, respiration of, 50, 243 urine of, 346 Cartilagine, 102 Caseine, 100 Cat, secretion of bile in, 188 closure of eyelids, after division of sympathetic, 522 Catalytic action, 98 of pepsin, 142 Centipede, nervous system of, 376 Centre, nervous definition of, 373 Cerebrum, 419. See Hemispheres. Cerebral ganglia, 382. See Hemi- spheres. , Cerebellum, 429 effects of injury to, 431 removal of, 431-434 function of, 430 development of, 638, 639 Cerebro-spinal system, 378, 379 development of, 637 Cervix uteri, 554 in foetus, 663 Cervical arches, 670 transformation of, 671 Changes, in egg, while passing through oviduct, 548, 551 in hepatic circulatiou at birth, 677 in comparative size of organs, after birth, 689 Chevreuil, -experiments on imbibition, 312 Chick, development of, 602-607 (Ihildren, Aztec, 426 Chloride of sodium, 71 its proportion in the animal tissues and fluids, 72 Chloride of sodium, importance of, in the food, 73 mode of discharge from the body, 74 partial decomposition of, in the body, 74 Chloride of potassium, 74 Cholesterin, 176 Chorda dorsalis, 591 Chorda tympani, 488 Chordae vooales, movement of, in respi- ration, 240 action of, in the production of vocal sounds, 464 obstruction of glottis by, after divi- sion of pneumogastric, 466, 467 Chorion, formation of, 608 villosities of, 610 source of vascularity of, 611 union with decidua, 619 * Chyle, 153, 169, 320 in lacteals, 170 absorption of, 171 by intestinal epithelium, 172 in blood, 173 Ciliary motion, in bronchi, 22] in Fallopian tubes, 573 Ciliary nerves, 5.14 "" Circulation, 264 in the heart, 270 in the arteries, 282 ' in the veins, 290 in the capillaries, 297 rapidity of, 302 peculiarities of, in different parts, 304 in liver, 339 in placenta, 621-629 Circulatory apparatus, development of, 665-686 "Civilization, aptitude for, of different races, 424 Classification of cranial nerves, 448 Clot, formation of, 225 separation from serum, 226 buffed and cupped, 230 Coagulation, 98 of fibrin, 223 of blood, 225 of white substance of Schwann, in nerve-fibres, 369 CoLiH, on unilateral mastication, 127 Cold, resistance to, by animals, 253 effect of, when long continued, 254 Colostrum, 333 Coloring matters, 103 of blood, 102, 218 of the skin, 103 of bile, 103 of urine, 103 Commissure, of spinal cord, gray, 381 white, 381 transverse, of cerebrum, 387 of cerebellum, 387 * 694 INDEX. Commissures, nervous, 373 olfactory, 382, 418 Congestion, of ear, &o., after division of sympathetic, 521 Convolvulus, sexual apparatus of, 540 Consentaneous action of muscles, 430 Contact, of chorion and amnion, 631 of decidua vera and reflexa, 632 Contraction, of stomach during diges- tion, 145 of spleen, 208 of hlood-clot, 226 of diaphragm and intercostal mus- cles, 236 of posterior crico-arytenoid muscles, 241 ' of ventricles, 275 of muscles after death, 389 of sphincter ani, 414 of rectum, 414 of urinary bladder, 415 of pupil, under influence of light, 367, 435, 501 after division of sympathetic, 522 Cooking, effect of, on food, 114 Cord, spinal, 379, 398 umbilical, 631 , withering and separation of, 689 Corpus callosum, 387 Corpus luteum, 576 of menstruation, 576-580 of pregnancy, 580-585 three weeks after menstruation, 578 four weeks after menstruation, 579 nine weeks after menstruation, 579 at end of second month of preg- nancy, 582 at end of fourth month, 582 at term, 583 disappearance of, after delivery, 584 Corpora Malpighiana, of spleen, 191 Corpora striata, 383, 419 Corpora olivaria, 384 Corpora Wolffiana, 655 CosTE, on rupture of Graafian follicle in menstruation, 572, 573 Cranial nerves, 446 classification of, 448 motor, 449 sensitive, 449, 450 Creatine, 346 Creatinine, 346 Cremaster muscle, formation of, 659 function of, in lower animals, 660 Crystals, of stearine, 87 and margarine, 88 of cholesterin, 177 of glykocholate of soda, l78 of biliary matters of dog's bile, 182 of urea, 343 » of creatine, 346 of creatinine, 346 Crystals, of urate of soda, 347 of uric acid, 354 of oxalate of lime, 360 of triple phosphate, 362 Crystallizable snstances of organic ori- gin, 79 Crossing of fibres in medulla oblongata, 385, 404 of sensitive fibres jn spinal cord, 405 of fibres of optic nerves, 436, 437 of streams of blood in foetal heart, 681, 682 Cebikshank, rupture of Graafian follicle in menstruation, 572 Cumulus proligerus, 567 Cutaneous respiration, 252 perapira1;ion, 330 Cuticle, exfoliation of, during fffital life, 643 after birth, 689 Cysticercus, 537 transformation of into taenia, 538 production of, from eggs of taenia, 539 Death, a necessary consequence of life, 626 Decidua, 614 ■ vera, 616 reflexa, 618 union with chorion, 619 its discharge in cases of abortion, 618 at the time of delivery, 633 Decussation of anterior columns of spinal cord, 385, 404 of optic nerves, 436, 437 Degeneration, fatty, of muscular fibres of uterus, after delivery, 635 Deglutition, 133 retarded by division of Steno's duct, 131 by division of pnenmogastrie, 463 Dentition, first, 689 second, 690 Descent of the testicles, 658 of the ovaries, 661 Destructive assimilation, 341 Development of the impregnated egg, 586 of allantois, 601 of chorion, 608 of villosities of chorion, 610, 611 of decidua, 615, 616 of placenta, 621-626 of nervous system, 637 , of eye, 640 of ear, 641 of skeleton, 641 • of limbs, 642 of integument, 643 of alimentary canal, 464, 646 INDKX. 695 Development of urinary passages, 646 of liver, 649, 675 of pliarynx and cesophagus, 650 of face, 651 of Wolffian bodies, 655 of kidneys, 656 of internal generative organs, 657 of circulatory apparatus, 665 of arterial system, 669 of venous system, 672 of hepatic circulation, 675 of heart, 678 of the body after birth) 687 Diabetes, 357 in foetus, 649 Diaphragm, action of in breathing, 237 formation of, 651 Diaphragmatic hernia, 651 Diet, iniluence of on nutrition, 108 on products of respiration, 243 on formation of urea, 345 of urate of soda, 348 Diffusion of gases in lungs, 239 Digestion, 115 of starch, 150 of fats, 153 of sugar, 150 of organic substances, 141 time required for, 146 Digestive apparatus of fowl, 117 of ox, 118 of man, 119 Discharge of eggs from ovary, 649 independent of sexual intercourse, 565 mechanism of, 568 during menstruation, 572 Discus proligerus, 567 Distahee and solidity, application of, by the eye, 501, 502 Distinction between corpora lutea of menstruation and pregnancy, 585 Diurnal variations, in exhalation of car- bonic acid, 252 in production of urea, 345 in density and acidity of urine, 351 Division of nerves, 371 of heart, into right and left cavities, 678 DoBSON, on variation in size of spleen, 208 Draper, John C, on production of urea, 345 Drugs, effect of, on newly born infant, 688 Ductus arteriosus, 679, 682 closure of, 679, 680 venosus, 676 obliteration of, 677 Duodenal glands, 152 fistula, 190 DniEooHBT, on temperature of plants, 256 on endosmosis of water with differ- ent liquids, 309* Ear, 505 muscular apparatus of, 506 development of, 641 ' Earthy phosphates, 74, 77 in urine, 353 precipitated by addition of an alkali, 354 Ectopia cordis, 651 Egg, 544 its contents, 545 where formed, 546 of frog, 547, 548 of fowl, 549 ■ changes in, while passing through the oviduct, 549-652 pre-existence of, in ovary, 563 development of, at period of puberty, 564 periodical ripening and discharge, 565 discharge of, from Graafian follicle, 568 impregnation of, how accomplished, 561 development of, after impregnation, 586 of fowl, showing area vasculosa, 603 ditto, showing formation of allantois, 604 of fish, showing vitelline circulation, 666 attachment of, to uterine mucous membrane, 617 discharge of from uterus, at the time of delivery, 633 condition of in newly bom infant, 663 Elasticity, of spleen, 208 of red globules of blood, 216 of lungs, 235, 237 of costal cartilages, 237 of vocal cords, 241 of arteries, 281 Electrical current, effect of on muscles. 389 on nerve, 391 different effects of direct and inverse. 394 • Electrical fishes, phenomena of, 396 Electricity, no manifestations of in irri- tated nerve, 397 Elevation of temperature, after division of sympathetic, 261, 521 Elongation of heart in pulsation, 275 anatomical causes of, 276 Embryo, formation of, 686 Embryonic spot, 590 Encephalon, 381, 417 ganglia of, 386 Endosmosis, 307 of fatty substances, 171 in capillary circulation, 314 conditions of, 308 cause of, 311 INDEX. Endosmosis of iodide of potassiiim, 313 of atropine, 313 of nux volnioa, 314 Endosmometer, 308 Enlargement of amnion, during preg- nancy, 630, 631 Entozoa, encysted, 534 . mode of production, 536 Epithelium, in saliva, 124 of gastric follicles, 134 » of intestine, during digestion, 172 Epidermis, exfoliation of, in foetal life, 643 after birth, 689 Epididymis, 659 Exoretine, 160 Excretion, 341 nature of, 342 importance to life, 342 products of, 343 by placenta, 628 Bxcrementitious substances, 342 mode of formation of, 342 effect of retention of, 342 Exfoliation of cuticle, during foeta} life, 643 after birth, 689 Exhalation, 307 of watery vapor, 71 from the lungs, 242 from the skin, 330 from the egg, during incubation, 606 of carbonic acid, 242 of nitrogen, 242 of animal vapor, 242 Exhaustion, of muscles, by repeated irri- tation, 390 of nerves, by ditto, 392 Exosmosis, 308 Expiration, movements of, 237 after section of pneumogastric, 469 Extractive matters of the blood, 225 Eye, protection of, by movements of pu- pil, 367, 435, 501, 518 by two sets of muscles, 519 Eyeball, inflammation of, after division of 5th pair, 455 Eyilids, formation of, 641 Face, sensitive nerve of, 451 motor nerve, 456 development of, 651 Facial nerve, 456 sensibility of, 459 influence of, on muscular apparatus of eye, 457 of nose, 458 of ear, 459 paralysis of, 458, 459 Fallopian tubes, 553 formation of, 661 « Farinaceous substances, 79 in food, 80, 106 digestion of, 150 Fat, decomposition of, in the blood, 166, 173 fats, 86 proportion of, in different kinds of food, 88 condition, in the various tissues and fluids, 88 internal source of, 93 decomposed in the body, 94 indispensable as ingredients of the food, 106 Patty matters of the blood, 224 Fatty degeneration of deoidua, 634 of muscular fibres of uterus, after delivery, 635 Feces, 160 Female generative organs, 546 of frog, 547 of fowl, 549 of sow, 553 of human species, 554 development of, 661 Fermentation, 99 of sugar, 85 acid, of urine, 359 alkaline, of ditto, 360 Fibrin, 100 of the blood, 223 coagulation of, 223 varying quantity of, in blood of dif- ferent veins, 224 Fifth pair of cranial nerves, 451 its distribution, 452 division of, paralyzes sensibility of face, 453 and of nasal passages, 454 produces inflammation of eye- ball, 455 lingual branch of, 456 small root 6t, 452 Fish, circulation of, 247 formation of umbilical vesicle jn, . 596 vitelline circulation,in embryo of,667 Fish, electrical, phenomena of, 396 Fissure, longitudinal, of brain and spinal cord, 379 formation of, 640 Fissure of palate; 654 Fistula, gastric. Dr. Beaumonf s case of, 135 mode of operating for, 136 duodenal, 190 Flint, Prof. Austin, Jr., steroorine, in contents of large intestine, 160 cholesterin, in blood of jugular vein, 177 not discharged with the feces, 177 effects of biliary fistula, 196 Foetal circulation, first form of, 665 second form of, 667 Follicles, of stomach, 134 of Lieberkunn, 151 INDEX. 697 Follicles, of Brunner's glands, 152 Gl-aafian, 546, 567 of uterus, 654, 615 Food, 105 composition of,'ll2 daily quantity required, 113 effect of cooking on, 114 Foramen ovale,' 680 valve of, 684 closure of, 684 Force, nervous, nature of, 395 Formation of sugar in liver, 200 in foetus, 64» Fossa ovalis, 686 Functions, animal, 59 vegetative, ^8 of teeth, 121 of saliva, 129 of gastric juice, 141 of pancreatic juice, 155 of intestinal juices, 153 of bile, 193 of spleen, 210 ^ eft' mucus, 328 of sebaceous matter, 329 of perspiration, 331 of the tears, 832 Galvanism, action of, on muscles, 389 ou nerves, 391 Ganglion, of spinal cord, 380 of tuber annulare, 438 of medulla oblongata, 439 Casserian, 451 of Andersoh, 460 pueumogastric, 461 ophthalmic, ^14 spheno-palatine, 489, 514 ^ submaxillary, 514 otic, 515 semilunar, 516 impar, 516 * Ganglionic system of nerves, 379, 514 Ganglia, nervous, 372 of ladiata, 373 of moUusca, 375 of artiQulata, 376 of posterior roots of spinal nerves, 380 of alligator's brain, 382 of rabbit's brain, 383 of medulla oblongata, 384 of human brain, 386 of great sympathetic, 514 olfactory, 382, 418 optic, 382, 434 Gases, diffusion of, in lungs, 239 absorption and exhalation of, by lungs, 244 by the tissues, 248 Gastric follicles, 134 Gastric juice, mode of obtaining, 137 composition of, 139 Gastric juice, action on food, 141 interference with Trommer's test, 143 interference with action of starch and iodine, 144 daily quantity of, 147 solvent action of, on stomach, after death, 149 Gelatine, how produced, 64 effect of feeding animals on, llJ9 Generation, 527 spontaneous, 527 of infusoria, 530 of parasites, 533 of encysted entozoa, 535 of taenia, 538 sexual, by germs, 540 Germ, nature of; 540 Germination, heat produced in, 238 Germinative vesicle, 545 disappearance of, in mature egg, 586 Germinative spot, 545 Gills, offish, 232 of menobranchus, 233 Glands, of Brunner, 162 mesenteric, 168 ' vascular, 210 Meibomian, 329 perspiratory, 330 action of, in secretion, 324 Glandulse solitarise and agminatae, 162 «Globule3, of blood, 213 red, 214 different appearances of, under microscope, 214, 215 mutual adhesion of, 215 color, consistency, and structure of, 216 action of water on, 217 composition of, 218 size, &c., in different animals, 219 white, 220 action of acetic acid on, 221 red and white, movement of, in circulation, 297 Globnline, 101, 218 Glomeruli, of Wolffian bodies, 656 Glosso-pharyngeal nerve, 459 action of, in swallowing, 460 Glottis, movements of, in respiration, 240 in formation of voice, 464 closure of, after section of pneumo- gastrios, 467 Glycine, 181 Glyco-cholic acid, 180 Glyco-cholate of soda, 180 its crystallization, 178, 179 Glycogenic function of liver, 200 in fcetus, 649 Glycogenic matter, 204 Its conversion into sugar, 204 GossELiN, experiments ou imbibition by cornea, 313 INDEX. Graafian follicles, 5^6, 567 structure of, 567 rupture of, and discharge of egg, 568 ruptured during menstruation, 672 condition of fcEtus at term, 663 Gray substance, of nervous system, 372 of spinal cord, 380 of brain, 386 its want of irritability, 417 Great sympathetic, 614 anatomy of, 515 sensibility and excitability of, 517 connection of, with special senses, 518 division of, influence on animal heat, 521 on pupil and eyelids, 522 reflex actions of, 524 Gubernaculum testis, 659 function of, in lower animals, 661 Gustatory nerve, 452, 483 Hammond, Prof. Wm. A., on effects of non -nitrogenous diet, 108 on production of urea, 344 Hsematine, 102, 218 Hairs, formation of, in embryo, 643 Hare-lip, 653 Harvey, on motions of heart, 273 Hearing, sense of, 505 apparatus of, 506 analogy of with touch, 610, 511 Heart, 265 of fish, 265 of reptiles, 266 of mammalians, 267 of man, 268 circulation of blood through, 270 sounds of, 270 movements of, 273 impulse, 279 development of, 651, 678 Heat, vital, of animals, 263 of plants, 256 how produced, 257 increased by division of sympathetic nerve, 261,621 Hemispheres, cerebral, 419 remarkable cases of injury to, 419, 420 effect of removal, on pigeons, 421 effect of disease, in man, 422 xjomparative.size of, in different races, • 423 functions of, 425 development of, 638 Hemorrhage, from placenta, in parturi- tion, 633 Hepatic circulation, 339 development of, 675 Herbivorous animals, respiration of, 50, 243 ? - uriue of, 346, 348 Hernia, congenital, diaphragmatic, 651 umbilical, 646 inguinal, 661 Hippurate of soda, 348 Hunger and thirst, continue after divi- sion of pneumogastrio, 473 Hydrogen, displacement of gases in blood by, 245 exhalation of carbonic acid in an atmosphere of, 249 Hygroscopic property ■ of organic sub- stances, 97 Hypoglossal nerve, 477 Imbibition, 307 of liquids, by different tissues, 312 by cornea, experiments on, 313 Impulse, of heart, 279 Infant, newly-born, characteristics of, 687 Infiammation of eyeball, after division of 5th pair, 465 Infusoria, 529 dififerent kinds of, 530 conditions of their produotionf 531 Schultze's experiment on generation of, 532 Inguinal hernia, congenital, 661 Injection of placental sinuses from ves- sels of uterusj 627 Inorganic substances, as proximate prin- ciples, 69 , their source and destination, 78 Inosculation, of veins, 291 of capillaries, 296 of nerves, 372 Insalivation, 123 importance of, 131 function of, 132 Inspration, how accomplished, 236 movements of glottis in, 240 Instinct, nature of, 444 Integument, respiration by, 252 development of, 643 Intellectual powers, 422 in animals, 444 Intestine, of fowl, 117 of man, 119 juices of, ,150 digestion in, 150-159 epithelium of, 172 disappearance of bile in, 196 development of, 693, 644 Intestinal digestion, 160 Intestinal juices, 151 action of, on starch, 163 Involution of uterus after delivery, 635 Iris, movements of, 367, 435, 501, 518 after division of sympathetic, 522 Irritability, of gastric mucous membrane, 137 of the heart, 276 ' , of muscles, 389 of nerves, 391 INDEX. 699 Jackson, Prof. Samuel, on digestion of fat in intestine, 1S4 Jaundice, 185 yellow color of urine in, 357 Kidneys, peculiarity of circulation in, 305 elimination of medicinal substances by, 357 formation of, 656 EncHENMEiSTEB, experiments on produc- tion of taenia from cysticercuS, 538 of oysticercus from eggs of tsenia, 539 Lachrymal secretion, 332 its function, 333 Lactation, 333 varia.tions in composition of milk during, 337 Laoteals, 168, 170, 319 and lymphatics, 167, 172 Larynx, action of, in respiration, 222 in formation of voice, 464 nerves of, 462, 464 protective action of, 466 movements in respiration, 466 Lassaighe, experiments on saliva, 132 analysis of lymph, 318 Layers, external and internal, of blasto- dermic membrane, 688 Lead, salts of, action in distinguishing the biliary matters, 183 Lehuann, on formation of carbonates in blood, 76 on total quantity of blood, 231 on effects of non-nitrogenous diet, 108 Lens, crystalline, action of, 495 Leuckaet, on production of oysticercus, 539 LiEBia, on absorption of different liquids under pressure, 310 Ligament of the ovary, formation of, 662 Limbs, formation of, in frog, 594 in human embryoj 642 Liver, vascularity of, 338 lobules of, 338, 339 secreting cells, 339, 340 formation of sugar in, 200 congestion of, after feeding, 207 development of, 649, 675 Liver cells, 92, 340 their action in secretion, 340 Liver-sugar, formation of, 200 after death, 203 in foetu^, 649 Lobules, of lung, 235 of liver, 338, 339 Local production of carbonic aoid, 248 of animal heat, 261 Local variations of circulation, 305 LoNGET, on interference of albuminose with Trommer's test, 143 on sensibility of glosso-pharyngeal nerve, 459 on irritability of anterior spinal roots, 401 LoHGET AND Mattbpcci, experiment on signs of electricity in an irritated nerve, 397 Long and short-sightedness, 496 Lungs, structure of, in reptiles, 234 in man, 235 alteration of, after division of pneu- mogastrics, 469 LympTi, 169,318 quantity of, 320 Lymphatic system, 168, 317 Magnus, on 'proportions of oxygen and carbonic acid in blood, 245 Male organs of generation, 556 development of, 658 Malpighian bodies of spleen, 209 Mammalians, circulation in, 267 Mammary gland, structure of, 333 secretion of, 334 Mabcet, on excretiue, 160 Mabey, M., experiments on arterial pul- sation, 285 Mastication, 121 unilateral, in ruminating animals, 127 retarded by suppressing saliva, 131 Meconium, 648 Medulla oblongata, 384, 439 ganglia of, 385, 386 reflex action of, 440 effect of destroying, 442 development of, 638 Meibomian glands, 329 Melanine, 103 • • Membrane, blastodermic, 688 Membrana granulosa, 567 Membrana tympani, action of, 508 Memory, connection of, with cerebral hemispheres, 425 Menobranchus, size of blood-globules in, 220 gills of, 233 spermatozoa of, 557 Menstruation, 570 commencement and duration of, 571 phenomena of, 571 rupture of Graafian follicles in, 572 suspended during pregnancy, 571, 581 Mesenteric glands, 168, 317 Michel, Dr. Myddleton, rupture of Graaf- ian follicle in menstruation, 572 MUk, 333 composition and properties of, 333, 334 microscopic characters, 335 - 700 INDEX. Milk, souring and coagulation of, 336 variations in, during lactation, 337 Milk-sugar, 83 converted into lactic acid, 336 Mollusca, nervous system of, 375 Moore and Penjsook, experiments on movements of heart, 275 Motion, 400 Motor cranial nerves, 449 ■ Motor nervous fibres, 403 Motor oculi communis, 450 extemus, 451 Movements, of stomach, 145 of intestine, 164 of heart, 273 of chest, in respiration, 237 of glottis, 240 associated, 408 of foetus, 631 Mucosine, 101 Mucous follicles, 327 Mucous membrane, of stomach, 133 of intestine, 151 of tongue, 483 of uterus, 554, 615 Mucus, 327 composition and properties of, 328 of mouth, 125 of cervix uteri, 554 Muscles, irritability of, 388 directly paralyzed by sulpho-cyanide of potassium, 390 consentaneous action of, 430 of respiration, 236 Muscular fibres, of spleen, 208, 209 of heart, spiral and circular, 277 Muscular irritability, 388 duration after death, 388 exhausted by repeated irritation, 390 Masculine, 102 Nails, formation of, in embryo, 643 Negeier, on rupture of Graafian follicle, in menstruation, 572 Nerve-cells, 372 Nerves, division of, 372 inosculation of, 373 irritability of, 390 spinal, 39S cranial, 446 .,.,.. olfactory, 446 ~~" optic, 447 auditory, 447 ooulo-motorius, 450 ^ patheticus, 451 motor extemus, 451 masticator, 452 facial, 456 hypoglossal, 477 spinal accessory, 474 trifacial (5th pair), 451 glosso-pharyngeal, 459 Nerves, pneumogastric, 461 superior and inferior laryngeal, 4G2 great sympathetic, 514 Nervous filaments, 368 of brain, 309 of sciatic nerve, 370 motor and sensitive, 375 Nervous force, how excited, 391 mature of, 395 Nervous tissue, two kinds of, 368 Nervous irritability, 390 how shown, 391 duration of, after death, 391 exhausted by excitement, 392 destroyed by wporara, 393 distinct from muscular, 395 nature of, 395 Nervous system, 365 general structure and functions of^ 365-387 of radiata, 373 of mollusca, 375 of articulata, 376 of vertebrata, 379 reflex action of, 374 Network, capillary, in Peyer's glands, 162 in web of frog's foot, 296 in lobule of liver, 339 Newly-born infant, weight of, 687 respiration in, 687. nervous phenomena of, 688 comparative size of organs in, 689 Newport, on temperature of insects, 256 Nitric acid, action of, on bile-pigment, > 184 precipitation of uric acid by, 354 Nitrogen, exhalation of, in respiration, 242 Nutrition,'-61-364 Obliteration^ of ductus venosus, 677 of ductus arteriosus, 680 Oculo-motorius nerve, 450 CEsophagus, paralysis of, after division of pneumogastric, 463 developBBient of, 645, 650 CEstruation, phenomena of, 569 Oleaginous substances, 86 in different kinds of food, 88 condition of, in the tissues and fluids, 88-93 partly produced in the body, 93 decomposed in the body, 94 in the blood, 170 indispensable as ingredients of the food, 107 insufficient for nutrition, 108 Olfactory apparatus, 489 protected by two sets of muscles, 520 commissures, 382, 418 Olfactory ganglia, 382, 418 their function, 418 INDEX. 701 Olfactory nerves, 489 Olivary bodies, 384 Omphalo-mesenterio vessels, 668 Ophthalmic ganglion, 514 Optic ganglia, 382, 434 Optic nerves, 447 decussation of, 436,437 Optic thalami, 41 8 development of, 638 Organs of special sense, 483, 489, 493, 507 development of, 640 Organic substances, 95 indefinite chemical composition of, 95 hygroscopic properties, 97 coagulation of, 98 catalytic action, 98 putrefaction, 99 source and destination, 104 digestion of, 141 Origin, of plants and animals, 527 of infusoria, 529 of animal and vegetable parasites, 532 of encysted entozoa, 534 Ossification of skeleton, 642 Osteine, 102 Otic ganglion, 515 Ovary, 541 of taenia, 541 of frog, 547 of fowl, 551 of human female, 553 Ovaries, descent of, in foetus, 661 condition at birth, 663 Oviparous and viviparous animals, dis- tinction between, 563 Oxalic acid, produced in urine, 360 Oxygen, absorbed in respiration, 242 daily quantity consumed, 242 state of solution in blood, 245 dissolved by blood-globules, 245 absorbed by the tissues, 248 exhaled by plants, 260 Palate, formation of, 653 Pancreatic juice, 153 mode of obtaining, 154 composition of, 165 action on fat, 155 daily quantity of, 155 Pancreatine, 101 in pancreatic juice, 155 Panizza, experiment on absorption by bloodvessels, 165 Paralysis, after division of anterior root of spinal nerve, 402 direct, after lateral injury of spinal cord, 405 crossed, after lateral injury of brain, 405 facial, 458 Paralysis of muscles, by snlpho-cyanide of potassium, 390 of motor nerves, by woorara, 393, 413 of sensitive nerves, by strychnine, 413 of voluntary motion and sensation, after destroying, tuber annulare, 438 of pharynx and oesophagus, after section of pneumogastrics, 463 of larynx, 465, 467 of muscular coat of stomach, 473 Paraplegia, refiex action of spinal cord in 411 Parasites, 532 conditions of development of, 533 mode of introduction into body, 534 sexless, reproduction of, 535 Parotid saliva, 126 Parturition, 633 Par vagum, 461. See Pneumogastric Patheticus nerve, 451 Pelouze, jomposition of glycogenic mat- ter, 204 Pelvis, development of, 642 Pbhmock and . Moore, experiments on movements of heart, 275 Pepsine, lOJ. in gastric juice, 139 Perception of sensations, after removal of hemispheres, 422 destroyed, after removal of tuber annulare, 438 Periodical ovulation, 563 Peristaltic motion, of stomach, 145 of intestine, 164 of oviduct, 54S, 550 Pebeins, Maurice, composition of parotid saliva, 126 Perspiration, 330 daily quantity of, 331 composition and properties of, 331 function, in regulating temperature, 831 Pettenkofer's test for bile, 185 Peyer's glands, 162 Pharynx, action of, in swallowing, 460, 461 formation of, 650 Phosphate of lime, its proportion in the animal tissues and fluids, 74 in the urine, 353 precipitated by alkalies, 354 Phosphate, triple, in putretying urine, 362 Phosphates, alkaline, 77 in urine, 353 earthy, 74, 77 in urine, 353 of magnesia, soda, and potassa, 77 Phosphorus, not a proximate principle, 63 Physiology, definition of, 49 702 IXDKX. Phrenology, 427 objections to, 428 practical difficulties of, 428, 429 Pigeon, after removal of cerebrum, 421 of cerebellum, 431 Placenta, 621 comparative anatomy of, 622 formation of, in human species, 623 foetal tufts of, 625 maternal sinuses of, 626 injection of, from uterine vessels, 627 function of, 628 separation of, in delivery, 633 Placental circulation, 624, 628 Plants, vital heat of, 256 , generative apparatus of, 540 Plasma of the blood, 223 Pneumic acid, 247 Pneumogastrio nerve, 461 its distribution, 462 action of, on pharynx and oesopha- gus, 463 on larynx, 464 in formation of voice, 464 in respiration, 466 effect of its division on respiratory movements, 466, 467 cause of death after division of, 470 influence of, on oesophagus and sto- mach, 473 Pneumogastrio ganglion, 461 PoGGiALB, on glycogenic matter in but- cher's meat, 205 Pons Varolii, 387 Portal blood, quantity of fibrin in, 224 temperature of, 262 Portal vein, in liver, 338 development of, 675 Posterior columns of spinal cord, 381 Primitive trace, 690 Production, of sugar in liver, 200 of carbonic acid, 246 of animal heat, 253 of urea in blood, 343 of infusorial animalcules, 529 of animal and vegetable parasites, 532 Proximate principles, 61 definition of, 63 mode of extraction, 64 manner of their association, 65 varying proportions of, 66 three distinct classes of, 67 Proximate principles of the first class (inorganic), 69 of the second class (crystallizable substances of organic origin), 79 of the third class (organic sub- stances), 95 Ptyaline, 124 Puberty, period of, 565 signs of, in female, 570 Pulsation, of heart, 270 in living animal, 274 of arteries, 282 Pupil, action of, 367, 435 contraction of, after division of sym, pathetic, 522 Pupillary menfbrane, 640 Putrefaction, 99 of the urine, 358 Pyramids, anterior, of medulla oblon- gata, 384 Quantity, daily, of water exhaled, 71 of food, 113 of saliva, 128 of gastric juice, 147 of pancreatic juice, 155 of bile, 188 of air used in respiration, 238 of oxygen used in respiration, 242 of carbonic acid exhaled, 250 of lymph and chyle, 320 of fluids secreted and reabsorbed, 323 of material absorbed and discharged, 363 of perspiration, 331 of urine, 349 of urea, 344 of urate of soda, 348 Quantity, entire, of blood in body, 231 Rabbit, brain of, 383 Races of men, different capacity of, for civilization, 424 Radiata, nervous system of, 373 Rapidity of circulation, 302 Reactions, of starch, 82 of sugar, 84 of fat, 86 of saliva, 124, 125 of gastric juice, 139 of intestinal juice, 153 of pancreatic juice, 155 of bile, 175 of mucus, 328 of milk, 334 of urine, 353 Reasoning powers, 425 in animals, 444 Red globules of blood, 213 Reflex action, 374 in centipede, 377 of spinal cord, 408 of medulla oblongata, 440 of tuber annulare, 443 of brain, 444 of optic tubercles, 435 in newly born infant, 688 Regeneration, of uterine mucous mem- brane after pregnancy, 633, 634 of walls of uterus, 635 Regnault and Beiset, on absorption of oxygen, 243 INDEX. 703 Reid, Dr. John, experiment on crossing of streams in foetal heart, 682 Reproduction, 525 nature and object of, 625, 627 of parasites, 533 of taenia, 538 by germs, 540 Reptiles, circulation of, 266 Respiration, 232 by gills, 233 by lungs, 234 by skin, 252 changes in air during, 241 changes in blood, 243 of newly born infant, 687 Respiratory movements of chest, 236 of glottis, 240 after section of pneumogastrics, 466 after injury of spinal cord, 441 Restiform bodies, 385 Rhythm of heart's movements, 279 Rotation of heart during contraction, 278 Round ligament of the uterus, formation of, 662 of liver, 67.7 Rumination, movements of, 118, 127 Rupture of Graafian follicle, 568 iu menstruation, 572 Rutting condition, in lower animals, 569 Saccharine substances, 83 iu stomach and intestine, 150 in liver, 200 in blood, 207 in urine, 357 Saliva, 123 different kinds of, 125 daily quantity of, 127 action on boiled starch, 129 variable, 130 does not take place in stomach, 130 physical function of saliva, 131 . quantity absorbed by difierent kinds of food, 132 Salivary glands, 125 Salts, biliary, 177 of the blood, 225 of urine, 353 Saponification, of fats, 87 SuH AELiNG, on diurnal variations in exha- lation of carbonic acid, 252 SoBDLTZB, experiment on^ generation of infusoria, 531 Scolopendra, nervous system of, 376 Sebaceous matter, 328 composition and properties of, 329 function of, 329 in foetus, 643 Secretion, 324 varying activity of, 326 of saliva, 125 of gastric juice, 137 of intestinal juice, 152 Secretion of pancreatic juice, 164 of bile, 188, 337 of sugar iu liver, 200 of mucus, ^27 of sebaceous matter, 328 of perspiration, 330 of the tears, 332 of bile in foetus, 649 Segmentation of the vitellus, 587 Seminal fluid, 566 mixed cnnstitutiou of, 560 Sensation, S98 remains after destruction of hemi- spheres, 422 lost after removal of tuber annulare, 438 special conveyed by pneumogastrio nerve, 440, 466, 468 Sensation and motion, distinct seat of, in nervous system, 400 in spinal cord, 403 Sensibility, of nerves to electric current, 391 and excitabilityj definition, of, 400 seat of, in spinal cord, 403 in brain, 417 of facial nerve, 459 of hypoglossal nerve, 477 of spinal accessory, 475 of great sympathetic, 517 Sensibilityj general and special, 478 special, of olfactory nerves, 446, 489 of optic nerves, 447 of auditory nerves, 447 of lingual branch of 5th pair, 484 of glosso-pharyngeal, 460 of pneumogastrio, 468 Sensitive nervous filaments, 375 Sensitive fibres, crossing of, in spinal cord, 405 of facial nerve, source of, 459 • Sensitive cranial nerves, 449 Septa, inter-auricular and inter-ventri- cular, formation of, 678, 680 S^QDARD, on crossing of sensitive fibres in spinal cord, 405 Serum, of the blood, 227 Sexes, distinctive characters of, 540 Sexless entozoa, 534 Sexual generation, 540 Shook, effect of, in destroying nervous irritability, 393 SiEBOLD, on production of tsenia from cysticercus, 638 Sight, 492 apparatus of, 493. See Vision. Sinus terminalis, of area vasculosa, 665 Sinuses, placental, 624, 626 Skeleton, development of, 641 Skin, respiration by, 252 sebaceous glands of, 328 perspiratory glands of, 330 development of, 643 704 INDEX. Smell, 488 ganglia of, 382, 489 nerves of, 446, 489 injured by division of 5th pair, 454 Smith, Dr. Southwood, on cutaneous and pulmonary exhalation, 331 Solar plexus of sympathetic nerve, 516 Solid bodies, vision of with two eyes, 502 Sounds, of heart, 270 how produced, 271 vocal, how produced, 464 destroyed by section of inferior la- ryngeal nerves, 465 of spinal accessory, 475 Sounds, acute and grave, transmitted by membrana tympani, 508 Special senses, 478 Species, mode of continuation, 527 Spermatic fluid, 656 mixed constitution of, 560 Spermatozoa, 556 movements of, 558 foi'matlon of, 559 Spina bifida, 641 Spinal accessory, 474 sensibility of, 475 communication of, with pneumogas- trio, 475 influence of, on larynx, 475 Spinal column, formation of, 591, 641 Spinal cord, 379, 398 commissures of, 381 anterior and posterior columns, 381 origin of nerves from, 380 sensibility and excitability of, 403 crossed action of, 404 reflex action of, 408 protective action of, 413, 414 influence on sphincters, 414 efi^ect of injury to, 414 qn respiration, 441 formation of, in embryo, 591, 645 Spinal nerves, origin of, 380 Spleen, 208 Malpighian bodies of, 209 extirpation of, 211 Spontaneous generation, 527 Starch, 79 proportion of, in different kinds of food, 80 varieties of, 80-82 reactions of, 82 action of saliva on, 129 digestion of, 150 Starfish, nervous system of, 373 Steroorine, 160 Stereoscope, 503 St. Martin, case of gastric fistula in, 135 Strabismus, after division of motor oculi * communis, 451 of motor externus, 451 Striated bodies, 419 Sublingual glaud, secretion of, 125 Submaxillary ganglion, 514 gland, secretion of, 125 Sudoriparous glands, 330 Sugar, 83 varieties of, 83 composition of, 84 tests for, 84 fermentation of, 85 proportion in different kinds of food, 86 source and destination, 86 produced in liver, 200 discharged by urine in disease, 357 Sugar in liver, formation of, 200 percentage of, 202 produced in hepatic tissue, 203 from glycogenic matter, 204 absorbed by hepatic blood, 206 decomposed in circulation, 206 Sulphates, alkaline, in urine, 353 Sulphur of the bile, 181 not discharged with the feces, 197 Swallowing, 131 retarded by suppression of saliva, 133 -^by division of pneumogastric, 463 Sympathetic nerve, 514 ^ its distribution, 515 sensibility and excitability erf, 517 influence of, on special senses, 518 on pupil, 518 on nutrition of eyeball, 455 on nasal passages, 520 on ear, 520, 521 on temperature of particular parts, 521 reflex actions of, 524 Tadpole, development of, 592 transformation into frog, 594 Tsenia, 536 produced by metamorphosis of cys- ticercus, 538 single articulation of, 541 Tapeworm, 636 mode of genlration, 537 Taste, 481 nerves of, 483 conditions of, 485 inj ury of, by paralysis of facial nerve, 487 Taurine, 181 . Tauro-cholate of soda, 181 microscopic characters of, 179 Tauro-cholic acid, 181 Tears, 332 ~ their function, 332 Teeth, of serpent, 121 of polar bear, 122 of horse, 122 of man, 123 first and second sets of, 689 Temperature of the blood, 254 INDEX. 705 Temperature of different species of ani- mals, 255 of the blood in different organs, 262 elevation pf, after section of sympa- thetic nerve, 261, 521 Tensor tympani, action of, 506, 508 Tests, for starch, 82 for sugar, 84 for bile, 184 Pettenkofer's, 185 Testicles, 559 periodical activity of, in fish, 561 development of, 658 descent of, 658, 659 Tetanus, pathology of, 410 Thalami, optic, in rabbit, 383 in man, 418 function of, 419 Thoracic duct, 168, 170 Thoracic respiration, 441 Tongue, motor nerve of, 477 sensitive, 452, 456, 483 Trichina spiralis, 535 Tricuspid valve, 268. See Auriculo-ven- trioular , Triple phosphate, in putrefying urine, 362 Trommer's test for sugar, 84 interfered with by gastric juice, 143 Tuber annulare, 386,438 effect of destroying, 438 action of, 439 Tubercula quadrigemina, 382, 431 refiex action of, 435 crossed action of, 436 development of, 637, 638 Tubules of uterine mucous membrane, 615 Tufts, placental, 625 Tunica vaginalis testis, formation of, 660 Tympanum, function of, in hearing, 508 Umbilical cord, formation of, 631 I • withering and separation of, 689 J Umbilical hernia, 646 Umbilical vesicle, 596 in human embryo, 597 in chick, 604 disappearance of, 631 Umbilical vein, formation of, 669 obliteration of, 677 Umbilicus, abdominal, 592 amniotic, 600 • decidual, 617 Unilateral mastication, in ruminating animals, 127 Urate of soda, 347 its properties, source, daily qu3.ntity, &c., 348 Urates of potassa and ammonia, 348 Urachus, 647 Urea, 343 source of, 343 mode of obtaining, 344 45 Urea, conversion into carbonate of am- monia, 344 daily quantity of, 344 diurnal variations in, 345 decomposed in putrefaction of urine, 360 Uric acid, 347, 354 Urine, 349 general character g,ud properties of, 349 quantity and specific gravity, 350 diurnal variations of, 351 composition of, 352 reactions, 353 . interference with Trommer's test, 355 accidental ingredients of, 355 acid fermentation of, 359 alkaline fermentation of, 360 final decomposition of, 363 Urinary bladder, paralysis and inflam- mation of, after injury to spinal cord, 415 formation of, in embryo, 647 • Urosacine, 103 Uterus, of lower animals, 553 of human female, 554 mucous membrane of, 614 changes in, after impregnation, 615 involution of, after delivery, 634 development of, in foetus, 661 position of, at birth, t)63 Uterine mucous membrane, 614 tubules of, 615 conversion into decidua, 615 exfoliation of, at the time of delivery, 633 its renovation, '634 Valve, Eustachian, 681 of foramen ovale, 684 Valves, cardiac, action of, 268 cause of heart's sounds, 271 Vasa deferentia, formation of, 658 Vapor, watery, exhalation of, 71 from lungs, 242 from the skin, 252 Variation, in quantity of bile in different animals, 188 in production of liver-sugar, 202 in size of spleen, 208 in rapidity of coagulatioii of blood, 227 in size of glottis in respiration, 240 in exhalation of carbonic acid, 250 in temperature of blood in different parts, 262 in composition of milk during lac- tation, 337 in quantity of urea, 345 in density and acidity of urine, 350 Varieties of starch, 80 of sugar, 83 of fat, 86 706 INDEX. Varieties of biliary salts in different ani- mals, 182 Vegetable foed, necessary to man, 106 Vegetable parasites, 532 Vegetables, production of heat in, 256 absorption of carbonic acid and ex- halation of oxygen by, 49, 260 Vegetative functions, 58 Veins, 290 their resistance to pressure, 290 , absorption by, 165 action of valves in, 293 motion of blood through, 291 rapidity of circulation in, 294 omphalo-mesenteric, 666 umbilical, 669 vertebral, 672 Vense cavse, formation of, 673 position of, in foetus, 681 Vena azygos, superior and inferior, for- mation of, 673 Venous system, development of, 672 Ventricles of heart, single in fish and reptiles, 265, 266 double in birds and mammalians, 267 situation of, 268 contraction and relaxation of, 269 elongation during contraction, 275 muscular fibres of, 277 Veruix caseosa, 643 Vertebrata, nervous system of, 378 Vertebrae, formation of, 691, 641 Vesicles, adipose, 90 pulmonary, 235 seminal, 560 . Vesiculffl seminales, 560 formation of, 660 Vicarious secretion, non-existence of, 325 Vicarious menstruation, nature of, 325 Villi, of intestine, 163 absorption by, 164 of chorion, 609-611 Vision, 492 ganglia of, 382, 434 nerves of, 447, 493 apparatus of, 493 distinct, at different distances, 496 circle of, 499 Vision, of solid bodies with both eyes, 502 Vital phenomena, their nature and pecu- liarities, 54 Vitellus, 641 segmentation of, 587 formation of, in ovary of foetus, 663 Vitelline circulation, 665 membrane, 544 spheres, 587 Vocal sounds, how produced, 464 Voice, formation of, in larynx, 464 lost, after division of spinal acces- sory nerve, 476 Volition, seat of, in tuber annulare, 438 Vomiting, peculiar, after division of pneumogastrics, 473 Water, as a proximate principle, 69 its proportion in the animal tissues and fluids, 70 its source, 70 mode of discharge from the body, 71 Weight of organs, in comparative, newly born infant and adult, 689 White globules cif the blood, 220 . action of acetic acid on, 221 sluggish movement of, in circula- tion, 297 White substance, of nervous system, 368 of Schwann, 368 of spinal cord, 380 of brain, insensible and inexcitable, 417 Withering and separation of umbilical cord, after birth, 689 WollEan bodies, 655 structure of, 656 atrophy and disappearance of, 656, 657 vestiges of, in adult female, 662 Wtman, Prof. Jeffries, on cranial nerves of Eana pipiens, 449 fissure of hare-lip on median line, ■ 637 Yellow color, of urine in jaundice, 357 of corpus luteum, 579 Zona pellucida, 544 THE END. (LATE LEA & BLAIJ-CHAKD'S,) MEDICAL MD SURGICAL PUBLICATION. TO THE MEDICAL PROFESSION. .The prices on this Catalogue are those a^ which these books can generally- be furnished by booksellers, who can readily procure for their customers any which they may not have on hand. Where access to bookstores is not convenieut, I will forward them at these prices, /ree % mail, to any post office in the United States. In all cases the amount must accompany the order, as accounts are opened only with dealers ; I assume no risks of the mail, either on the money or the books, and can supply nothing but my own publications. Gentlemen desirous of purchasing will, therefore, find it more advantageous, to deal with the nearest bookseller whenever practicable, or to send orders through tfieir merchants visit- ing the larger cities. HENRY 0. LEA. fHiLADBLFHiA, February, 1866. *^,* Just issued, a revised Illustrated Catalogue of Medical and Soientifio Publications, forming an octavo pamphlet of 80 large pages, containing specimens of . illustrations, notices of the medical press, &c. &c. It has been prepared with- out regard to expense, and will be found one of the handsomest' specimens of typographical execution as yet presented in this country. Copies will be sent to any address, by mail, free of postage, on receipt of nine cents in stamps. Catalogues of publications in miscellaneous and educational literature forwarded on application.. Gy The attention of physicians is especially solicited to the following important new works and new editions, just issued or nearly ready : — Ashton on the Rectum, new edition, * .. . Seepage 3 Brintun on the Stoaiach, Bennett on i he Uterus, fiixth edition Bowman's Medical Chemistry, fourth edition, Bumstead on Venereal, second edition, Barclay on Medical Diagnosis, third edition, • Brande and Taylor's Chemistry, . . . Dalton's Human Physiology, 3d edition, . . Dunglison's Medical Dictionary, a revised edition, 1S6S, • Ellis' Formulary, new edition, . . . ^ . Krichsen's System of Surggry, a revised edition, Flint's Practice of Medicine, now ready, Gross's System of Surgery, third edition,' Gray's Anatomy, Descriptive and Surgical, 2d edition, Hillier on Skin Diseases^ Hamilton on Fractures and Dislocations, second*edition, Hodge's Obstetrics, f ereira's Materia Medica, condensed edition, preparing, Roberts on Urinary Diseases, nearly ready, Parrish's Practical Pharmacy, a new edition, . Smith on Consumption, Slille's Therapeutics and Materia Medica, second edition, Wilson on the. Skin, filth edition, . . . . , Win slow on the Brain and Mind, second edition, West on Children, fourth edition, . ' . 4 4 4 5 3 6 11 la 13 14 IS IB 17 18 18 19 ■M 24 25 26 27 31 32 32 THE AMERICAN JOURNAL OF THE MEDICAL SCIENCES AND THE MEDICAL NEWS AND LIBEAEY, For FITE D01 has lendt red necessary — - London Med. Times and Gazette, Nov. 29, 1663. Nor is any thing huiried over, anything shirked ; open the book where; you will,|yuu find the same careful treatment of the subject manifested, ana the beat process for the attaiLment of the particular ob- , BY THE SAIUB AUTHOR. INTRODUCTION TO PRACTICAL CHEMISTRY,. INCLUDINO ANA- LYSIS .Third American, from the third and revised London edition. With numerous illus- trations' In one neal vqI., royal 12ino., e:Ura qloih S-i !i5 This iavorite little manual has received a very thurungh and careful revision at the hands of a competent editor, and will be found luUy brought up to the present condiiion of chemical science. Many portions have been fewrittcn, the subjects ol the blow-pipe and volumetric analysis have re- ceived special altentiun, and an addilional chapter! has been appended, .Siudenls uf practical cheni- ulry will, therefore find it, a!> here ofore, a most ci^nvenicnt pud condensed test-book and guide in the opeiations of thelaboraiory. ' ' ' I AND SCIENTIFIC PUBLICATIONS. BUMSTEAD (FREEMAN J.) M. D., Lecturer on Venereal DiaeaaeB at the College of Puygicians and Surgeons, New York, 4;o. THE PATHOLOGY AND TEEATMENT OP VENEREAL DISEASES, including the results of recent investigations upon the subject. Second edition, thoroughly re- viR(:d and much improved. With illustratiuns on wood. In one very handsome octavo volume, of about 700 pages $5 00 {Just Issued.) By far the moBt valuable contribution to this par- ticular branch of practice that has seen the light wUbin the last score of years. His clear and accu- rate descriptions of the various forms of venereal disease, and especially the methods of treatment he proposes, are worthy of the highest encomium. In tlieae respects it is better adapted for the assistance u- f every-day practitioner than any other with which we are acquainted. In variety of methods proposed, m minuteness of direction, guided by care- (ttl ducriraination bf varying forms and compUca tiona, we w^ite down the book as unsurpassed. It is a work which should be in the possession of every pj-actitioner.- Cfeicag-o Med. Journal. Nov. 1881. Tne fencing admirable volume comes to us, em- bracing the^whole subject of syphilology, resolving many a doubt, correcting and confirming many an entertained opinion, and in our estimation the best, eompletest, fullest monograph on this subject in oar language. ■ As far as the.author*s labors themselves are concerned, we feel it u duty to say that he has not only exhausted his subject, but he has presented to us, without the slightest hyperbole, the best di- gested treatise on these diseases in our language He has carried its literature down to the prestnt moment, and has achieved his task in a manner which cannot but redound to his Qz&iit.— British American Journal^ Oct. 1861. We believe this treatise will come to be reerarded as high authority in this branch of medicsl practice, and we cordially commend it to the favorable notice of ourbrethren in the profession. For our own part, we candidly confess that we have received aany new ideas fromits perusal, as well as modified many views which we have long, and, as we now thinK erroneously entertained on the subject of syphilis. To sum up all in a few words, this book is one which no practising physician or medical student can very well afford to do without. — American Med Times, Npv.a, 1861. The whole work presents a complete history of venereal diseases, comprising much interesting and vaBl,uable material that hasbeen spread through intd- ical journals within the last twenty years— the pe- riod of many eyperiments and investigations on the subject— the whole carefully digested b^ t^e aid of the author^s extensive personal experience, and offeied to the profession m an admirable form. Its completent^ss is secured by good plates; which are especially full in the anatomy of the genital organs. W^e have examined it with great satisfaction, and congratulate the medical profession in America on the nationality of a work ihat may fairly be sailed original. — Berkshire Med. Journal^ Dec IbSL. One thing, however, we are impelled to say, that we have met with no other book on syphilis, in the English language, which gave so full, clear, and impartial views of the important subjects on wiiiuh it treats. We cannot, however^ refrain from ex- pressing oursatisfaction with the full and persiiicu- ous manner in which the subjecthas been presented, and the careful attention to minuie details, so use^ ful — not to say indispensable — inu practical Lreatise. In conclusion, if we may be pardoned the use of a phrase now become stereotyped, bat which we here employ in all seriousness and sincerity, we do not hesitate to express tite opinion Ihat Dr. BuinsteadU Treatise on Venereal Diseases is a '* work without which no medical library will hereafter be consi- dered complete." — Boston Med. and Surg. Journal Sept. 5, 1861. BARCLAY (A. W.), M. D., Assistant Physician to St. George's Hospital, &c.; A MANUAL OF MEDICAL DIAGNOSIS; being an Analysis of the Signe and Symptoms of Disease- Third American from the second and revised London edition. Id one neat octavo volume, extra cloth, of 451 pages. S3 50 (Just Issued ") The demand for another edition of this work %hows tbat the vacancy which it attempts to sup* ply has been recognized by the profession, and that the efforts of the author to meet the want have Deen successful. The revision which it has enjoyed will render it better adapted than before to afford assistance to the learner intheprosecutionof his studies, and to thupractilioner who requires a convenient and accessible manual for speedy reference in the exigencies of his daily duties. For this tatter purpose it-t complete and extensive Index renders it ei^pecially valuable, offering facilities for immediately turning to any class of symptoms, or any variety of disea!$e. We hope the volume will have an extensive c'ii- culation, not among students of medicine only, but practitioners also. They will never regret a faith - ful study of itspages. — Cincinnati Lancet. An important acquisition to medical litentur^. The task of composing such a wOtk is neither an easy nor a light one; but Dr.Burclay has performed it in a manner which meets our most unqualified approbation. He is no mere theorist; he knows his work thoroughly, and in attempting to perform it, hasnot exceeded his powers. — British Med. Journal. We venture to predict that the work will be de- servedly popular, and soon become, like Watson'i^ Practice, an indispensable necessity to the practi- tioner. — N.A.Med Journal. , An inestimable work of reference for the young practitioner and student. — Nashville Med. Journal . It IS a work of high merits both from the vast im-, por',ance of the sabject upon which it treats, au(l' also from the real ability displayed in ■t'lt elabora- tion. In conclusion, let us bespeak for this volume that attention of every SLudent of our art which it 80 richly deserves — that place in every meaical library which it can so well adorn.' Peninsul^f Medical Journal. BARTLETT (EUISHA), M. D, THE HISTORY, DIAGNOSIS, AND TREATMENT OF THE FEVERS OF THE UNITED STATES. A neW and revised edition. By Alonzo Clark , IVf. D.j Prof, of Pathology and Practical Medicine in the N. V. College of Physicians and Surgeons, dec. lit one octavo volume, of six hundred pages, extra cloth. Price $4 25. BROWN (ISAAC BAKER), Suigeon- Accoucheur to St. Mary's Hospital, &o. ON SOME DISEASES OF WOMEN ADMITTING OF SURGICAL TREAT- MENT. With handsome illustrations. One vol. Svo., extra cloth, pp. 276. $160. Mr. Brown lias earned f'lr himBelf a high reputa- tion inthe operative treatment of sundry diseases and injuries to which females are peculiarly- subject. Wc can truly say of his work that lit is an important addition to obstetrical literature. The operative suffffestions and contrivances which Mr. Brown de- scribes, exhibit much practical rfgacity and skUl, and merit the careful attentioib of every saiKeon. accoucheur. — Association Journal. We have no hesitation in recommending this book tn tje careful attention of all surgeoiia who make female coniplaints a part of their study and practice —DuUin i^uarteriy Journal. 6 HENRY C. LEA»S MEDICAL BRANDE {WM, T..1 D, C. Of her MaJJBsty's Mint, &c. ■ A»D ALFKED S. TAYLOR, M. D., F. R. S. ProfesBurof GhemiBtry and Medical Juriaprudencein Guy'aHoBpital. CHEMISTRY.. In one handsome 8vo. volume of 696 pages, extra cloth. $4 50. *' Having been engaged in teaching Chemistry in this Metropolis, the one for a period of forty, and the other for a period of thirty years, it has appeared to us that, in spite of the number of books already existing, there was room, for an additional volume, which shtuld be especially adapted for. the ute of students. In preparing such a volume for the press, we have endeavored to bear in mind, that the student in ihe present day lias much to learn, and but a short lime at his disposal for the acquisition of this learning."— ^Authors' Preface. In reprinting this volume, its passage through the press has been superintended by a competent chemtst, who has sedulou^ly endeavored to secure the accuracy so necesi-ary in a work of this nature. No notes or additions have been introduced, bul the publishers have been favored by tlie authors with some corrections and revisions of the iitsl twenty-one chapters, which have been duly inserted. In so progressive a science as Chemistry, the, latest work always has the advantage of presenting the subject as modified by the results of the latest investigations and discoveries. That this advan- tage has been made the most of, and that the work possesses superior attractions arising from its clearness, simplicity of style, afld lucid arrangement, are manifested by the unanimous testimony of the Engtis^h medical press. , It needs uo great sagacity t9 foretell that this book will be, literally, the Handbook in Chemistry of the student and practitioner. For clearness of language, accnracy of description, extent of information, and freedom from pedantry and mysticism of modern chemistry, no other text -book comes into< competition with it. The result is a work which for fulness of matter, for lucidity of arrangement, for clearness of Btyle, is as yet without a rival. And long will it be without a rival. For, although with the necessary advance of chemical knowledge addenda will be re- qaired, there will be little to take away. The funda- mental excellences of the book will remain, preserv- ing it for years to come, what it now is, the best guide to the study of Chemistry yet given to the world. — London Lancet, Dec. 20,1862. Most assuredly, time has not abated one whit of the fluency, the vigor, and ihe clearness with which they not only have composed the work before ua, but have, 80 to say, cleared the ground for it, by hittin^right and left at the affectation, mysticism, and obscurity which pervade some late chemical treatises. Thus conceived, and worked oat in the moat sturdy, com- mon sense meihodj this boofegives, in the cl'earest and most summary method possible, all the facts and doc- trinesof chemistry, with more especial reference to the wants of the medical student. — London Medical Times mid Gazette, Mov. 29, 1862. If we are not very much mistaken, this book will occupy ^a place which none has hitherto held among chemises ; for, by avoiding the errors of previous au- thors, we have a work which, for its size, is certainly the mq^t perfect of any in the English language. There are sevei'al points to be noted in this volume which separate it ■widely from any of ita compeers- its wide application, not to the medical student only, nor to the student in chemistry merely, but to every branch of science, art, or commerce which is in any way connected with the domain of chemistry. — Lon- donMed. Seview, Feb. 1863. BARWELL (RICHARD,) F- R. C. S., Assistant Surgeon Charing Cross Hospital, &,c. A TREATISE ON DISEASES OF THE JOINTS. Illustrated with engrav- ings on wood. In one very handsome octavo Volume, of about 500 pages, extra cloth ; $5 00. ing^ and faithful delineations of disease. — London Med; Times and Gazette^ Feb. 9, lti6l. At the outset we may state that the work Is worthy of much praise, and bears evidence of much ithoughtful and careful inquiry, and here and there ■of no siight originality. We have already carrud ttiie notice further than we intended to do, but not to the extent the work deserv<:8. We can only add, that the perusal of itl^as afl^rdcd us gi'eat pleasure. The author has evidently worked very hard at his subject, and his investigations into the Physiology and Pathology of Joints have been carried on in a manner which entitles him to be listened to with attention and respect. We must not omit to men- tion the very admirable plates with which th& vo- lume iseoriched. We seldom meet with suchstnk- This volume;wilt be welconfedj as the record of much honest research and careful investigation into the nature and treatment of a' most imfjortant class of disorders. We cannot conclude this notice of a valuable and ustful' book without calling attention to the amount o{bon&Jide work it contains. It is no slight matter for a volume to show laborious iuvea- tigation., and at the same time original thought, on the part uf its auDhor, whom v e may congratulate on the successful completion of his arduous task.— London Lancet, March.9, 1861. CARPENTER (WILLIAM BJ, M. D., F. R. S., «£C., Examiner in Physiology and Comparative Anatomy in the University of London. THE MICROSCOPE AND ITS EEVELATIONS. With an Appendix con- taining the Applications of the JVIicroscope to Clinical Medicine, &c. By F. G. Smith, M. D. Illustrated by four hundred and Ihirly-four beautiful engraving* on wood. In one large and very handsome octavo volume, of 724 pages, extra cloth, $5 25. The great importance of the niicroscope as a means of diagnosis, and the number oi microsco- pists who are also physicians, have induced the American publishers, with the author's approval, to ■add an Appendix, careluily prepared by Professor Smith, on the applications of Ihe inf-lrument to clinical medicine, together with an account of Auierican Microscopes, the:"/ modifications and accessories. This portion of the work ,ia illustrated with nearly one hundred wood-cuts, and, it is hoped, will adapt the volume more particularly to the use of the American student. Those who are acquainted wiih Dr. Carpenter's revious writings on Animal and Vegetable Physio- ogy, will fully understand how vasla store of know- lt„,, . - ledge he is able to bring to bear upon su comprehen< dsive a subject as the revelations of the fnicroscoj;>e ; and even those who have nu pievious acquaintance with Che construction or uses of 'tHis instrument, will find abandanceof information conveyed in clear and siinple language. — Med. Ttmes and Gazette. The additions by, Prof. Smich g^iye it a positive claim upnn the profession', for which v/e uoubt not he will rebtiive their sincere thanks. Indeed, we know not where the student of medicint will find such a eouiplete and satisfactory collection of micro- BCtipic facis bearing upon physiology and practical medicine us Is contained in Prof. Smith's appendix; and this of itself, it seems to us, is fully worth the cost of; the volume. — Louisville Medicai Heviev, ANU SCIJSKTIFIC F UJBLIG ATIONS. CARPENTER (WILi.lAM B.), M. D., F. R. S., Examiner in Phyniology and Comparative Anatomy in the University of London. PRINCIPLES OF HUMAN PHYSIOLOGY; with their chief applications to Psychologyj PatholMy, Therapeutics, Hygiene, and Forensic Medicine. A new American, from the last and revised London edition. Withnearlythree hundrediUustrations, Edited, witha4di- ''^**°-^' -U ^J^^^^^^ Gurnet Smith, M. D., Professor of the Institutes of Medicine in the Pennsyl- vania Medical College, &c. In one very large and beautiful octavo volume, oi about nine hundred large pages, handsomely printed, extra cloth, $5 50 For upwards of thirteen years 0r. Carpenter's work has-been considered by the profession gene- ' rally, both m this country and England, as the most valuable compendium on the subject of physiology in our language . This distinction it owes to the high attamments and unwearied industry of its accom- plished author. The present edition (which, like the ^ast American one, was prepared by the author him- selt), IS the result of such extensive revision, that it may almost be considered* a new work. We need hardly say, in concluding thisbrief notice, that while the work is indispensable to every student of medi- cine in this country, it will amply repay the practi- tioner for its perusal by the interest and value of its contents.^£os£on Med. and Surg. Journal, Thiais a standard work— the text-book used by all medical students who read the English language. It has passed through several editions in order to keep pace with the rapidly growing science of Phy- siology. Nothing need be said in its praise, for its merits are universally known ; we have nothing to \ say of its defects, for they only appear where the science of which it treats is incomplete. — Western Lancet. The most complete exposition of physiology which any language can at present give. — Brit, and For. ilSed.-Ckirurg. Review. The greatest, the most reliable, and the best book oa the subject which we know of in the English language. — Stethoscope . To eulogize this great work would be superfluous. We should observe, however, that in this edition the author hsis remodelled a large portion of the former, and the editor has added much matter of in- terest, especially In the form of illustrations. We may confidently recommend it as the most complete work on Human Physiology in our language.— Southern Med. and Surg. Journal. The most complete work on the science in our language. — A?n. Med. Journal. The most complete work now extant in our lan- guage. — N. O. Med. Register. The best text-book in the language on this ex tensive subject. — London Med. Ttmes. A complete cyclopsedia of this branch of scienott. —N. Y. Med. Times. The profession of this country, and perhaps also of Europe, have anxiously and for sometime awaited the announcement of this new edition of Carpenter's Human Physiology, His former editions have for many years been almost the only text-book on Phy- siology in all our medical schools, and its circula- tion amon^ the profession has been unsurpasBed by any work m any department of medical science. It is quite unnecessary for us to speak of this work as its merits would justify. The mere an- nouncement of its appearance will afford the highest pleasiure to every student of Physiology, while its perusal will be of infinite service in advancing physiological science. — Ohio Med.andSurg. Journ BY THE SAME AUTHOR. ELEMENTS (OR MANUAL) OF PHYSIOLOGY, INCLUDING PHYSIO- LOGICAL ANATOMY. Second AirieriCati, from a new and revised London edition. With one hundred and ninety illustrations. In one very handsome octavo volume, leather, pp. 566. $4 00. In publishing the first edition of this work, its title was altered from that of the London volume, by the substitution of the word " Elements" for that of " Manual," and with the author's sanction toe title of " Elements" is still retained as being more expressive of the scope of the treatise. BV THE SAME A.CTHOK. PRINCIPLES OF COMPARATIVE PHYSIOLOGY. New Atnerican, from the Fourth and Revised London edition. In one large and handsome octavo volume, with over three hundred beautiful illustrations, pp. 752. Extra cloth, $5 00 This book should not only be read bat thoroughly studied by every member of the profcBBion. None are too wise or old, to be benefited thereby. But egpeoially to the younger class would we cordially commend it as best fitted of any work in the English language to qualify them for the reception and com- prehension of those truths which are daily being de- veloped in physiology. — Medical Counsellor. Without pretending to it, it is an encyclopedia of (he subject, accurate and complete in all respects— a truthful reflection of the advanced state at which the science has now arrived. — Dublin Quarterly Journal of Medical Science. A truly magnificent work — in itself a perfect phy- siological BivAy.— Ranking' s Abstract'. This work stands without its fellow. It is one few men in Europecould have undertaken; it is one no man, we believe, could have brought to so sno- cessful an issue as Dr. Carpenter, ft regnired'for its production a physiologist at once deeply read in the labors of others, capable of taking a general critical, and unprejudiced view of those labors, an ofthiswork. — N Y. Journal of Medicine. A veritable pebdiatric encyclopsedia, and an hunoi 10 American medical literature. — Ohio Medical and Surgical Journal. Wefeel persuaded thatthe American medical pro- fession will 80(m regard it not only aa a very good, but as the very best "Practical Treatise im tht Diseases of Ghildren." — American Medical, Journal In the department of infantile therapeutics, the work of Dr. Gondie is considered one ot the best which has been published in the English language, ^Tke Stethoscope. We pronounced the first edition to be the best work on the diseases of children in the English language, and, nut withstanding all that has been published, we still regard it in that light. — Medical Examiner. The value ofworks by native authors on the dii- eases which the physician is called up^n to eombat, will be appreciated by all ; an'i the work of Dr. Con- die has gained for Itself the character of a safe guide tor students, and a useful work for consultation by those engaged in practice,. — N. Y.. Med Times. This is the fourth edition of this deservedly popu- lar treatise. During the interval since the last edi- tion, It has been subjected to a thorough revision by the author; and all new observations in the pathology and therapeutics of children hav« been included in the present volume. As we said btfore, we do not know of a better book on diseases of chil- dren, and to a large part of its recommendations we yield an unhesitating concurrence. — Buffalo Med. ToumaJ^ . Perhaps the mostfuU and complete work now be- lOre the profession of the United States; indeed, we iuay aay in the English language. It is vastly supe- rior to most of its predecessors. —Tronsylwawa Med, Journal CHRISTISON (ROBERT), M. D., V. P. R. S. E., fcc. A DISPENSATORY; or, Commentary on the Pharmacopoeias of Great Britain and th« United Stales ; comprising the Natural History, Descriptiou, Chemistry, Pharmacy, Ac- tions, Uses, and Doses of the Articles of the Materia Medica. Second editioii, revised and im- proved, with a Supplement containing the most important New Remedies. With uopiank;^ddi- tions, and two hundred and thirteen large wood-engravings. By R- Eslesfeld Geifehth, M. D. In one very large and handsome octavo volume, extra cloth, of over 1000 pages. W CO COOPER (BRANSBY B.), F. R. S. LECTURES ON THE PRINCIPLES AND PRACTICE OF SURGERY- In one very large octavo volume, extra cloth, of 750 pages. $2 00. COOPER ON THE ANATOMY AND DISKASEg OF THE BREAST, with twenty-five Mlscellane- oas and Surgical Papers. One large volume, im- perial 8vo,, extra cloth, with 3S2 figures, on 36 plates. S3 00. COOPER ON THE STRUCTURE AND DIS- EASES OF THE TESTIS, AND ON THE THYMUS GLAND. One vol. imperial 8vo., ex- tra cloth, with 177 £gures on 29 plates, f 2 50. CLYMEB ON FEVERS: THEIR DIAGNOSIS. PATHOLOGY, AND TREATMENT. In OM octavo volume, leather, of 600 pages. SI 73. COLOMBAT DE L'ISERE ON THE DISEASES OF FEMALES, and on the special Hygiene of their Sex. Translated, with many Notes and Ad- ditions, by C. D. Meiss, M.D. Second edition, revised and improved. In one large voluhie, oc- tavo; leather, with nnmerous wood-cuts. pp. 720. S3 73. CARSON (JOSEPH), M. D., Professor of Materia Medica and Pharmacy in the University of Pennsylvania, SYNOPSIS OF THE COURSE OF LECTUREis ON MATERIA MEDICA AND PHARMACY, delivered m the .University of Peanisylvania. With three Lectures on the Modus Operandi of Medicines. Third edition, revised. In one handsome octavro volume. »2 50. CURLING (T. B.), F. R.S., Surgeon to the London Hospital, President of the Hunterian Society, &c. A PKACTICAL TREATISE ON DISEASES OF THE TESTIS, SPERMA- TIC CORD, AND SCROTUM. Second American, from the second and enlarged English edi- Moa. In one handsome octavo volume, extra cloth, with nnmerous illustrations, pp. 420. S2 00 AND SCIENTIFIC PUBLICATIONS. 9 CHURCHILL (FLEETWOOD), M. D., M. R. I. A. ON THE THEORY AND PRACTICE OF MIDWIFERY. A new American from the fourth revised and enlarged London edition. With Notes and Additions, by D. Francis OoNDiE, M. D., author of a "Practical Treatise on the Diseases of Chil4ren," &c With 194 illustrations. In one very handsome octavo volume of nearly 700 large pages, extra cloth, $4 00. This virork has been so long an established favorite, both as a text-boob lor the learner and as a reliable aid in consultation tor the practitioner, that in presenting a new edition it is only necessary* to call attention to the very extended improvements which it has received Having had the benefit of iwo revisions by the author since the last American reprint, it has been materially enlarged, and Dr. Churchill's woU-known conscientious mduslry is a guarantee that every portion has been tho- roughly brought up with the latest results of European investigation in all departments of the sci- ence and art of obstetrics. The recent dale of the last Dublin edition has not left much of novelty for the American editor to introduce, but he l?as endeavored to insert whatever has since appeared, together with such matters as his experience has shown him would be desirable fur the American student, including a large number of illustrations Wiih the sanction of the author he has added in the form of an appendix, some chapters from a little "Manual for Midwivei^ and Nurses," re- cently isi^ued by Dr. Churchill, believing- that the details there presented can hardly fail to prove nt advantage to the junior practitioner. The result of ajj these auditions is that the work now con- tains fully one-half more matter than the last American edition, with nearly one-half more illus- trations, so that notwithstanding tlfe use of a smaller type, the volume contains almost two hundred pages more than before. No effort has been spared to securS an improvement in the mechanical execution of the work equal to that which the text has received, and the volume is confidently presented as one of the handsomest that has thifs far been laid before the American profession; while the very low price at which it is offered should secure for it a place in every lecture-room and on every office table. A better book in which to learn these important points we have not met than Dr. Churchill's. Every pn^e of it is full of ihBtruction ; the opinion of all wntefs of authority is given on questions of difH- cuity, as well as the directions and advice of the learned auttior himself, to w^hich he adds the result of statiBticalintiuiry, putting statistics in tiieir pio per place and giving them their due weight, and no more. We have never read a book more free from profesBional jealousy than Dr. Churchill's. It ap- pears Co be written with the true design of a book on medicine, viz: to give all that is known on the sub- Jeet of which he treats, both theoretically and prac- tically, and to advance such opinions of his own as he J)elieves will benefit medical science, and insure the safety of the patient. We have said enough to convey to the profession that this book of Dr. Chur- chill's is admirably suited for a book. of reference for the practitioner, as well asAtextTbook for the student, and we hope it may be extensively pur- chased amongst our readers. To them we most strongly recommend it. — Dublin Medical Press To bestow praise on a book that has received such marked approbation would be superfluous. We need enly say, therefore, that if the first edition was thought worthy of a favorable reception by the medical public, we can confidently affirm that this will be found much more bo. The lecturer, the practitioner, and the student, may all have recourse to its pages, and derive from their perusal much in- terest and instruction in everything relating to theo- retical and practical midwifery.— i)«6K» Quarttrly Journal of Medical Science. A work of -very great merit, and such as we can confidently recommend to the study of every obste- tric practitioner -T-I-ottrfonilfedicoZ Gazette. Few treatises y^ill be found better adapted as t text-bo&kfor the student, or as a manual for tfai frequent consultation of the young practitioner.- American Medical Journal. Were we redaced to the necessity of having bu^ me work on midwifery, and permitted to choosef ve would unhesitatingly take Churchill. — Western Med. and Surg. Journal. It is impossible to conceive a more useful and ilegant manual than Dr. Churchill's Practice of Midwifery. — Provincial Medical Journal. Certainly, in our opinion, the very best work on ^le subject which exists. — N. Y. Annalist. No work holds a higher position, or is more de- serving of being placed in the hands of the tyro, the advanced student, or the practitioner. — Medical Examiner Previous editions have been received witli mark- ed favdr, and they deserved it; but this, reprinted from a very late Dublin edition, carefally revised and brought up by the author to the present' timn, does present an anusualLy accurate and able expo- sition of every important particular embraced in the departmentof midwifery. * * The clearnessj directness, and precision of its teachings, together with the great amount of statistical research which its text exhibits, have served to place it already in the foremost rank of works in this department of re- medial science. — Y Meit and Sv.rg Tournal. In our opinion, it forms one o( the best if not t e very best text-book and epitome of obstetric scieuco which we at present possess in the English lan- guage.— ilfon(Aiy./o«r»oi of Medical SeteiMd: The clearness and precision of style in which it ia written, and the greatamount(ff statistical research which it contains, have served to place it in the first rank oT works in this departmentof medical science. — JV. Y. Journal of Medicine. This is certainly the most perfect system extant. It ia the best adapted for the purposes of a text- 30ok, and that which he whose necessities confine lim to one book, should select in preference to all others. — Soutkern Medical and Surgical Journal. BY THE SAME AUTHOR ON THE DISEASES OF INFANTS AND CHILDREN. Second American Edition, revised and enlarged by the author. Edited, with Notes, by W. V. Keatins, M. O. In one large and handsome voiume, extra cloth, of over 700 pages. $4 50. In preparing this work a second time for the American profession, the author has spared no labor in giving it a very thorough revision, introducing several new chapters, and rewriting others, while every portion of the volume has been subjected to a severe scrutiny. The efforts of the American editor hate been directed to supplying such information relaiive to matters peculiar to this country as might have escaped the attention of the author, and the whole may, there- fore be safely pronounced one of the most complete works on the subject accessible to the Ame- rican Profession. By an alteration in the size! of the page, these very extensive additions have been accommodated without unduly increasing' the size of the work. BY THE SAMS AUTHOB.. ESSAYS ON THE PUERPERAL FEVER, AND OTHER DISEASES PE- CIILIAR TO WOiWEN. Selectedfromthe writings of British Authori- previous to the clos*: of tbeSiffhteenth Century. IaoneneBtocUvovo[ame,extracloth,olabout 450 pages. S2 50. 10 HENRY C. LEA'S MEDICAL CHURCHILL (FLEETWOOD), M. D., M. R. I. A., &o ON THE DISEASES OF WOMEN; iifcluding those of Pregnancy and Child- bed. A new American edition, revised by the Autiior. With Notes and Additions, by D Fram- cis CoNDiE, M. D., author o) " A Practical Treatise on the Uiseases of Children." With nume- rous illustrations. In one large and handsome octavo volume, extra cloth, of 768 pages. $4 00. This edition of Dr. Churchill's very popular treatise may almost be termed a new work, so thoroughly has he revised it in every portion. It will be fouifd greatly enlarged, and completely brought up to the most recent condition of the subjeot, while the very handsome series of illustra- tions introduced, representing such pathologigal conditions as can be accurately portrayed, present a novel feature, and afford valuable assistance to the young practitioner. Such additions as ap- peared desirable for the American student have been made by the editor, Dr. Condie, while a marked improvement in the mechanical execution keeps pace with the advance in all other respects which the volume has undergone, while the price has,been kept at the former very moderate rate. it compriBes, unquestionablj^, one of the meet ex- act and comprehensive expositions of the present state of medical knowledge in respect to the diseases of women that has yet been published. — Am. Joum, Med. Sciences. This work is the most reliable which we possess on this subject; and is deservedly popular with the profession. — Charleston Med. Joumaly July, 1857. We know of no author who deserves that appro- bation, on " the diseases of females," to the same extent that Dr. Churchill does. His, indeed, is ths only thorough treatise we know of on the subject : and it may be commended to practitioners and stu- dents as a masterpiece in its particular department, — Tkt Western Journal of Medicine and Surgery. As a comprehensive manual for students, or i work of referedce for practitioners, it surpasses any other that has ever issued on the same subject from the British press. — Dublin Quart. Journal. DICKSON (S. H.), M. D., Professor of Practice of Medicine in the Jefferson Medical College, Philadelphia. ELEMENTS OF MEDICINE; a Compendious View of Pathology and There- peutics, or the History and Treatment of Diseases. Second edition, revised. In one large and handsome octavo volume, of 750 pages, extra cloth. $4 00. 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 needed— -an elementary manual of praclice, which should present the leading principles of medicine with the practical results, in a condensed and perspicuous manner. Disencumbered of unnecessary detail and fruitless speculations, it embodies what is most requisite for the student to learn, anci at the same time what the active practitioner wants When obliged, m the daily calls of his profession, to refresh his memory on special points. The clear and attractive style of the author renders the whole easy of comprehension, while his long experience gives to his teacbiiigs an authority every- where 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 de- voted 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. DRUITT (ROBERT), M.R.C.S., &c. THE PRINCIPLES AND PRACTICE OF MODERN SURGERY. A new and revised American from the eighth enlarged and improved London edition. Illustrated with fQur hundred and thirty-two wood-engravings. In one very handsomely printed octavo volume of nearly 700 large pages, extra cloth, $4 00. A work which like Druitt's Surgery h«s for so many years mamtamed the position of a lead- ing favorite with all classes of the profession, needs no special recommendation to attract attention to a revised edition. It is only necessary to state that the author has spared no pains to keep the work up to its well earned repulalion of presenting in a small and convenient compass the latest condition of every department of surgery, considered both as a science and as an art; and that the services of a competent American editor ^ve been employed to introduce whatever novelties may have escaped the author's attention, or may prove of service to the American practitioner, As several editions have appeared in London since the issue of the last American reprint, the volume has had the benefit of repeated revisions by the author, resulting in a very thorough alteration and improvement. The extent of these additions may be estimated from the fact that it now contains about one-third more matter than the previous American edition, and that notwithstanding the adoption of a smaller type, the pages have been increased by about one hundred, while nearly two hundred and fifty wood-cuts have been added to the former list of illustrations. A marked improvement will also be perceived in the mechanical and artistical execution of the work, which, printed in the best style, on new type, and fine pap^r, leaves little to be desired as regards external finish; while at the very low price affixed it will be found one of the cheapest volumes accessible to the profession. This popular volume, now a moat comprehenaive work on surgery, has undergone many corrections, improvemeuts, and additions, and the principles and the practice of the art have been brought down to the latestrecordandobservatiou. Of the operations in surgery it is impossible to speak too highly. The descripiions are so clear and concise, and the illus- trations BO accurate and numerous, ttiat the student can have no difficulty, with instritmentinhand, and book by his side, over the dead body, in obtaining a proper knowledge andisufficient tact in this much neglected department of medical education. — British and Foreign Medico-Chirurg , Review^ Jan. 1S60 In the present edition the author has entirely re- written many of the chapters, and has incorporatsd the various improvements and additions in modern •urgery. On carefully going over it, we find that nothing of real practical importance has been omit- ted ; it presents a faithful epitome of everything re- lating t ) surgery up to the present hour. It is de- servedly a popular manual, both with the student and practitioner.— Loff<2on Lancets Nov. 19, 1859. In closing this brief notice, we recommend as cor- diallv as ever this most useful and comprehensive hand-bouk. It must prove a vast assistance, not only to the student of surgery, but also to the busy practitioner who may not have the leisure to devote himself to the study of more lengthy volumes.— London Med. Times and Gazettgj Oct. 23, 1859. In a word, this eighth edition of Dr. Bruitt'i Manual of Surgery is all that the surgical student or practitioner could desire. — Dublin Quarttrif Journal of Mid. Sciences, Nov. 1859. AND SCIKNTIFIC PUBLICATIONS. 11 DALTON, JR. (J. C), M. D. PrdfesBor of PhyBiology in the College of FhysicianB, New York. A TREATISE ON HUMAN PHYSIOLOGY, designed for the use of Students and Practitioners of Medicine. Third edition, reviped, with nearly three hundred illustrations on wood. In one very beautiful octavo volume, of 700 pages', exira cloth, S5 ^5, {Just Issued.) The rapid demand for another edition of this work sufficiently shows that the author has suc- ceeded in his eflbrls to produce a text-book of standard and permanent value, embodying within a moderate compass all that is definitely and positively known within the domain of Human Pfaysiology. His higblk-eputation as an original observer and investigator, is a guarantee that in again revising it he has introduced whatever is necessary to render it thoroughly on a level with the advanced science of the day, and this has been accomplished without unduly increasing the size of the volume. No exertion has been spared to maintain the high standard of typographical execution which has rendered this work admittedly one of the handsomest volumes as yet produced in this couniry. Itwilibeeeen, therefoxe, that Dr.Dalton'e best efforts have been directed towards ptjifecting his work. The.additiong are marked by the same fea- tures which characterize the remainder of the vol- ume, and render it by far the most desirable text- K„„l, _l ^:-l ^._ .1 j_ ^L. 1 ... _^.^L- book on physiology to place in the hands of the student which, so far as we are aware, exists in the English language, or'perhgps in any other. We therefore have no hesitation in recommending Dr. Dal ton's book for Che claBseB for which it is intend- ed, satisBed as we are ihat it is better adapted to their uec than any other work of the kind to which they have access. — American Journal of the Med, jSlctenc«5, April, 1861. * It is, therefore, no disparagement to the many books upon physiology, most excelien'; in theirday, to say that Daiton^s is the only one that gives us the icience as it was known to the best ptiilosophers throughout the world, at the beginning of the cur- rent year. It states in comprehensive but concise diction, the facts established by experiment, or other method of demonstration, and details, in an nnderstandable manner, how it is done, but abstains from thediscussion of unsettled or theoretical points. Herein it ie unique; and these characteristics rrn der it a text-book without a rival, for those who desire to stujdy physiological science as it is known to its most successful. cultivators. And it is physi- ology thus presented that lies at the foundation of correct pathological knowledge; and this in turn is the basis of rational therapeutics; so that pathi>lo- gy, in fact, becomes of prime importance in the proper discharge of our every-day practical^nCies. — Cincinnati LoMcet^ May, 1861. Dr. Dalton needs no word of praise from us. He is universally recognizee as among the 'first, if not the very fii st, of American physiologists now living. The first edition of his admirable work appeared but two year^ since, and the advance of science, his own original views and experiments, together with a. desire to supply what he considered some deficien- cies in the first edition, have already made the pre- sent one a necessity, and it will no doubt be even mure eagerly souf;bt for than the first. That it is not merely a reprint, will be seen from the author's statement of the following priucipal additions und alterafioDB which he has made.' The present, likfe the first edition, is printed in the highest style of the printer's art, and the illustrations are truly admira- ble tor their clearness in expressing exactly what their author intended. — Boston Medical and Surgi- cal Journal, March 28, 1861. It is unnecessary to give a detail of the additions; suffice it to say, that they are numerous and import- ant, and such as will render the work still' more valuable and acceptable to the profess ion, as -a learn- ed and cT^ginal treatiseon this all-important; branch of mcJicine. All that was said in comcncndation of the getting up of the first edition, and the superior style of the illustrations apply with equal force to this. No better work on physiology can be placed in the hand of the student. — St. Louis Medical and Surgical Journal, May,, 1861. Theseadditioiis, while testifying to the learning and industry of the author, render the book exceed- ingly useful, as the most complete expose of a sci- ence, of which Dr. Dalton is doubtless the ablest representative on this side of the Atlantic. — New Orleans Med Times, May, 1861. A second edition of this deservedly popular work having been called for in the short space of two years, the author has supplied deficiencies, which existed in the former volume, and has thus more completely fulfilled his design of presenting to the profession a reliable and precise text- book, and one which we consider the best outline on the subiect of which it treats, in any laiisuage. — N. Afnerican Medico- Chirurg. Review, May, lb61. DUNGLISON, FORBES, TWEEDIE, AND CONOLLY. THE CYCLOP JEDIA OF PRACTICAL MEDICINE: comprising Treatises on the Nature and Treatment of Diseases, Materia Medica, and Therapeutics, Diseases of Women and Children, Medical Jurisprudence, &c. &:c. In four large super-royal octavo volumes, of 3254 double-columned pages, strongly and handsomely boiind, with raised bands. ffilS 00. K *^(!* This work contains no less than four hundred and eighteen distinct treatises, contributed by •ixty^^ighl distinguished physicians, rendering it a complete library of relerence for the country practitioner. The most complete work on Practical Medicine •xtant; or, at feast, in our language.— Btt^oio Medical and Surgical Journal. For reference, it ia above all price to every prac- titioner. — Western Lancet. One of the most valuable medical publications of the day as a work of reference it is invaluable. — Western .Tournal of Medicine and Surgery. It has been to ub, both as learner and teacher, a work for ready and frequent reference, one in which modern English medicine is exhibited in the mpst advantageous light. — Medical Examiner. The editors are practitioners of established repu- tation, and the list of contributors embraces many of the most eminent professors and teachers of Lon- don, Edinburgh, Dublin, and Glasgow. It is, in- deed, thegreat raerito) this work that the principal articles have been furnished by practitioners who have not only devoted especial attention to the dis- eases about which they have written, hut have also enjoyed opportunitie* for an extensive ptucti- eal acquaintance with them and whose reputation carries the assurance of their competency justly to appreciate the opinions o' others, while it stamps their own dotftrineswitl high and just authority AfTU rican Medical Journal . DEWEES'S COMPREHENSIVE SYSTEM OF MIDWIFERY. Illustrpted' by occasional cases and many engravings Twelfth edition, with the author's last improvements and corrections In oneoctavovolume, extracloth.offtOOpages. S3 50. PBWEBS'S TREATISE ON THE PHYSICAL AND MEDICAL TRBATWENT OF CHILD REN. The last edition In one volume, octavo extra cloth, 548 pages 363 80 ' OEWEES'S TREATISE ON THE DISEASES OF FEMALES Tenth edition In one volume, octavo extra cloth, 532 pages, witli plates «3 00. 12 HENRy C. LEA'S MEDICAL DUNGLISON (ROBLEY), IV). D., PTofeBSor of Institutes of Medicine in the Jefferson Medical College, Philadelphin ENLARGED AND REVISED EDITION OP 1865— : Just Issued.) MEDICAL LEXICON; a Dictionary of Medical Science, containing a concise Explanation of the various Subjects and Terms of Anatomy, Physiology, Pathology, Hygiene, Therapeutics. Pharmacology, Pharmacy, Surgery, Obstetrics, Medical Jurisprudence, and Den- tistry. Notices of Climate and of Mineral Waters; Formulse for Officinal, Empirical, and Dietetic Preparations; with ihe Accentuation and Etymology of I he Terms, and the French and other Synonymes; so as to constitule aFrenth as well as Engli^h Medifeafc; Lexicon. Thoroughly revised and very greatly m-^dified and augmented. In one very large and handsome royal octavo volume, of 1048 double-columned pages, in small type . strongly done up in extra clolh, Sb 00; leather, raited bands, $6 75 Preface to the New Edition ** The author has again been required to subject his Medical Lexicon to a thorough revision. The progress of Medical Science, and the cnnsequeiit introduction of new subjects and terms, demanded this; and he has embraced the occa^on to render more complete the etymology and accentuation ofth* terms. On no previous revision has so much time and labor been expended by him. Some idea may be formed of this, from the fact, that although the page has been augmented in all iis dimensions, not fewer than between sixty and seventy pages have been added. "As the author has remarked on former occasions^ it has ever been his ardent wish to make the work a caiislactorvand desirable — if not indispenirable — lexicon, in whinh the inquirer may search, without disappointment, for every term that has been legitimated in the nomenclature of the science; and he confidently presents this edition as having more claims on the attention of the practitioner and student than its predecessors. 'Once more the auihor gladly seizes the opportunity afforded him to express his gmteful acknow- ledgments for the vast amount of favor which has been extended to the Dictionary." January, 1S65. The object of the author from the outset has not been to make the work a mere lexicon or dic- tionary of terms, but to afford, under each a condensed view of its various medical relations, and thus to render the work an epitome of the existing condition of medical science. Starting with this view, the immense demand whic.i has Existed for the work has enabled him, in repeated re- visions, to augment its completeness and usefulness, until at length it has attained the position of a recognized and standard authority wherever the language is spouen, * This has only been accom- plished by the earnest determination to bring each successive edition thoroughly on a level with the most advanced condition of contemporary medical science, and on no previous occasion has tnis demanded a more patient and laborious effort than in rendering the present edition fully equal to the wanis of the student of the present day, and in no previous editions has the amount of i^ew matter introduced been so large. While, therefore, the reader who merely desire* a voeabuJary explaining the terms in common use can satisfy himself with the smaller works, such as Hoblyn*s, the student and practi'ioner who wish a work to which they can at all times refer with unfailing confidence for all which it is the province of such a book to supply, must still, as heretofore, keep the latest edition of *' Uunglison's DicTioNARy" within reach. The mechanical execution of.this edition will be found greatly superior to that of previous im- pressions. By enlarging the size of the volume to a royal octavo, and hy the employment of a small but clear ty^ on extra fine paper, the additions have been incorporated without materially increas- ing the bulk of the volume, and the matter of two or three ordinary octavos has been compressed into the space of one not unhandy for consultation and reference. A few notices' of the previous editions are subjoined. mthor, and have furnished ns with a volnme indis- This work. cAe appearance of the fifteenth edition of which it buB become uur duty and pleasure to announce, ie perhaps the most stupendous monument of labor and erudition in medical literature. One would hardly suppose after constant use of the pre- ceding editions, where we have never failed to find a sufficiently full explanation of ever} medical term, that in this edition '^'^ about six thousand subjects and terms have been added^'^with a careful revisioi] and correction of the entire work. It is only neces- Bary 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 appeared. — Buffalo Med. Joum.i Jan. ,1858. The wiirk is a muuument of patient research, skilful judgment, and vast physical labor, that will perpetuate the name of the author more effectuaUy than any possible device of stone or metajl. Dr. Donglison deserves the thanks not only of the Ame- rican profession, butbf the whole medical world. — North Am. Medieo-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 which places it far above comparison, and competition. — Am Joum. Med. SeienceSj Jan. 1BS6. We need only say, that the addition of 6,000 new terms, with their accompanying definitions, may be said to constitute a new work, hy itself. We have examined 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 perfectjion] redound to the lasting credit of its pensable at the present day, to all who would find themselves au niveau with the highest standards of medical information. — Soston Medical cmd Surgical ^(>ttrMa;,Dec.31, 1857. Good lexicons and encyclopedic works generally, ire the most labor-eaving contrivances which lite- rary men enjoy ; and the labor which is required to ;>roduce them in the perfect manner of this example 9 something 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 coitsidered universally as the best work of the kind in any language. — Sillima»^s Journal, March, 1858. A complete and thorough exponent of medical terminology, without rival or possibility of rivalry. — Nashville Joum. of Med. and Surg., Jan. 18S8. It is universally acknowledged, we believe, that this work is incomparably the best and most com- plete Medical Lexicon m the English language, Comment and commendation areunnecessary, as no one at the present da^ thinks of purchasing any other Medical Dietionar} than this.— 5t. Louis Med. and Surg. Journ.j Jan 1858. It is the foundation stone of a good medical libra- ry, and should always be included in the first list of books purchased by the medical student.— ^m. Mtd. Monthly, Jan. 1858. It is scarcely necessary to remark that any medi- cal library wanting a copy of Dunerlison's Lexicon must be imperfect. — Cin. Laneetj /an. 1858. The present edition we may safely say has no equal in ttie world. — Peninsular Med. Jowmal Jan. IbSS. AND SqiENTlFIC PUBLICATIONS. 13 DUNGLISON (ROBLEY), M.D., ProfeBBOT of InatitttteB of Medicine in tlie Jefferson Medical College, Philadelphia. HUMAN PHYSIOLOGY. Eighth edition. Thoroughly revised and exten- sively modified and enlarged, with five hundred and thirty-two illustrations.- In two large and handsomely printed octavo volumes, extra cloth, of about 1500 pages, f 7 00. In revising this work lor its eighth appearance, the author has spared no labor to render it worthy a continuance o£the very great favor which has been extended to it by the profession. The whole contents have been rearranged, and to a great extent remodelled ; the investigations which of late years have been so numerous and so important, have been carefully examined and incorporatedi wid the work in every respect has been brought up to a level with the present state of the subject. u "'V®'" "'^ "'^ author has been to render it a concise but compreihensive treatise', containing the whole body of physiological science, to which the student and man of science can at all times refer with the certainty of finding whatever they are in search of, fully presented in all its aspects ; and on no former edition has the author bestowed more labor to secm:e this result. We believe that it can truly be said, no more com- plete repertory of faotB upon the subject treated, ean anywhere be found. The author has, moreover, that enviable tact at deBcription and that facility and ease of expreBsion which render hint peculiarly acceptable to the caxial, or the studious reader. This facuUy, so reQuisite in setting forth many graver and less attractive subjects, lends additional ehaims to one always fascinating Boston Mei,. Mid Surg. Journal. The most complete and satisfactory system of Physiology in the English language. — Amer. Med. Journal. The best work of the kilid in the English lan- guage. — Silliman^s Journal. The present edition the author has made a perfect mirror of the science as it is at the present hour. As a work upon physiology proper, the science of the functions performed by the body , the student will find it all he wishes. — Nashville Journ. of Med. That he has succeeded, moat admirably succeeded in his purpose, is apparent from the appearance of an eighth edition. It is now the great encyclopsedia on the subject, and worthy of a place iii every phy- sician's library. — Western Lancet. BY THE SAME AUTHOR. aENEKAL THEKAPEUTICS AND MATEKIA MEDICA; adapted for a Medical Text-book. With Indexes of Remedies and of Diseases and their Remedies. Sixth Edition, revised and improved. -With one hundred and ninety-three illustrations. In two large and handsomely printed ottavo vols., extra cloth, of about 1100 J>ages. $6 50. In announcing a new edition of Dr. Dungliaon's general Therapeutics and Materia Medica, weiiave no words of commendation to bestow upon a w^ork whose merits have been heretofore so often and so justly extolled. It must not be supposed, however, that the present is a mere reprint of the previous edition: the character of the author for laborious researciL, judicious analysis, and clearness of ex- pression, is fully sustained by the numerous addi- tions he has made to the work, and the careful re- vision to which he has subjected the whole.— Jf. A. Medieo-Chir. Revievf, Jan. 1858. The work will, we have little doubt, be bought and read by the majority of medical studentsj its size, arrangement, and reliability recommend it to all; no one, we venture to predict, will study it without profit, and there are few to whom it, will not be in sume measure useful as a work of refer- ence. The youn^practitioner, more eBpeciall}^, will find the copious indexes appended to this ediiion of great assistance in the selection and preparation of suitable formulae. — Charleston Med. Journ. and Rs' view. Jan. 1858. BT THE SAME AUTHOR- NEW REMEDIES, WITH FORMULA FOR THEIR PREPARATION AND ADMINISTRATION. Seventh edition, with extensive Additions. In one very large octavo volume, extra cloth, of 770 pages. $4 OO. One of the most useful of the author's works.— Southern Medical and Surgical Journal. This elaborate and useful volume should be found in every medical library, for as a book of re- ference, for physicians, it is unsurpassed by any other work in existence, and the double index for diseases and for remedies, will be found greatly to anhance its value. — New York Med. Gaxetu, The great learning of the author, and his remark- able industry in pushing his researches into every suurce whence information is derivable, have enabled him to throw together an extensive mass of facts and /statements, accompanied by full reference to authorities; which last feature renders the work practically valuable to investigators who desire to examine the original papers.— 'iZ'As Afnerican Journal of Pharmacy. ELLIS (BENJAMIN), M.D. 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 me usual Dietetic Preparations and Antidotes for Poisons. To which is added an Appendix, on the Endermic use of Medicines, and on the use of Ether and Chloroform. The whole accompanied with a few brief Pharmaceutic and Medical Obseryations. Eleventh edition, carefully revised and much extended by Robert P. Thomas, M. D., Professor of Materia Me- dica in the Philadelphia College of Pharmacy. In one volume, Svo., of about350 pages. $3 00. (Jtist Isstted.) On no previous edition of this work has there been so complete and thorough a revision. The extensive changes in the new United States Pharmacopceia have necessitated corresponding alter- ations m the Formulary, to conform to that national standard, while the progress made in the materia medica and the arts of prescribing and dispensing during the last ten years have been care- fully noted and incorporated throughout. It is therefore presented as not only worthy a continuance of the favor so long enjoyed, but as more valuable than ever to the practitioner and pharmaceutist. Those who possess previous editions will find the additional matter of sufficient importance to warrant their adding the present to their libraries. 14 HENRY C. LEA'S MEDICAL, ERICHSEN (JOHN), ProfesBor of Surgery in University College, London, *e. THE SCIENCE AND ART OF SURGERY; being a Trbatisb on Surgioah Injuries, Diseases, and Operations. New and improved American, from the second enlarged and carefully revised London edition. Illustrated with over fojir hundred engravings on wood. In one large and handsome octavo volume, of one thousand closely printed pages, extra cloth, $6 CO. The very distingudshed favor with which this work has been received on both sides of the Atlan- tic has stimulated the author to render, it even more worthy of the position which it has so rapidly attained as a standard authority. Every portion has been carefully revised, numerous additions have been made, and the most watchful care has been exercised to render it a complete exponent of the most advanced condition of surgical science. In this manner the work has been enlarged by about a hundred pages, whilie the series of engravings has been increased by more than a hundred, rendering it one of the most thoroughly illustrated volumes before the profession. The additions of the author having rendered unnecessary most of the notes of the former American editor, but little has been added in this country ; some few notes and occasional illustrations have, however, been introduced to elucidate American modes of practice. It is, in our humble jndgment. decidedly the best book of the kind in the English'language. Strange that just such books are notoftener produced by pub- lic teachers of surg:ery in this country and Great Britain, Indeed, it is a matter of great astonishment but no less true than astonishing, that of the many works on surgery republished in this country within the last fifteen or' twenty years as text-books for medical students, thisis the only one that even ap- proximates to ihe fulfilment of the peculiar wants of youngmen just enteringupon the study ofthisbranch of the profession. — WesumJour.ofMtd. and Surgery. Its value is greatly enhanced by a very copious well-arranged index. We regard this as one of the most valuable contributions to modern surgery. To one entering his novitiate of practice, we regard it the most serviceable guide which he can consult. He will find a fulness of detailleadinff him throLgh everv step of the operation, and not deserting him until the final issue of the case is decided. — Sedwscope. Embracing, as will be perceived, the whole snrgi cal domain, and each division of itself almost com plete and perfect, each chapterfull and explicit, eacl subject faithfully exhibited, we can only express ou estimate of it iJt the aggregate. We consider it ai excellent contribution to surgery, as probably the best single volume now extant' on the subject, and with great pleasure we add it to our text-booEs.— if ashmUe Journal of Medicine and Surgery. Prof Eriohsen's work, for its size, has not been surpassed; his nine hundred and eight pages, pro- fusely illustrated, are rich in physiological, patholo- gical, and operative suggestions, doctrines, details, and processes ; and will prove a reliable resourcs for information, both to physician and surgeon, in the hour of peril. — N. O. Med. and Surg;. Journal. We may say, after a careful perusal of some of the chapters, snd a more hasty examination uf the remainder, that it must raise the character of the author, and reflect gre^t credit upon the college to which he is professor, and we can cordially recom- mend it as a work of reference beta to students and p'acti tinners. — Med. Times and Gazette. We do not hesitatS to say that the volume before us gives a veiy (Admirable practical view of the sci- ence and art ot Surgery Of the present day, and we have no doubt that it will be highly valued as a surgical guide as well by the surgeon. as bv the student of surgery. — Edinburgh Med, and Swg. Journal. FISKE FUND PRIZE ESSAYS. — THE EF- FECTS OF CLIMATE ON TUBERCULOUS DISEASE. By Edwin Lee, M.R.C.S , London, and THE INFLUENCE OF PREGNANCY ON THE DEVELOPMENT OF TUBERCLES By Edwam Wabrbh, M. D., of Edenton,N. C. To- gether in one neat 8vo. volume^ extracloth. SI 00, FRICK ON RENALAFFECTJON?; their Diag- nosis and Pathology. With illustrations. Qua volume, royal 12mo., extra cloth, 75 cents. FERGUSSON (WILLIAM), F. R. S., Professor of Surgery in King's College, London, &c. ' A SYSTEM OF PRACTICAL SURGERY. Fourth American, from the third and enlarged London edition. In one large and beautifully printed octavo volume, of about 700 pages, with 393 handsome illustrations, leather. $4 CO. FOWNES (GEORGE), PH. D., &c. A MANUAL OF ELEMENTARY CHEMISTRY; Theoretical and Practical. With one hundred and ninety-seven illustrations. Edited by Robert Bridges, M. D. In one large royal ISmo. volume, of 600 pages, extra cloth, $2 00. ' The death of the Aithor having placed the editorial care of this work in the practised hands of Drs. Bence Jones and A. W. Hoffman, everything has been done in its revision which experience could suggfest to keep it on a level with the rapid advance of chemical science. The additions requisite to this purpose have necesi-itated an enlargement of the page, notwithstanding which the work has been increased by about fifty pages. At the same time every care has been used to maintain its distinctive character as a condensed manual for the student, diyested of all unnecessary detail or mere theoretical speculation. The additions have, of course, been mainly jn the depart- ment of Organic Cheniistry, which has made such rapid progress within the last few years, but yet equal attention has been bestowed on the other branct(es of the subject— Chemical Physics and Inorganic Chemistry— to present all investigations and discoveries of importance^ and to keep up the reputation of the volume as a complete manual of the whole science, admirably adapted for the learner. Bytheuse of a small but exceedingly cjear type the matter of a large octavo is coihpreSsed within the convenient and poi^table limits of a moderate sized duodecimo, and' at the very low price affixed, it is offered as one of the cheapest volumes before the profession. The work of Dr Dr. Fownes' excellent work'has been universally recognized everywhere in Jiis own and this country, as the best elementary treatise on chemistry in the English topgu^, and is yery generally adopted, we believe, as the standard text- book in all i ur colleges, both literary and scientific .^Charleston Meet. Joum. and Review Fownes has long been before the public, and its merits have been fully appreci- ated as the ,beat text-bqok on chemistry now ia existence. We do. not, of course, place it in a rank superior to the works of Brande, Graham, Turner, Gregory, or Gmelin, but we say that, as a work for students, it is preferable to any of them.— ioi» don Journal of U:ediei»t, AND SCIENTIFIC PUBLICATIONS. 15 FLINT (AUSTIN), M, D., Professor of the Principles and Practice of Medicine in Bellevae Hosp. Med. College, New Yorlt. Now Aeady, 1866. THE PRINCIPLES AND PRACTICE OP MEDICINE. For the use of , Practitioners aad Students. In one large and handsome octavo volqme of over 850 closely printed pages, leather, raised bands, $7 ; handsome extra cloih, $6. The want has for some time been felt in this country of a volume' Jvhich, Within a moderate coinpas's, should give a clear and connected view of general and special pathology and therapeutics m their most modern aspect. Re Bent researches have modified many opinions which were formerly universally received on important points both of theory and practice, and these changes have per- haps as yet scarcely received the attention due to' them in the works accessible to the,profefsioh. 1 he author s reputation as a teacher is a guarantee that the present volumfe will be fully up to the most advanced state of the science of the day, while his long and varied experience as a practi- tioner will insure that m all practical details his work will be a sound and trustworthy guide. BY THE SAME AUTHOB. (Preparing.) PHYSICAL EXPLORATION AND DIAGNOSIS OP t)ISBASES APPEOT- ING THE RESPIRATORY ORGANS. Second edition. In one large and handsome octavo volume, extia cloth. BY THE SAME AUTHOE. A PRACTICAL TREATISE ON THE DIAGNOSIS, PATHOLOGY, AND TREATMENT Of- DISEASES OF THE HEART. In one neat octavo volume, of about 500 pages, extra oloth.. $3 50. We do not know that Dr. Flint has written any- thing which is not first rate; but this, his latest con- tribution to medical literature, in our opinion, sur- passes all the others. The work is most comprehen- sive in its scope, and most sound in the views it enun- ciat£B. The descriptions are clear and methodical ; the statements are substantiated by facts, and are made with s4ch simplicity and slhoisrity, that with- out them they would 6arry conviction. The styl^ is admirably clear, direct, and free from dryness. With Dr. Wai^he's excellent treatise before us, we have no hesitation in saying that Dr. Flint's book is the best work on the heart in the English language'. — Boston Med. and Surg. Journal. GRAHAM (THOMAS), F. R. S. THE ELEMENTS OP INORGANIC CHEMISTRY, including the Applioar tions of the Science in the Arts. New and much enlarged edition, by Henry Watts and Robekt Beidses, M. D. Complete in one large and handsome octavo volume, ol over 800 very large pages, with two hundred and thirty-two wood-cuts, extra cloth. $5 50. »*^ Part II., completing the work from p. 431 to end, with Index, Title Matter, &o., may be had separate, cloth backs and paper sides. Price $3 00. From Prof. E. N. Horsfordi Harvard College. It has, in its earlier and less perfecteditions, been familiar to me, and the excellence of its plan and the clearness and completeness of its discussions, have long been my admiration. No reader of English works on this science can afford to be without this edition of Prof. Graham's Elements.^ — Silliman's JournaJ, March, 1858. From Prof. Wolcott Gibhs, N. Y. Free Academy The work is an admirable one in all respects, and its republication here cannot fail to exert a positive influence,upon theprogrcss of science in this country. GRIFFITH (ROBERT E,), M. D., tc. A UNIViERSAL FORMULARY, containing the methods of Preparing and Ad- ministering Officinal and other Medicines. The whole adapted to Physicians and Pharmaceu- tists. Second Edition, thoroughly revised, with numerous additions, by Robert P. Thomas, M. D., Professor of Materia Medioa in the Philadelphia College of Pharmacy. In one large and handsome octavo volume, extra cloth, of 650 pages, double columns. $4 00. It' was a work requiring much perseverance, and when published was looked upon as by far the best work of its kind that had issued from the Americaii press. Prof Thomas has certainly "improved," as well as added to this Formulary, and has rendered jj additionally deserving of the confidence of pharma- ceutists and physicians. — Jyn. Joumalof Pharmacy . We are happy to announce a new and improved edition of this, one of the most valuable and usefal works that have emanated from an American pen. It would do credit to any country, and will be found of daily usefulness to practitioners of medlcme; it is better adapted to their purposes than the dispensato- ries.— So«[?' "sble text-book for the student and an unfailing counsel for the practitioner in the emergencies ^!U? *° frequently arise in obstetric practice. The illustrations form a novel feature in the work. The lithographic plates are all original, and to insure their absolute accuracy they, have all been copied from photographs taken expressly for the purpose. In ordinary obstetrical plates, the positions of the foetus are represented by dia- grams or sections of the patient, which are of course purely imaginary, and their correctness is scarcely more than a matter of chance with the artist. Their beauty as pictures is thereby increased without corresponding utility to the student, as in practice he must for the most part depend for his diagnosis upon the relative positions of the foetal skull and the pelvic bones of the mother. It is, therefore, desirable that the points upon which he is in future to rfely, should form the basis of his instruction, and consequently in the preparation of these illustrations the skeleton has alone been used, and the aid of photography Invoked, by which a series of representations has been secured of Ijie strictest and most rigid accuracy. It is easy to recognize the value thus added to the very full detai s on the. subject of the Mechanism of Labour with which the work abounds it may be added that no pains or expense have been spared to render the mechanical execution oi the volume worthy in every respect of the character and valuS of the teachings it contains. HABERSHON (S. O.), M. D., AesiBtant Physician to and Lecturer on Materia iVIedica and Therapeutics at Guy's Hospital, &c. PATHOLOGICAL AND PKACTfCAL OBSERVATIONS ON DISEASES OF THE ALIMENTARY CANAL, (ESOPHAGUS, STOMACH, C^CUM, AND INTES- TINES. With illustrations on wood. In one handsoine octa\ro volume of 312 page», extra cloth. $2 50. HOBLYN (RICHARD D,), M. D. A DICTIONARY OF THE TERMS USED. IN MEDICINE AND THE COL LATERAL SCIENCES. A new American edition. Revised, with numerous Addftions, by Isaac Hays, M. D., editor of the " American Journal of the Medical Sciences." In one large royal l2mo. volume, cloth, of over 500 double columned pages. $1 50. To both practitioner and student, we recommend this dictionary as being convenient in size, accurate in definition, and sufficiently full and complete for ordinary conaultfLtion.— -Charleston Med. Joum, We know of no dictionary better arranged and adapted . 1 1 is not encumbered with the obsolete terms of a bygone age, but it contains all that are now in use ; embracing every department of medical sciencfl down to the very latest date. — Western Lancet. Hoblyn'B Dictionary has long been a favorite with us. It la the best book of dennitions we have^ and ought always to be ^upon the atudentU table.— Southern Med. and Surg. Journal. JONES (T. WHARTON), F. R, S., Professor of Ophthalmic Medicine and Surgery in University College, London, &,a. THE PRINCIPLES AND PRACTICE OP OPHTHALMIC MEDICINE AND SURGERY. With one hundred and seventeen illustrations. Third and revised Ameri- can, with additions from the second London edition. In one handsome octavo volume, extra cloth, oi 4S5 pages. $3 '^5. Seven years having elapsed since the appearance of the last edition of this standard work, very considerable additions have been found necessary to adapt it thoroughly to the advance of ophthal- mic sclencb The introduction of the ophthalmoscope has resulted in adding greatly to our know- ledge of the pathology of the diseases of the eye, particularly of its more deeply seated tissues, and corresponding improvements in medical treatment and operative procedures nave been introduced. All these malters the editor has endeavoured to add, bearing in mind the character of the volume as a condensed and practical manual. Toaccommodate this unavoidable increase in the size of the work, its form has been changed from a duodecimo to an octavo, and it is presented as worthy a continu- ance of the favour which has been bestowed on former editions. A complete, series of " test-types" for examining the accommodating power of the eye, will be found an important and useful addition. JONES (C. HANDFIELD), J^. R. S., St EDWARD H. SIEVEKING, M.D., Assistant Physicians and Lecturers in St. Mary's Hospital, London. A MANUAL OF PATHOLOGICAL ANATOMY. First American Edition, Revised. With three hundred and ninety-seven hand.some wood engravings. In one large and beautiful octavo volume of nearly 750 pages, extra cloth. $3 50. As a concise text-book, containinp:, in a condensed form, a complete outline of what is known in the domain of Pathological Anatomy, it is perhaps the best work in the English languaia. Itsgreat merit consists in its completeness and orevity^ and in this respect it supplies a great desideratum in our lite- rature. Heretofore the student of pathology was obliged to glean from a great naraber of monographs, and the field was so extensive that but few cnltivatea it with any degree of success. As a simple work of reference, therefore, it is of great value to the student of pathological anatomy, and should be m every physician's library .—W«jier«La»ci<. 20 HENKy C. LEA'S MEDICAT. KIRKES (WILLIAM SENHOUSE), M. D,, Demonstrator of Morbid Anatomy at St, Bartholomew's Hospital, &.C. A MANUAL OF PHYSIOLOGY. A new Amerixjan, from the third and improved London edition. With two hundred illustrations. In one large and handsome royal 12moj volunrte, extra cloth, pp. 586. $2 25. a. This ia a new and very much improved edition of Dr. Klrkea' well-known Handbook of Physiology. It combines conciseness with completeness, and is, therefore, admirably adapted for consultation by thQ busy practitioner. — Dublin Quarterly Journal . One of the very best handbooks of Physiology wt possess— presenting just such an outline of the sci- ence as the student requires during his 'attendance. upon a course of lectures, or for reference whilst preparingfoT examination.— ^Im. Mtdical Journal ' Its excellence is in its compactness, its clearness, and its carefijlly c-itEsd authorities. It ia the most convenient of text-books. These gentlemen, Messrs. Kirkesand Paget, have the gift of telling us what we want to know, without thinking ' it necessary to tell us all they know*. — Boston Med and Surg. Journal . For the student beginning this study, and the practitioner who has^ but leisure to refresh his memoryj this book is invaluable, as it contains all that it 'IS important to know. — Charleston iHirJ, Journal KN APP'S TECHNOLOGY ; or, Chemistry applied t(i the Arts and to Manufactures. Edited by Dr. Ronalds, Dr. Richardsom, and Prof. W. R. JoHNson. In tw,o handsome 8vo. vols. , extra cloth, with about 500 wood engravings. $6 00. LAYCOCK'S LECTURES ON THE PRINCI- PLES AND METHODS OF MEDICAL OB- SERVATION AND RESEARCH. For the Use of Advanced Students and Junior Practitioners. In one royal 12mo. volume, extra cloth. PriceSl. LALLEMAND AND WILSON, A PKAGTICAL TREATISE ON THE CAUSES, SYMPTOMS, AND TREATMENT OF SPERMATORRHCEA. By M. Lallbmand. Translated and edited by Hknry J McDouGALL. Third American edition. To which is added ON DISEASES OF THE VESICUL^ SEMINALES; and their associated organs. With special refer- ence to the Morbid Secretions of the Prostatic and Urethral Mucous Membrane. By Marris Wilson, M. D. In one neat octavo volume, of about 400 pp., extra cloth. $2 75. LA ROCHE (R.), M. D., &c. YELLOW FEVEE, 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 examination of the connections between it and the fevers known under the same name in other parts of temperate as well as in tropical regions. In two larg:e and handsome octavo volumes of nearly 1500 pages, extra cloth. $7 00. We have not time at present, engaged as we are, by da^y and by night, in the work of combating this very disease, now prevailing in oui city, to do more than give this cursory notice of what we consider as undoubtedly the most able and erudite medical publication our country has yet produced But in view of the startlinf; fact, that this, the most malig- nant and unmanageable disease of modern timesj has for several year^ been prevailing in our country to a greater extent than ever before; that it'''a'>'ework upon the most remark. ^„nS "">«', .>niPortanl; class of diseases to which May! 1861!" '''"'*— -M^"*- ■^»«™- »/ ^- Carolina We have great pleasure in recommending Dr. Lyons' work on Fever to the attention of the pro- fession. It is a work which cannot fail to enhance the author's previous well-earned reputation, as a diligent, careful, and accurate observer. — British, Mea. Journal, March 2, 1861. MONTGOMERY (W. F.), M. D., M. R. I. A., «tc., Professor of Midwifery in the King and Queen's College of Physicians in Ireland, to. AN EXPOSITION OF THE SIGNS A.ND SYMPTOMS OF PREGNANCY. With some other Papers on Subjects connected with Midwifery. From the second and enlarged English edition. With two exquisite colored plates, ai)d numerous wood-cuts. In one very handsome octavo volume, extra cloth, of nearly 600 pages, f 3 75. A book unusually rich in practical SUggestiong.— Am Journal Med. Sciences, Jan. 1857. These several subjects so interesting in them- selves, and so important, every one of them, to the most delicate aild precious of social relations, con- trolling often the honor and domestic peace of a family, the legitimacy of offspring, or the life of its parent, are all treated with an elegance of diction, fulness of illustrations, acuteness and justice of rea- soning, unparalleled in obstetrics, and unsurpassed in medicine. The reader's interest can never flag, so fresh, and vigorous, and classical is our author's style J and one forgets, in the renewed charm of ■-Very page, that it, and every line, and every word has been weighed and reweighed through years of p'reparation ; that this is uf all others the book of Obstetric Law, on each of its several topics ; on all points connected with pregn;incy, to be everywhere received as a manual of special jurisprudence^ at once announcing fact, affordingiaTgument, establish- ing precedent, and governing alike the juryman, ad- vocate, and judge — N. A. Sied.-Chir. Revieip. MEIGSiCHARLES D.), M. D., Lately Professor of Obstetrics, &c. in the Jefferson Medical College, Philadelphia. OBSTETRICS : THE SCIENCE AND THE ART. Fourth edition, revised and improved. With one hundred and twenty-nine illustrations. In one beautifully printed octav* volume, of seven hundred and thirty latge pages, extra cloth, $500. Fkom the Author's Peepace. " (n this edition I have endeavored to amend the wort by changes in its form ; by careful cor- rections of many expressions, and by a few omissions and some additions as to the text. "The Student will find that I have repast the article on Placenta Prsevia, which I was led to do out of my desire to notice certain hew modes of treatment which I regarded as not only ill founded as to the philosophy of our department, but dangerous to ihe people. " In changing the form of my work by dividing it into paragraphs or sections, numbered from 1 to 959, 1 thought to present to the reader a oommon-plnoe book of the whole volume Such a table of contents ought to pnove both convenient and useful to a Student while attending public lectures." A work which hat enjoyed so extensive a reputation and has been received with such general favor, requires only the assurance that the author has labored assiduously to embody in his new edition whatever has been found necessary to render it fully on a level with the moat advanced slate of the subject. Both as a text-book for the student and as a reliable work of reference for the practitioner, it is therefore to be hoped that the volume will be found worthy a continuance 0/ the confidence reposed in previous editions. BY THE SAME ATXTHOR. WOMAN : HER DISEASES AND THEIR REMEDIES. A Series of Lee tares to his Class. Fourth and Improved edition. In one large and beautifully printed octave volume, extra cloth, of over 700 pages. 55 00. which cannot fail to recommend the volume to the attention of the reader. — Ranking's Abstract. In other respects, in our estimation, too much can- not be said in praise of this work. It abounds with beautiful passages, and for conciseness, for origin- ality, and for all that is commendable in a work on the diseases of females, it is not excelled, and pro- bably not equalled in the English language. On the whole, we know of no work on the diseases of wo- men which we can so cordially commend to the student and practitioner as the one before us. — Ohio Med. and Surg. Journal. The body of the book is worthy of attentive con- sideration, and is evidently the production of a clever, thoughtful, and sagacious physician. Dr. Meigs's letters on the diseases of the external or- gans, contain many interesting and rare cases, and miiny instructive observations. We take our leave of Dr. Meigs, with a high opinion of his talents and originality. — The British and Foreign Medico-Chi- rurgical Revievj. Every chapter is replete with practical instruc- tion, and bears the impress of being the composition of an acute and experienced mind. There is a terse- ness and at the same time an accuracy in his de- ■cription ol symptoms, and in the rules for diagnosis, It contains a vast amount of practical knowledge. 3y one who has accurately observed and retained the experience of many years. — Dv^Hn Quarterly Journal. ' Full of important matter, conveyed in a ready and agreeable manner. — St. Louis Med. and Surg. Jour. There is an off-hand fervor, a glow, and a warm- leartednesB infecting the effort of Dr. Meigs, which is entirely captivating, and which absolutely hur- ries the reader through from beginning to end. Be- sides, the book teems with solid instruction, and It shows the very highest evidence of ability, viz., the clearness with which the information is pre- sented. We know of no better test of one's under- standing 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 is such bold relief, as to produce distinct impressions upon the mind and memory of the reader. Tin CharltstoH Med. Journal, 22 HENRY 0. LEA'S MEDICAL MEIGS (CHARLES D.) M. D., Lately Professor of Obstetrics,. &c., in Jeiferson Medical College, Philadelphia. ON THE NATURE, SIGNS, AND TEEATMENT OP CHILDBED FEVER. In a Series of Letters addressed to the Students of his Class. In one handsome octavo volume, extra cloth, of 365 pages. %2 Oi>. lectable book. '# * # This treatise upoa child- The instrVictlve and' interesting author of this work, whose previouslaborshave placed his coun- trymen under deep and abiding oblig-ations, again challenges their admiration in the fresh and vi^or ousj attractive and racy pages before as. It is a de- bed fevers will have an extensive sale, being des- tined, as it deserves, to find a place in the library of every practitioner who scorns tolag in the rear.— Nashville Journ(il of Medieint andSurgery. MACLISE (JOSEPH), SURGEON. SURGrlCAL ' ANATOMY. Forming one volume, very large imperial quarto. With sixty-eight large and splendid Plates, drawn in the best style and beautifully ci)lored. Con- taining one himdred and ninety Figures, many of them the size of life. Together with copious and explanatory letter-press. Strongly and handsomely bound in extra doth, being one of the cheapest and best executed Surgical works as yet issued in this country. $14 00. These plates will be found of the highest practical value, either for consulta- tion in enjergencies or to refresh the recollections of the dissecting room. *^* The size of this work prevents its transmission through the post-office as a whole, but those who desire to have copies forwarded by mail, can receive them in five parts, done up in stout wrappers. Price $12 00. A work which has no parallel in point of accu- racy and^cheapness in the English language.— iV. T. Journal of Medicine. ' * We are extremely gratified to annoance to th« profession the completion of this truly magnificent work, which, as a whole, certainly stands unri- valled, both for accur'acy of drawing, beauty of coloring^ and all the requisite explanations of tha subject in hand. — TA« ifcw Orleans Medical and Surgical Journal . One of the greatjest artistic triumphs of the age in Surgical Anatcnny. — British American Medical Joumm^ No practitioner whose means will admit should fstil t^'puBsess it. — Ranhing^s Abstract, Too much cannot be said in its praise; indeed, we have not language to do it justice. — Ohio Medi- tal and Surgical Journal. The most accurately engraved and beautifully colored plates we have ever seen iji an American book — ^^one of the best and cheapest surgical works •ver published. — Buffalo Medical Journal. It is very rare that so elegantly printed, ao well illustrated, aiid so useful a work, is offered at so moderate a price.— CAaWss ton Medical Journal . Its plates can boast a superiority which places them almost beyond the reach of eompetition, — Medi- tal Examiner, Country practitioners will find these plates of im- mense value.— if. Y. Medical Gazette. This is by far the ablest work on Surgical Ana- tomy that has come under our observation. Wi know of no other work that would justify a stu- dent, in any degree, for neglect of actual disseo-. tion. In those sudflei? emergencies that so often arise, and which require. theinstantaneouscommand of minute anatomical knowledge, a work of this kind keeps the details Of the dissecting-room perpetually fresh in the memory . — The Western Journal of Medi- cine and Surgery. MILLER (HENRY), M, D., Professor of Obstetrics and Diseases of Women and Children in the University of Loutsville. PRINCIPLES AND PEACTICE OF OBSTETRICS, &c. ; including the Treat- mentof Chronic Inflammation of the Cervix and Body of the Uterus considered as a frequent cause of Abortion. With about one hundred illustrations on wood. In one very handsome oc- tavo volume, of over 600 pages, extra cloth. $3 75. tion to which its merits justly entitle it — The- Cin- cinnati Lancet and Observer. ■ A most respectable and valuable addition to our home medical literature, and one reflecting credit alike on the, author and the institution to which he is attached. The student will find in this work a most useful guide to his studies; the country prac- titioner, rusty in his reading, can obtain from itB pages a fair rfesumfe of the moaern literature of the science; and we hope to see this American produc- tion generally consulted by the profession, — V», Med. Journal. We congratulate the author that the task is done. We congratulate him that he has given to the medi- cal public a work which will secure for him a high and permanent position among thp standard autlm- rities on the principles and practice of obstetrics. Congratulations are not less due to the medical pro- fession of this country, on the acquisition of a trea- tise embodying the results of the studies, reflections, and experience of Prof. Miller.'— .?w/faZo Medical Journal. In fact, thisvolumemjist take its place among the standard systematic treatises on obstetrics ; a posi- MACKENZIE (W.), M. D,, Surgeon Oculist In Scotland in ordinary to Her Majesty, &c.&c. A PRACTICAL TREATISE ON DISEASES AND INJURIES OF THE EYE. To which IS prefixed an Ajiatomical Introduction explanatory of a Horizontal Section of the Human EyebaU, by Thomas Wharton Jones, F. R. S. From the Fourth Revised and En- larged London Edition. With Notes and Additions by Addinell Hewson, M. D., Surgeon to Wills Hospital, &c. &c. In one very large and handsome octavo volume, extra cloth, with plates and numerous wood-clits %Q 50. The treatise of Dr. ftlackenzle indisputably holds We consider it the duty of every one who has the the firstplace, and forms, in respect of learning^and love of his profession and the welfare of his patient research, an GncyclopEedia unequalled ;in extent by at heart, to make himself familiar with this the most any other work of the kind, either English or foreign, complete work in the English language upon thedii- —Bixon on Diseases of the Ey*. eases of the eye. — Med. Times and Gazette. AND SCIENTIFIC PUBLICATIONS. 25 PARRISH (EDWARD), . Professor of Materia Medica in the Pbiladelpliia College of Pliarmacy. A TREATISE ON PHAllMACV'. Desigaed as a Text-book for the Student, and as a Guide for the Physician and Pharmaceutist. With many Formulae and Prescriptions. Third ediiion, greatly improved. In one handsome octavo volume, of 850 pages, with several niindred Illustrations, extra cloth. $5 00. {Just Issued.) Though for some time out of print, ihe appearance of a new edition of this work has been de- layed for ihe purpose of embodying io it the results of the new U.S. Pharmacopoeia. The pub- lication of this latter has enabled the Eiuthor to complete hisTevision in the most thorough manner, rhose who have been waiting for the work may therefore rely on obtaining a volume compLCtely On a level with the most advanced condition of pharmaceutical science. The favor with which the work has thus far been received shows that the author was not mis- taken in his estimate of the waot of a treatise which should serve as a practical text-l/ook for all engaged in preparing and dispensiag medicines. Such a guide was indispensable not only to the educuied pharmaceutist, but also to that large class of practitioners throughout the country who are obliged to comptnmd their own prescriptions, and who during their collegiate course have no opportunity of obtaining a practical familiarity with the necessary processes and manipulatidns. The rapid exhaustion of two large editions is evidence that the author has succeeded in thoroughly carrying out his object. Since the appearance of the last edition, much has been done to perfect the science ; the new Pharmacopoeia has introduced many changes to which the profession must conform ; and the authnr has labored assiduously to embody in his work all that physicians and pharmaceutists can ask for in such a volume. The new matter aldne will thus be found worth more than the very moderate cost of the Work to those who have been using the previous editions. All that we can say of it is that to trie practising physician, and especiall) the country physician, who is generally his own apotliecary, there is hard- ly any book that mightnotDettev be dispensed with It is at the same time a dispensatory and a pharma- cy — Louisville Review. A careful examination of this wi»rk enables us to speak of it in the highest terms, as being the best treatise on practical pharmacy with which we are acquainted, and an invaluable vxtZe-mecuTn, not only to the apothecary and to those practitioners who are accustomed to prepare tl eir own medicines, but to every medical man and medical student. — Boston Med. and Surg. Journal. This is altogether one of the most useful hooka we have seen. It is just what we have long felt to be neeijed by apbthe^caries, students, and practition- ers of medicine, most of whom in this country have to put up their own prescriptions. It bears, upon every page, the impress of practical knotvledge, conveyed in a plain common sense manner, and ada^teU to the oomprehension of all who may read it.— Southern Med. and Surg. Journal. That Edward Parriah, in writing a book upon practical Pharmacy some few years ago — one emi- nently original and unique — did the medical and pharmaceutical profesaiunB a great and Valuable ser- vice, no one, we think, wao has had access to its pages will deny; doubly welcome, then, is this new edition, containing the added results of his recent and rich experience as an observer, teacher, and practical operator in the pharmaceutical laboratory. The excellent plan of the first is more thoroughlyj — Peninsular Med. Journal, Jan. 1860. Of course, all a,pothecarieB who have not already a copy of the.^rst edition will procure one of this ; it is, therefore, to physicians residing; in the counti^ and in small towns, who cannot avail themselves of the skill of ah educated pharmaceutist, that we would espfcially commend this work. In it they will find all that' they desire to know, and should know, but very little of which they do really Know in reference to this important collateral branch of their profession; for it is a well established fact, that, m the eaucation of physicians, while the sci- ence of, medicine is generally well taught, very little attention is paid to the art of preparing them for use, and we know hot how this defect can be so well remedied as by procuring and consulting Dr. Parrish'sexcellent work.— Si. Louis Med. Journal. Jan. I860. We know of no work on the subject which would be more indispensable to the physician or student desiring information bn' the subject ofwhich it treats. With Griffith's " Medical Formulary" and this, the practising physician would be supplied with nearly or quite all the mostuseful information on the sub- ject. — Charleston Med. Jour. and Revieto, Jan. 1860, PEASLEE (E. R.), M. D,, Professoi' uf Physiology and General Pathology in the New York lyiedicai CoUjege. HUMAN tilCjTULOGY, ia its relations fco Anatomy^ Phygiology, and Pathology; for the use of Medical Students. With four hundred and thirty-four iUustratioii$. In one hand- some octavo volume, extra cloth, of over 600 pages. $3 75. , We would recommendit ascontainingasummary of all that 18 known of the importantsubjects which It embraces a library upon che copies discussed within itself, and is just what the teacher and learner need. Wp have not only the whole subject of His- tology , interesting in itself,ably and fully discussed, but what 18 ot infinitely greater interest to the stu- dent, because of greater practical value, are its re- lations to Anatomy, Physiology, and Pathology, which are here fully and satisfactorily set forth. — Nashville Journ. of Med. and Surgery. it treats ; of all that is In the great worka of Simon and Lehmann, and the organic chemists in general. Master this one volume, and you know all that is known of the great fundamental principles of medi- cine, and Are have no hesitation in saying that it is an honor so th^ American medical prufession.— St, Louis Mid. and Surg. Journal. ROKITANSKY Curator of the Imperial Pathological Museum, A MAJSUAL OM PATHOLOGICAL Bound m two, extra cloth, of about 1200 pages. KING, G. H. MooBB, and G. E. Day. $7 50. The profesaioli is too well acquainted with tne re- oatation of Rokitansky's work to neoa our asBur- aaee that this is one of the mostprofoand, thorough, and valuable books ever issued from the medical nreas It is sutffeiKi-t.'.andhasnostaBdardof oom- Darison. It is only necessary to announce that it is issued in a form as cheap as is compatible with its (CARL), M.D., and Professor at the University of Vienna, &a ANATOMY. Four volumes, octavj, Translated by W. E. Swaine, Edwakd Sieve- size and ipreservation, and its sale follows as a matter. of course. .Mo library can be called co n ilete without it. — Buffalo Med. Journal An attempt to give our readers any adequate idea jf the vast amount of instruction accumulated in .nese volumes, would be feeble and hopeless Wiite.n Lancet BOYLE'S JlATJfiKlA MEDICA AND THERAPEUTICS; including the P^eoaratioIl.•^ oi the PharmacoixEias of London^ Edinburgh, Dublin, and of the United States With many n=w ine.diciu^b. Kdited by Joseph Carson, M. D. With ninety-eight illustrationi' In one large octavo voluuic, extra cloth, of about 700 pages. S3 00. 24 HENRY C. LEA'S MEDICAL NEILL (JOHN), M. O., Surgeon to thePenuBylvania Hospital, &c,; and FRANCIS QURNEY SMITH, M.D., Professor of Institutes of Medicine in the Pennsylvania Medical College. AN ANALYTICAL COMPENDIUM OF THE VAEIOUS BRANCHES ' OF MEDICAL SCIENCE ; for the Use and Examination of Students. A new edition, revised and improved. In one very large and handsomely printed royal 12ino. volume, of about one thousand pages, with 374 wood-cuts, extra cloth, $4 00. Strongly bouiid in leather, with raised bands. $4 75. This work is agtiin presented as eminently worthy ot 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 changed and improvement in professional science, its reputation is permanently established. The beat work of the kind with which we are acquainted.! — Med. Exwminer. > Having made free use of this volume in our ex- aminations oXf pupils, we can epeak from experi- ence in recunim ending it as an admirable compend for students, and as especially useful to preceptors who examine their pupils. It will save the teacher much -labor by enabling him readily to recall all of the points upon which his pupils should be ex- amined. A work oO^his sort should be in the hands of every one who takes pupils into his office witlj a view of examining them ; and thisisunquestionably thebestofits class. — TTonsylvania Med. Journal In the rapid course of lectures, where work for the students is heavy, and review necessary for an examination, a compend is not only valuable, but it is almost a sine qua non. The one before ub. is, in most of the divisions, the. most unexceptionable of all books of the kind that we know of. The newest and soundest doctrines and che^ latest im-, provements and discoveries are 'explicitly, though concisely, laid before the student There is a claai to whom we very sincerely commend this cheap book as worth its weight in silver — that class is the gradu- ates in medicine of more than ten years' standing. who have not studied medicine since. They wul perhaps Knd outfromitthat the science is not exactly now what it was when they left it off,— TA< S(it4«- scopt PI RRIE (WILLIAM), F. R. S, E., Professor of Surgery Ln the University of Aberdeen. THE PRINCIPLES AND PEACTICE OF SURGERY. Edited by John Neill, M. D., Professor of Surgery in the Penna. Medical College, Surgeon to the Pennsylvania Hospital, (fee. In one very handsome 8vo. volume, extra cloth, oi 780 pages, with 316 illustrations. $3 75. We know of no other surgical work of a reason- able size, wherein there is so much theory and prac- tice, or where subjects are more soundly or clearly taught. — The Stethoscope . Prof. Pirrie, in the work before us, has elabo- rately discussed the principles of surgery, and a safe' and effectual practice predicated upon them. Perhaps ho work upon this subject heretofore issued is so full upon the science of the art of surgery.— Nashville Journal of Medicine and Surgery . PARKER (LANGSTON), Surgeon to the Queen's Hospital, Birmingham, THE MODEEN TKEATMENT OF SYPHILITIC DISEASES, BOTH PBI- MARY AND SEC0NDARY; comprising the Treatment of Constitutional and Confirmed Syphi- lis, by a safe and suocessfui method. With numerous Cases, Formulae, and Clinical Observa- tions. From tiie Third and entirely rewritten London edition. In one neat octavo volume, , extra 6lothj of 316 pages. S2 50. •* ■* PEREIBA (JONATHAN), M.D. MATERIA MEDICA AND THERAPEUTICS; being an Abridgment of the late Dr. Pereira's Elements of Materia Medica, arranged in conformity with the briii-h Pharma- copoeia; and adapted to the use of Medical Practitioners, Chemi-^ts, and Drug'gis's, Medical and Pharmaceutical Students, &c. By F. J. Farre, M. D., Senior Physician to' St. Bartholottiew's Hospital, and London Editor of the British Pharmacopeeia; assisted by Robert Bentley, M.R C. S.,(^r6fe__ssor of Materia Medica and Botany to the Pharmaceutical Society of Great Britain; and by Robert Warikgtoh, F. R. S., Chemical Operator lo the Society of Apothecaries. With numerous additions -and references to the United Slates Pharmacopoeia, by Horatio C. Wood, M. D., Professor of T3olany in the Univer-sity of Pennsylvania. , In one large and hand- some octavo vdume of about yOO pages, with numerous illustrations, (fr^arin^.)- ROBERTS (WILLIAM) M. D., Fhysiciaa to the Manchester ^uyal Infirmary, Lecturer'on Medicine in the Manchester School of . Medicine, Sec. A PRACTICAL TREATISE ON URINARY AND RENAL DISEASES, i, including Urinavy Deposits. Illustrated by numerous ca'esand engravings. In one. handsome octavo volume of over 500 pages, extra cloth, (^st Ready.) AND SCIENTIFIC PUBLICATIONS. a7 STILLE (ALFRED), M.D,, niTTTTr* Professor of the Theory and Practice of Medicine in the University of Pennsylvania. THERAPEUTlOa AMD MATEKlA MJDDlCAj a Systematie Treatise on the Action and Uses of Medicinal Agents, including- their Description and History. Second Edilion, revised and enlarged. In twolarg&and handsome octavo volumes, extra cloth., $10 00. uTust Issued,.) This work is designed especially for the student and practitioner of medicine, and treats the various ' articles of the Materia Medica from the point of view of the bedside, and not of the shop or oi the^ le^iure-room. While thus endeavoring to give all practical information likely to be useful with' respect to theemployment of special remedies in special affections, and the results to be anticipated from their administratibn, a copious Index of Diseases and their Remedies renders the work emi- nently fitted for reference by showing at a glance ihe difTerent means which, have been employed, and enabling the practitioner to extend his resources in difficult cases with all that the experience of the profession has suggested. The speedy demand for another edition of this work sjhows thatit haS acceptably filled an acknow- ledged want Noexertion of the author ha^been wanting to render it worthy a continuance of t-he favor with which it has been received, while an alteration in the typographical arrangement has accommodated the additions without increasing unduly the size of the volumes. Rarely, indeed, have we had Bubmiited to ub a work on medicine so ponderous in its dimensions as that now before ub, and yet so fascinating in its contencs. It is, therefore, with a peculiar gratifi- cation that we recognize in Dr. Siillfe the poases- Bion of many of those more distinguished qualifica- tions which entitle him to appYohation,'and which justify him in coming before his medical brethren as an instructor. A comprehensive knowledge. tested by a sound and penetrating judgment, joined to a love of progresfl— which a uiecriminating spirit of inquiry has tempered bo as to accept nothing new because it Is new, and abandon nuthmg old because it is old, but v^hichestimateB either accurcing Eo its relations to a just logic and experience — manifests icaelf everywhere, and gives to the guidance of the author all 'he assurance of safety which ihe diffi- culties of his subject can allow^. In conciUBion, we earnestly advise our readers to ascertain for them-' selves, t}y a study of Br. Still6*s vulumea, the great value and interest of the stores of knowledge they present. We have pleaenre in referring rather to the ample treasury of undoubted truth8,'the real and assured conquest of medicine, accumulated by Dr. Stille in his pages ; and commend the aum ofhis la- bors to the attention of our readers, as alike honor- able to our science, and creditable to the zeal, the candor, and the judgment of him who has garnered the whole so CAief ally ,->~Edinburgh Med. Journal. The most recent authority is the one last men- tioned, Stille. His great work on <* Materia Medi- ca and Therapeutics," published last year, in two octavo volumes, of some sixteen huii;4red pages, whileitembodies the results of the labor of others up to the time of publication, is enriched with a great amount df original observation and research. we would draw attention, by the way, to the very convenient mode in which the Index is arranged in this work. There fs firstan "Index of Remedies;' next an "Index of Diseases aud their Remedies." Such an arrangement of the Indices, in our opinion, g reatly enhances the practicai value of books of this kind. In tedious, obstinate cases of disease, where we have to try one remedy after another until our stock is pretty nearly exhausted, and we are almost driven to our wit^s end, such an index as the second of the two just mentioned, is precisely what we want. — London Med. Timesand GazettBjAprilj 1861. We think this work will do much to obviate the reluctance to a tihuroughiin vestigation o( this branch of scientific study, for in the wide range of medical literature treasured in the Englisa tongue, we shall hardly And a work written in u style more clear and simple, cunveymg. forcibly the facts taught, and yet free from turbidity and redundancy. There is a fas- cination, in Its pages that will insure to it a wide [lopjilarity and attentive perusal-., and a decree of usafulnesg not often attained througn the influence of a single work. SIMPSON (J. Y.), M. D., Professor of Midwiiery,&c., in the University of Edinburgh, &c. CLINICAL LECTDKES OM THE DISEASES OF WOMEN. With nu- meious illustraliona. In one handsome octavo volume, of over 500 pag-es, extra cloth, $4 00. The principal topics embraced in the Lectures are Vesico-Vaginal Fistula, Cancerof the Uterus, Treatment of Oarcnoma by Caustics, Dysmenorrhosa, Amenorrhoea, Closures, Gontraoiions, &c., of tne Vagina, Vulvitis, Causes of Death after Surgical Operations, SuBgieal Fever, Phlegmasia Dolens, Coccyodinia, Pelvic Cellulitis, Pelvib Hsemaioma, Spurious Preghancy, Ovarian Dropsy, Ovariotomy, Cranioclasm, Diseases of the Fallopian Tubes, Puerperal Mania, Sub-InvolUtion and Super-involution of the Uterus, &c. &o. As a series of monographs on these important topics — many of which receive little attention in the ordinary text-books — elucidated wilh the extensive experience and readiness of resource for which Professor Simpson is so distinguished, there are lijw practilioners who will not find in its pages matter of the utmost importance in the treatment of obscure and difficult cases. SALTER (H. H.), M. D. ASTHMA; its Pathology, Causes, Consequences, and Treatment. 8vo., extra cloth (Just Issued.) $2 50. ThelBoni'm of Dr. Salter's worn which is devoted to ircSHiient, is ot great practical inieresi und value. It wouTd lie necessary to loilow him step by slep in lilB remarks, nut only ou the medicinal, bat also on the dii tctie and hygienic trealmeut uf ttie disease, in order lo convey a justnotiunoi mt practical value of this part of his work. This o4r spa ce forbnig, ^^ SLAOE lO. D.), M. p. DIPHTHERIA : its Nature and Treatment, with an account of the History of its Prevalence in various countries. Second and revised edition. In one neat royal 12mo. volume, extra cloth. $1 25. (.Just Issued.) In one vol. and this we shall little rtgret, if, by our silence, wc Bhou Id ind uce our readers to possess Lhemuelvea of the book itself; a bauk which, without uuubt,de-, serves to be r«nked aming the moat valuable of re- cent,cKs that we have met with ; sup- plying so Inuch both to interest and instruct, that we do not hesitate to {affirm that after having once commenced its perusal, few coulil be prevailed upon to desist before completing it. In the last London edition, all the newly observed and accurately rc- ctiroed facts have been inserted,' including much that is recent of Chemi^cal, Microscopical, and Pa- thological research, besidts papers on numerous subjects never before published Ckarleston Med. Journal and Revieiio . It is at once comprehensive and eminently prac- tical, and by universal consent ttands attheheadof BM THE SAME AUTHOR. ON POISONS:, IN RELATION TO MEDICAL JURISPRUDENCE AND MEDICINE. Second American, from a secoiid and revised London edition. In one large octavo volume, of 765 pages, extra cloth. $5 00. Mr. Taylor's position as the leading fnedical jurist of England, has conferred on him extraordi- nary advaiilages in acquiring experience on these subjects, nearly^all cases of moment being referred to him for examination, as an expert whose testimony i^generally accepted as final. The results of his labors, therefore, as gathered together m this volume, carefully weighed and silled, and presented in the clear and intelligtble style for which he is noted, may be received as an acknowledged authority, and as a guide to be followed with implicit confidence. BY THE SAME AUTSOB. AMD WM. BRANDK. CHEMISTRY. In one volume 8vo. See "Bbande," p. 6. AND SCIENTIFKmrBTilCATIONS. 31 WILSON (ERASMUS), F. R. S. ON DISEASES OF THE SKIN. Fifth American, from the Fifth enlarged London edition. In one handsome octavo volume, of nearly 700 large pages, with illustrations on wood, extia cloth $4 50. ■ '^'^^Vp^^s.sical work, which for twenty years has occupied the position of the leading authority in the English language on its important subject, has just received a thorough revision at the hands of the author, and is now presented as embodying the results of the latest investigations and expe- rience on all matters connected with diseases of the skin. The increase in the size of the work shows the industry of the author, and his determination that it shall maintain the position which it has acquired as thoroughly on a level with the most advanced condition of medical science. A few notices of the last edition are appended. No matter what other treatises may he in thd libra- ry of the medical attendaat, he needs the clear and suggestive counsels of Wilson, who is thoroughly- posted up on all subjects connected with cutaneous pathology. We ^jave, it is very true, other valuable works on the maladies that invade the skin ; but, compared with the volume under consideration, they .are certainly to be regarded as inferior lights in guid- ing the judgment of the medical man. — Boston Med. and Swg. Journal, Oct. 1857. The author adopts a simple and entertaining style. He strives to clear away the complications of ,hla subject, and has thus prodWed a book Ulled with a vast amount of information, in a form so agreeable as to make It pleasant i-eading, even to the uninitiated. More especially doe.s it deserve our praise because of its beautiful and complete atlas, which the American publishers have successfully imitated from the origi- nal plates. We pronounce them by ikr the best imi- tations of nature yet published in our country. With the text-book and atlas at hand, the diagnosis is ren- dered easy and accurate, and the practitioner feels himself safe in his treatment. W.e will add that this work, although it must have been very expensive to the pulTlishers, is not high priced. There is no rea- son! then, to prevent every physicianfrom obtaining a work of such importance, and one which will save him both labor and perplexity. — t^a. Med. Journal, As a practical gviide to the classification, diagnosis, and treatment of the diseases of the skin, the book is complete. We know not-hing, considered in this as- ''peet, better in our language; it is a safe authority oa all the ordinary matters which, in this range of dis- eases, engage the practitioner's attention, and pos- sesses the high quality — unknown, we believe, to every older manual, of being on a level with science's high-watermark; a sound book of practice. — London Med. Times. ThewritingB of Wilson, upondiaeasesof the skin, are by far the most scientific and practical that have ever been presented to the meaical world on this subject. The pirelenteditionisagreat improve- ment on all its predecesBors. To dwell upon all the great merit's and high claims of the work before us, seriatim^ would indeed be. an agreeable service j it would be a mental homage which we could freely offer, but we should thus occupy an undue amount of space in this Journal. We will, however look a± Bome of the more salien.t points with which it abounds, and which make itincompuva oiy superior to all other treatises on the subject of dermatology N o mere speculative views aire allowed a place in, this volume, which , withoutadoubt;Will,foravery long period, be acknowledged as the chief standard work on dermatology. The principles of an enlighiened and rational therapeia are introduced on ^very ap- propriate occasion. — Am. Jour. Med Science. When the first edition of this work appeared about fourteen years ago, Mr Erasmus Wilson hau iBilready given some years to the study of piseases of theSkiUj and he then expressed his intention of devoti&g his filcure life to the elucidation of this branch of TVIedical Science In the present edition Mr. Wilson presents us with the results of his ma- tured experirnce, and, we have now before us not merely a reprint of his former publications, but an en,tirely new and rewritten voiume. Thus, the whole history of the diseases affecting the skin, whether they originate in that structure ur are the mere mani- festations of derangement of internal oirgans,' is brought under notice, and the book includes a mass of ^formation 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 inexistencein the En- glish language. — Lmidon Med. Times and G-azette. ALSO, NOW RE4;DY, A SEEIES OF PLATES ILLUSTRATINa WILSON ON DISEASES OP THE SKIN; consistingof twenty beautifully execuled plates, of which thirteen are exquisitely colored, presenting the Normal Anatomy and Pathology of the Skhi, and' containing accurate re- presentations of about one hundred varieties of disease, most of them the size of nature. Pricb id cloth. $5 50. ' ' ,. ' . In beauty of drawing and accuracy and finish of coloring these plates will be found equal to Miything o/ the kind as yet issued in this country. The value of the new edition is enhanced by an Eidditional colored plate. The plates by which this edition is accompanied leave nothing to be desired, 60 far as excellence of delineation and perfect accuracy of illustration are concerned.— Medico-Ctirurgicaf Review. Oftheseplatesitisimposaible to sneak loo highly The representations of the various forms of cutane- ous disease are suigularly|)iccurate, and the color- ing exceeds almost anything we have met with.— British and Foreign Medical Review. We hsve already expressed oar high appreciation of Mr. Wilson's treatise on Diseases of the iSkin. The plates arfe comprised in a separate volume, .which we counsel all those who possess the text to purchase. It is. a beautiful specimen of cqlor print- ing, and the representations of the various forms of skin disease are as faithful as is possible in plates of the size.— Soston Med. and Surg. JoumaK Anril 8, 1858. IP" Also, the TEXT and PLATilS done up in one handsome volume, exira cloth, price $9 50. BY THE SAME AUTHOR. THE DISSECTOR'S MANUAL; or, Practical and Surgical Anatomy. Third American, from the last revised, and enlarged English edition. Modided and rearranged, by WlIAIAM Hunt, M. U., Demonstrator of Anatomy in the University vi Pennsylvania. In one large and handsome royal 12mo. volume, extra cloth, of 582 pages, with 154illu8tration8. S2 00. BY THE SA^E AUTHOR. HEALTHY SKIN; A Popular Treatise on the Skin and Hair, their Preserva- tion and Management. Second American, from the fourth London edition. One neat volume royal 12mo.,e;ctra cloth, ol about 300 pages, with numerous illustrations. $1 00. ' 38 HE^UY C. LEA'S MEDICAL New and mucli enlarged edition. WATSON (THOMAS), M. D., tc, Late Physician to tlie Middlesex Hospital, &c. LECTURES ON THE PRINCIPLES AND PRACTICE QP PHYSIC. delivered at King's Colleg-e, London. A new Americanj fromtbe last revised and enlarged Eng'li&h edition, with Additions, by D. Francis Condib, M. D., author of " A Practical Treatise on the Diseases of Children," &c. With one hundred and eighty.five illustrations on wood. In one very large and handsome volume, imperial octavo, of over 3200 closely printed pages in small type; extra cloth, $6 50; strongly bound in leather, with raised bands, ^7 ao. That the high reputation of this work might be fully maintained, the author has subjected it to a thorough revision ; every portion has been examined with the aid pf the most recent researcl]ies m 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 cfimate which are little known in England, as well is 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 v&luabteL assistance may be conveyed to Ihe student in elucidating the text. The work will, therefore, be found thoroughly on a level with ■ the most advanced state of medical science on l;)oth sides of the Atlantic. The addition's which the work has received ar^ shown by the fact that notwithstanding an en- largement in the size of the page, more than two hundred additional pages have been necessary to accommodate the two large volumes of the London edition (which sells at ten dollars), within the compass of a single volume, and m its present form it contains the matter of at least three ordinary octavos. Believing it to be a work which should lie on the table of every physician, 'and be in the hands of 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 ol its rnechanical execution renders it an exceedingly attractive volume. The fourth edition now appears, so carefully re- vised, as to add considerably to the value of a book alteady acknowledged, wherever the ISrigliah lan- guage is readj to be beyond all comparison the beet systematic work on the Principles and Practice of Physic in the whole range of medical literature. Every lecture contains proof of the extreme anxiety of tjie author to keep pace with ihe advancing know- ledge of the day One scarcely knows whether to admire most the pure, simple, forcible English — the Vast amount of useful praetrcal information condensed into the Lectures^— or the manly, kind- hearted,' unassuming character of the lecturer shin- ing through his work.— LoMd. Med. Times. Thus these admirable volumes come before the profession in their fourth edition, abounding in those, distinguished attributes of moderation, judgment, erudite cultivation, clearness, and eloQuence, with which they were from the i&tst in^'es'.eil, but yet richer than'before in the results of m(.rft prolonged observatiuD, and in the able appreciation of the latest advances in pathology and medicine by one of the most profound medical thinkers of the day .— London Lancet. The lecturer's skill, his wisdom, his learning, are equalled by the ease of his graceful diction; his elo- quence, and the far higher qualities of candor^' of courtesy J of modesty, and of gener6u6 appreciation of merit in others.— iv A. Med -Chir Review. • Watson's unrivalled, perhaps unapproachable work on Practice — the copious addiUons made to which (the fourth edition) have given it 4II the no- velty and much of the interest of a new book.— Charleston Med. Journal. Lecturers, practitioners, and students. of medicine will equally hail the, reappearance of tUe work of Dr. Watson in theformo^anew — af"ourth— edition. We merely do justice to our own feelings, and, we are sure, of the whole profession, if we thank him for having, in the trouble and turmoil of a large practice, made leisure to supply the hiatus caused by the exhaustion of the third ediliun. For Dr. Watson has not merely caused the lectures to be reprinted, but scattered through the whole work we find additions or alterations which pruve that the author has in every way sought to bring up his teach- ing to the level of !^he most recent acquisitiunsin science.-rfrit, and For. Medico-Chir. Review. New and much enlarged edition. WILSON (ERASMUS), F. R.S. A SYSTEM OF HUMAN ANATOMY, General and Special. A new and re- vised American.from tke last and enlarged Englii^h Edition. Edited by 'W.H.Gobiiecht,M.D.,, Professor of Anatomy in the Pennsylvania JVIedical College, fee. Illustrated with three hundred and ninety-seven engravings on wood. In one large and handsome octavo volume, ol over 600 large page^; extra cloth, $4 00. The publishers trust that the well earned reputation so long enjoyed by this work will be more than maintained by the present edition. Besidefca very thorough revision by the author, it has-been mo:^t careJ'uUy exumined by the editor, and the efibrtss ot both have been direcied to introducing everything which increased experit nee in its U!*e nas suggested as desirables to render it a complete text-book lot jhose [peeking to obtain or to renew an acquaintance with Human Anatomy. The amount of additions which it has.thus received may be estimated from the fact that the present efdiiion contains over one-fourlb more matter than the la^l, rendering a smaller type and an enlarged flage requisite to keep the volume within a convenient size. The editor has exercised the utmost caution to obtain entire accuracy in the text, and, ha? largely increased the number of illustra- tions, of which there are about one hundif'ed and ^fty more in this edition than in the last, thus bringing distinctly before the eye of the student everything of interest orlrapottance. It may be recommended to the student as no less distingui^ed by its accuracy and clearnesB of de- scription tlian by its typographical elegance. The wood-cuts are exquisite. — Brit, and For. Medical Review . An elegant edition of one' of the most useful and accurate systems uf unatomicini science which has been issued from the press The illuBtrutions are really beautiful. In its style the work is extremely Concise and intelligible. No one can poasibjy take up this Vdlume without being struck with the great bMuty of its mechanical execution, and the clear- ness of the{ descriptiims which it contains lb equally evident. Let students, by all mean« exuinuie tae claims of this work on thfcir notice, before cliey pur- chasti a text-bnok of tihe vitally important science which this volume so fully and easily unfolds.— Lancet. We regard it as the best system now extant for students. — Westerii Lancet. U therefore receives ourhighestcommendation.— Southern Med. and Surg. Journal. 33 HENRY C. LEA'S MEDICAL P UBLICATIONST WINSLOW (FORBES), M. D., O. C. L., &c. ON OBSCURE DISEASES OP THE BRAIN AND DISORDERS OF THE MIND; their incipienl SymptomB, Pathology, Diagnosis, Treatment, and Prophylaxis. Second American, from the third and revised English edition. In one handsome octavo volume, of nearly 600 pages, extra cloth. $4 25. {Just Ready.) Pathology. It completely exhausts the sukject, in We close this brief and necessarily Very imperfect notice of Dr. Winslow's great and classical work, by expressing our conviction that it is long since so important and beautifully written a volume has is- BuedTrom the British medical press. — DvhlinMed. Press, July ?S, 1660. We honestly believe this to be the best book of th6 season hanlcing^s Aistract, July, 1860. The latter portion of Dr. Winslow's work is ex- .clusively devoted to the consideration of Cerebral the same manner as'the previous seventeen chapters relating to morbjd psychical phenomena left nothing unnoticed m reference to the mental symptoms pre- monitory of cerebral disease. It is impossible, to overrate the benefits likely to result from a general perusal of Dr. Winslow's valuaole and deeply in- teresting work. — London Lancet, June 23, 1860. It contains an immense .mass of information.— Brit, ami For. Med.-Chir. Beviev, Oct. 1860. WEST (CHARLES), M. D., Accoucheur to and Lecturer on Midwifery at St. Bartholomew's Hospital, Fhysician to the Hospital for \ Sick Children, &c. LECTURES ON THE DISEASES OF WOMEN. SecoDd American, from the second London edition. In one handsome octavo volume, extra clotli, of about 500 pages ; price $3 25. * ' *^* Gentlemen who received the first portion, as issued in the *' Medjcal News and Library," can now c6laiplete their copies by procuring Part II, being page 309 to end, with Index, Title matter, &c., 8vo.; cloth, price $1 25. We mustnow conclude this hastily written sketch with the confident assurance to our readers that the work will weli repay perusal. The conscientious, painstaking, practical physician isapparent on every page. — N. Y. Journal of Medicine. We know of no treatise of the kind so complete and yet so comp&ct.~r-Chicago Med. Jour. A fairer, more honest, mure earnest, and more re- liable investigator of the 0iany diseases of women and children is not to be found in any country.— Southern Med. and Surg. Journal. We have to say of it, briefly and decidedly, that it is the best work on„the subject in any language ,' and that it stamps Dr. West as the facile princeps of British obstetric authors. — Edinb. Med. Journ. . We gladly recommend his Lectures as in thehigh- est degree instructive to all who are interestea in obstetric practice. — London Lancet. IJappy in his simplicity of manner, and moderate in his exjpression of opinion^ the author is a sound reaaoner and a good practitioner, and his book is worthy of the handsome garb in. which it has ap- peared. — "Virginia Med. Journal. We must lake leave of Dr. West's very useful work, with «iur commendation of the clearness of ita style, and theincustry and sobriety of judgment of which it gives evidence.— ^Lontion Med Times. Sound judgment and good sense pervade every chapter of-the. book. From its perusal we have de- rived unmixed satisfaction. — Dublin Quart. Jown, BY THE SAME AUTHOIL. {Just Reodjf.i LECTUEES ON THE DISEASES OF . INFANCY AND CHILDHOOD. Fourth American, from the fifth enlarged and improved London edition. In one handsome octavo volume, extra clotli, of about six hundred arid fifty pages, f 4 50. The numerous editions through, which this work has passed on both sides of the Atlantic, are the best evidence that it has met a want felt by the profession: Few practitioners, indeed, have had the opportunilieb of obt-ervation and experience enjoyed, ty the author. Jn his Preface he remarks: 'The preserjt edition embodies the results of 1200 recorded cases, and of nearly 400 post-mortem examinations,* collected from between 30,000 and 40,000 children, who, during the past twenty -SIX years have come under my care, either in public or in private practice." The universal favor with which the work has been received shows that the author has made good use of these unusual advantages. The three former editions of the work now before as have placed the, author, in tue foremost rank of those physicians who have cievoted special attention to the diseases of early life We attempt no. ana- lysis of this edition, but mayrefer the reader to some of the chapiters to which the largest additions have been made — those on Diphtheria, Disorders of the Mind, and Idiocy, for instance — as a prooi that the work is re.ally a new editiuu; not a mere reprint. in its pretent shape it will be lound of the greatest possible service in the every-day practice of nine- tenths of the profession. — Med. Tim,ei and Gazette^ London, Dtc. 10, 1859. ■■ . All things considt red this book of Dr. West is by far theoe&t treatise in our language upon such njodificatiuDB of morbid action and dise&se as are wiiintssedwhen w'e have to deal with infancy and Childhood. It is true that it confines itself to such disorders as come within the province of the phy- tictan, and even with respect to these it is unequal as regards minutentss of consideration, and some diseases it omits to notice altogether. But those who know anything of the present condition of paediatrics will readily admit that it would be next to impossible to efiect more, or effect it better, than the accoucheur of St. Bartholomew's has done in a smgle volume. The lecture (XVI.) upon Disordtra of the Mind in chile rcn is an admirable specimen of the value oi the later information conveyed in the Lectures of Dr. Gharlea West London Limeetf Oct. 22, 1859. Since the appearance of the first edition, about eleven years, ago,, the experiencjB of the author, has doubled ; so that, whereas the lectures at first were founded on six hundred observations, and one hun- dred and eighty dissections made among nearly four- teen thousand children, they now embody the results of nine hundred observations, and two hundred and eighty-ei^htpost-mortem examinations made among nearly thirty thtmsand children, who, during the past twecty years, have been under his care.— , British Med. Journal^ Oct. 1, 1859. BY THE SAME AUTHOR. AN ENQUIRY INTO THE PATHOLOGICAL IMPORTANCE OF ULCEB- ATIOW Ot THE OS UTERI. In one neat octavo volume* extra cloth. ^1^.