COLUMBIA LIBRARIES OFFSITE HEALTH SCIENCES STANDARD HX64085651 QM451 .W581899 Anatomy of the brain 4* *"S 5 iWftARlCS ^ «* BY J. RYLAND WHITAKER, B.A, M.B.(Lond.) FELLOW OF THE EOTAL COLLEGE OF PHTSICIAXS, EDIXBUBGH; LECTCKEE OX iXilOUT, SCEGEOXS' HALL, EDIXBUEGH, AXD LADIES' SCHOOL OF 1IEDICIXE, 1IIXTO HOrSE, EDIXBUBGH; EXA3IIXEE IX AXATOMT, EOTAL COLLEGE OF PHTSICIAXS, EDIXBCBGH. SIXTH THOUSAND . EDINBURGH: E. & S. LIVINGSTONE. 1899. ("all eights eeserved] ISV TO Sir JAMES ALEXANDER RUSSELL, LL.D., M.A., M.B., B.Sc, F.R.C.F.E., F. R.S.Ed., Late Lord Provost of Edinburgh, ®Ijis littk fttatmal is respsrtfulhr gauicat^u IN TOKEN OF MANY KINDNESSES. THE AUTHOR. PKEFACE TO THIED EDITION. This little book, first written whilst a student at the earnest request of students, has now reached a third edition, and has become a favourite, perhaps undeservedly, with those for whom it was originally intended. Although fully conscious of its many defects, I fear that my time, now so much occupied with teach- ing, leaves me no chance of remodelling the book in a way that more mature years and experience would suggest. It must practically remain in the same garb, however imperfect, in which it first appeared. This the student, at least, will probably not regret. In this edition my work has been confined to the removal of the more glaring blemishes and errors, and to the rewriting and bringing up to date some of the sections. As the book is now much used by senior students and by practitioners, a general summary, for their special use, has been added at the end. Though the actual number of pages has been slightly increased, the matter has been shortened, for the individual pages have been made smaller, more margin viii PREFACE. having been allowed, so that the book reads more easily and more room has been obtained for extra plates. To Dr W. F. Harvey, and to my other demon- strators and students who have so kindly, more than once, read over the proofs, I cannot sufficiently express my gratitude. Finally, it remains for me to thank Messrs M 'Lagan & Cumming for the trouble they have taken in the reproduction of the plates, and the Messrs Livingstone and staff for their uniform courtesy and excellent execu- tion of the work. Surgeons' Hall, Edinburgh, 1st May 1899. TABLE OF CONTENTS. Dedication, Preface, ... List of Plates, l'AGE V vii xiii SECTION I. The Spinal Cord, its Membranes, and its Vessels, Chapter I. Membranes of the Spinal Cord, I. Dttra Mater, II. Pia Hater, III. Arachnoid, Cerebro-Spinal Fluid. 3 4 6 9 10 Spinal Vessels, Chapter II. 11 Chapter III. The Spinal Cord, 12 I. Fissures of the Cord, 14 II. Spinal Nerves, 16 III. White and Grey Matter of Cord, 17 1. White Matter, IS White Columns, IS White Commissure. 25 Structure of White Matter. 26 2. Grey Matter, 27 Cornua, 28 Grey Commissure, i".' Neuroglia, 30 Nervous Constituents, 31 Vesicular Columns, 32 IV. Deep Origins of Spinal Nerves, ... 37 TABLE OF CONTENTS. SECTION II. The Brain, its Membranes, and its Vessels, Dissection to Expose Membranes, Dissection to Remote Brain, ... PAGE 43 43 44 Chapter I. Membranes of Brain, 45 I. Dura Mater, 45 Falx Cerebri, 46 Falx Cerebelli, ... ... 47 Tentorium Cerebelli, ... 47 II. Pia Mater, 49 III. Arachnoid, ... 50 IV. Venous Sinuses, Chapter II. 52 Vessels of the Brain, ... 57 I. Arteries, ... 57 II. Veins, 61 Chapter III. Sub-divisions of the Brain, I. Medulla Oblongata, 1. Fissures of Medulla, 2. White Matter — Areas of Medulla, 3. Grey Matter of Medulla, Recapitulation,... II. Pons Varolii, 1. White Matter of the Pons, _'. Grey Matter of the Pons, III. Cerebellum, 1. Lobes of Cerebellum, ... 2. Peduncles of Cerebellum, 3. Medullary Vela, I. Grey Matter of Cerebellum, Minute Structure of Grey Matter, 5. White Matter of Cerebellum, The 4th Ventricle, 63 65 66 67 77 85 87 89 91 92 95 97 100 101 102 105 106 TABLE OF CONTENTS. XI PAGK IV. Cerebkum, ... 113 I. Exterior of Cerebrum, 114 1. Fissures of Cerebral Hemispheres, 114 2. Lobes and Convolutions 116 Structure of Cerebral Cortex, ... 128 II. Base of Cerebrum, 132 III. Interior of Cerebrum, 138 1. Ventricles, ... 141 Lateral Ventricle, 142 The 3rd Ventricle, .. 146 The 5th Ventricle and Septum Lucidum, 147 Velum Interpositum, 148 Choroid Plexus, 149 Veins of Galen, 150 Great Transverse Fissure, 150 2. Basal Ganglia, 151 Corpora Striata, 151 ' Optic Thalami, 153 Corpora Geniculate, ... 155 Pineal Gland, 156 3. White Strands, 157 Corpus Callosum, 157 Anterior Middle, and Posterior Com- missures, 160 Fornix, 160 Internal and External Capsules, 163 IV. Crura Cerebri, &c, 165 1. Crura, 165 2, Corpora Quadrigemina, 169 3. Aqueduct of Sylvius, ... 170 Origins of Cranial Nerves, 170 General Summary, ... 180 SECTION III. Outlines of Development or — I. The Spinal Cord, II. The Brain, Index, 190 191 193 199 LIST OF PLATES. PLATE PAGE I. — Fig. 1, General View of the Brain and Spinal Cord (Hirschfeld and Leveille). Figs. 2 and 3, Mem- branes of the Spinal Cord, ... ... ... 4 IJ. — Figs. 4, 5, and 6, Membranes of the Spinal Cord. Fig. 7, Central Canal of Spinal Cord (Foster). Fig. 8, Sections of Spinal Cord, ... ... 8 III. — Sections of Spinal Cord at various levels, ... 14 IV.— Fig. 13, Tracts of Spinal Cord. Fig. 14, Vesicular Columns and Deep Origins of Spiual Nerves. Fig. 15, Blood- Vessels, ... ... ... 22 V. — Various kinds of Nerve Cells of Spinal Cord, ... 28 VI.— Figs. 23, 24, 25, and 20, White Tracts of Spinal Cord (Bristowe). Fig. 27, Pia Mater. Fig. 28, White Nerve Matter of the Spinal Cord. Fig. 29, Anterior Horn of the Spinal Cord, ... ... 32 VII.— Fig. 30, Different kinds of Cells of the Spinal Cord, Neuroglia, &c. (Van Gehuchten). Figs. 31 and 32, Collaterals (Edinger), ... ... ... ... 34 VIII.— Fig. 1, Collaterals. Fig. 2, Spiual Ganglion. Fig. 3, Neuroglia Cells (Van Gehuchten). Fig. 4, Nerve Cells,... ... 40 IX. — Fig. 33, Diagrammatic View of Parts of the Brain. Fig. 34, Membranes of the Brain, ... ... 41 X. — Fig. 35, Membranes aud Vessels of the Brain (H. & L.). Fig. 36, Venous Sinuses (Wilson's Plates), ... ... ... ... ... 48 XL — Figs. 37 and 38, Venous Sinuses (Wilson), ... 52 XII.— Figs. 39 and 40, Blood- Vessels of the Brain (II. & L.), 58 xiv LIST OF PLATES. PLATE PAGE XIII.— Blood- Vessels of the Brain, ... ... ... 60 XIV. — Blood- Vessels of the Brain, and Areas of Motion and of Sensation, ... ... ... ... ... 62 XV.— Views of the Medulla, ... ... ... ... 66 XVI. — Fig. 50, Diagram of Course of Fibres in the Spinal Cord,... ... ... ... ... ... 70 XVII.— Sections of the Medulla, ... ... ... ... 78 XVIII. — Figs. 57 and 58, Nuclei in the 4th Ventricle (Edinger), ... ... ... ... ... 82 XIX.— Figs. 59 and 60, Pons. Fig. 61, Crura Cerebri, ... 90 XX— Figs. 62 and 63 (H. & L.), Figures of Cerebellum, 94 XXI.— Figs. 64 and 65, Cerebellum. Fig. 66 (Edinger), and Fig. 67, Structure of Cerebellum (Ferrier), 98 XXII. — Structure of Cerebellum (Edinger) ... ... 102 XXIIa.— Figs. 68 and 69, Structure of Cerebellum (Van Gehuchten), ... ... ... ... ... 104 XXIII. — Convolutions of the Brain, ... ... ... 118 XXIV. — Convolutions of the Brain, ... ... ... 124 XXV.— Figs. 74 and 75, Relation of Fissures, &c, to Skull and Scalp, ... ... ... ... ... 128 XXVI. — Figs. 76, 77, and 78, Structure of Cerebrum (Ramon y Cajal), — ••• ... ... ... 130 XXVIL— Fig. 79, Base of the Brain, ... ... ... 134 XXVIII.— Fig. 80, Side View of the Brain, ... ... 135 XXIX. — Diagram of Ventricles, ... ... ... ... 133 XXX. — Diagram of Ventricles, ... ... ... 142 XXXI. — Diagram of Ventricles, ... ... ... ... 144 XXXII. — Velum Interpositum, &c. ... ... ... 145 XXXIII. — Diagram of Ventricles, ... ... ... 143 XXXIIIa.— Fig. 1, Velum Interpositum, Fig. 2, Fornix, &c, ... 150 XXXIV.— Capsules, k ... 154 LIST OF PLATES. xv PLATE PAGE XXX V— Fig. 87, Nuclei of 3rd and 4th Nerves (Edinger). Fig. 88, Optic Nerves (Hill). Fig. 89, Fornix. Fig. 90, Nucleus Oaudatus (Edinger). Fig. 91, Fornix (own specimen, and Hill), ... ... 160 XXXVI.— Fig. 92, Blood- Vessels to Capsules. Fig. 93, Ver- tical Section of the Cerebrum. Fig. 94, Diagram of the 3rd Ventricle. Fig. 95, Fascia Dentata, &c, 164 XXXVII.— Fig. 96, Diagram of Tracts of Fibres (Hill). Fig. 97, Tracts of Medulla (Edinger, Hill, Bruce). Fig. 98, Inter-peduncular Space (Holden). Fig. 99, Eesti- form Body (Edinger), ... ... ... ... 168 XXXVIII. — Nuclei of Origin of Cranial Nerves, ... ... 172 XXXIX. — Nuclei of Origin of Cranial Nerves, ... ... 176 XL.— Figs. 100 and 101, Tracts of Degeneration in the Brain and Spinal Cord, ... ... ... 182 XL I. — Diagram of Course of Motor Fibres (Van Gehuchten), 184 XLII. — Diagram of Course of Sensory Fibres (Van Gehuchten), 186 XLIII. — Diagrams of Development of the Brain and Spinal Cord (after Quain, Kolliker, Sclrsvalbe, and Edinger), .. ... ... ... ... 192 XL IV. — Diagrams of the Vesicles of the Brain and the Parts Developed therefrom (after Van Gehuchten), ... 194 ANATOMY OF THE BRAIN AND SPINAL CORD. THE nervous system of man comprises two por- tions, the Sympathetic and the Cerebrospinal, The latter, the cerebro-spinal, has two parts: a central, which consists of the brain and spinal cord, and a peripheral, which consists of the several nerves of the body. NERVOUS SYSTEM. 1. Sympathetic. 2. Cerebro-spinal. I I I (i) Central Part- (2) Peripheral Part- Brain and Spinal Cord. The Nerves of the Body. The Central Nervous System of man, the only part treated of in this hook, is, as we have just seen, made up of the Brain and Spinal Cord. Sur- rounded by the bony wall of the cranium in the one case, and of the spinal canal in the other, these two parts of the great cerebro-spinal nervous system are continuous with each other through the foramen magnum. They are each enveloped in three distinct membranes, the Meninges, which form additional pro- tective sheaths around them, and help to support them in their respective cavities. B 2 THE SPINAL CORD The Brain and Spinal Cord are, moreover, each composed of two kinds of nervous substance, known from their colour as the White and Grey Matter respectively ; but with this difference in their arrangement, namely, that in the Brain the grey matter is situated chiefly on the outside, forming the cortex ; whilst in the Spinal Cord the white substance is external, and the orev matter forms the central core or pith. We shall describe, firstly, the Spinal Cord, with its Vessels and Membranes; and, secondly, the Brain, with its Vessels and Membranes. AND ITS MEMBRANES. SECTION I. SPINAL CORD AND MEMBRANES. Dissection. — To see the spinal cord and its membranes it will be necessary to open the vertebral canal. To do this, remove the muscles from the vertebral grooves, and saw through the laminae of the vertebras on each side, close to their union with the pedicles, being careful, especially in the dorsal region, to direct the edge of the saw inwards. Carry the incision downwards as far as the lower end of the sacral canal, and upwards as high in the neck as may be convenient. Break through with the chisel any partially sawn arches, cut through the various ligaments, and remove the pieces of bones thus detached. Carefully clear away the connective tissue, veins, and fat covering the outer aspect'of the dura mater, snip through the articular processes with the bone forceps, and dissect out one or more of the processes of the dura mater which pass through the inter-vertebral foramina. Examine the outer surface of the dural sheath, and then slit it open in its entire length, CHAPTER I. MEMBRANES OF SPINAL CORD. (Plates I. and II.) These membranes are identical in many respects with those of the Brain, and are therefore simi- larly named. They are the Dura Mater, the Pia Mater, and the Arachnoid. The Dura Mater is the most external, the Pia Mater is in close contact with the cord, and the Arachnoid forms a vertical, tubular partition between the others, dividing the 4 THE SPINAL CORD space between them into two — viz., the sub-dural and the sub-arachnoid. Thus we speak of the sub- dural space, meaning that between the dura mater and the arachnoid, and of the sub-arachnoid, or that between the arachnoid and the pia mater. For convenience we shall describe (1) the Dura Mater; (2) the Pia Mater ; (3) the Arachnoid. I.— THE DURA MATER. (Pigs. 2 to 6.) The Dura Mater, the most external and the strongest covering of the cord, is a firm fibrous mem- brane, continuous at the foramen magnum with the similar membrane which lines the cranial cavity. Enclosed in the spinal canal, it does not, however, form an endosteum to the bones, and in this respect differs from the cranial dura mater. Its outer surface has a shining, pearly-white appearance, and is separated from the walls of the spinal canal by loose areolar tissue with a little fat, and by a plexus, of veins. Slender fibrous bands, especially at its lower end, attach it closely to the posterior common ligament of the vertebras. In extent the dural sheath reaches from the foramen magnum to the second or third piece of the sacrum, where it ends in a conical cul-de-sac. As you will see upon open- ing it, the cavity which it encloses is much longer and wider than its contents — the spinal cord — for this latter ends at the first or second lumbar vertebra. Below the pointed termination of the spinal marrow — conns meclullaris — the cavity of Fir,. I Fig. 2. ki/perior I gin us. iteral * inus. Subdural space Plate I. A interior median f. j 'erebrum Subarachnoid space ^Cerebellum Spinal nerv Medulla . Dura mater. irachno'ld. Pia mater. Spinal Cord m terminale. tda equina. M Conns medullaris. Spinal __ j nerve roots. \ Ligament um dentieulatum.. i Epithelium. Posterior median f. Fig. 3. ira mater. achnoid. Via mater. Linea plendens AND ITS MEMBRANES. 5 the dural sheath is occupied by bundles of nerve roots — cauda equina (Fig. 1) — in the midst of which you will be able to pick out a slender, silvery- looking thread, the filura terminate or central ligament (Fig. 1, fil. ter.). Continuous with the apex of the conus medullaris, this terminal filament runs down the middle line amonsrst the nerve roots to the lower end of the dural cavity. It there pierces the dural sheath, receives an investment from it, and passes along with this investment to be attached to the back of the coccyx (filum dura? matris spinalis, see page 13). Thus, we see that the dura mater forms a very loose "covering to the cord, and we find, moreover, that it has a greater capacity in the neck and back than it has in the loins. Smooth and glistening on its inner aspect, it presents on each side a longitudinal series of rounded openings arranged an pairs, one for each spinal nerve root. These roots, as they pass out to the inter-vertebral foramina, carry with them a tubular prolongation of the dura mater which, in part, ensheaths the spinal ganglia, and, in part, blends with the periosteum of the neighbouring bones (Figs. 4 and 6, page 8). It is important to recollect that, although the spinal cord itself ends at the spot indicated — viz., the second lumbar vertebra, the dura mater, the arachnoid, and the cerebro-spinal fluid, extend as far as the second piece of the sacrum, so that injuries inflicted upon the spine as low down as this latter point may cause death by inducing inflammation of the meninges. 6 THE SPINAL CORD Minute structure of the dura mater. — Under the microscope the dura mater will be seen to consist of white fibrous and elastic tissues, arranged in longitudinal bands or lamellae, with flattened, branched, connective tissue corpuscles, clasping the bundles of fibrils. Both its inner and outer surfaces are smooth and covered by epithelial plates. Many lymphatics and blood-vessels, as well as slender nerve filaments derived from both spinal and sym- pathetic systems, are furnished to its substance. II.— THE PIA MATER. (Figs. 2 to 6.) Disskotion, — After slitting open the dura mater, the first mem- brane you will see is the delicate arachnoid. Pierce it with a sharp-pointed blow-pipe and inflate, as far as you can, the sub- arachnoid space. Next, to expose the pia mater, remove a small piece of the arachnoid from any part of the cord, leaving it intact elsewhere. The Pia Mater is a delicate, highly - vascular, fibrous membrane, which so closely surrounds the spinal cord that it cannot easily be stripped off. It also furnishes sheaths to the roots of the spinal nerves. A distinct process or fold of it passes into the anterior median fissure of the cord, and other smaller septa penetrate the spinal marrow at various points, carrying with them blood-vessels and lymphatics for the nutrition of both its white and grey matter. The largest of these septa is seen at the posterior median fissure (Fig. 6, Plate II., page 8). It is not, however, a fold of pia mater like that in the anterior fissure, but consists of the AND ITS MEMBRANES. 7 supporting tissue of the cord called neuroglia (see page 30). If you compare the pia mater of the cord with that of the brain, you will find that the former is thicker, less vascular and more adherent to the subjacent nervous tissue. The outer surface of the pia mater is compara- tively rough, and has the three following structures connected with it — the linea splendens, the liga- mentum denticulatum, and the septum posticum. 1. The linea splendens (Fig. 3, page 4) is a thickened fibrous band along its anterior aspect. It is sometimes difficult to make out. 2. The ligamentum denticulatum is a well- defined structure (Figs. 3 and 5, lig. dent.) which runs longitudinally on each side of the cord in the form of a toothed white band, having its serrated edge turned outwards. It helps to support the spinal marrow within its dural sheath. Internally, it is attached to the pia mater, about midway between the lines of origin of the anterior and posterior nerve roots, reaching upwards as high as the medulla oblongata, and ending below on the pointed extremity of the cord, the conus medullaris. Externally, its outer margin forms a series of tooth- like processes, about twenty-one in number, which are fixed to the inner surface of the dura mater, in the intervals between the points of exit of successive nerve roots. The highest of these denticulations is attached opposite the margin of the foramen magnum, between the last cranial and first spinal nerve, and the lowest between the twelfth dorsal and first lumbar nerves. The ligamentum denticulatum 8 THE SPINAL CORD thus partially divides the sub-arachnoid space into an anterior and a posterior compartment. 3. At the back of the cord is another process or partition — the sejrtum posticum (Figs. 4, 6, page 8, sep. post.) — which crosses the sub-arachnoid space and serves to connect the pia mater with the arachnoid. Below the end of the cord, the pia mater, though at first retaining its tubular form, afterwards becomes suddenly reduced in size, and is finally prolonged as a sheath to the delicate thread-like continuation of the spinal cord, the filum terminate or central ligament (Fig. 1) ; the silvery hue of which will enable you to distinguish it among the surrounding bundles of nerve roots (cauda equina). Lying between the pia mater on the one hand, and the arachnoid on the other, but connected with both, is a quantity of delicate connective tissue arranged in the form of a spongy network, the sub - arachnoid trabecular, which are clothed with epithelial plates (Fig. 4, page 8, sub-arach. trab.). The lacuna? or areola?, thus formed, contain the greater part of the cerebro-spinal fluid. Minute structure of the pia mater (Fig. 27, Plate VI., page 32). — The pia mater consists of a basis of white fibrous connective tissue, which is arranged in interlacing bundles, and supports a plexus of blood-vessels. Both its surfaces are covered by epithelial cells. It possesses a com- plete network of lymphatics ; and its nervous supply is derived from the sympathetic system. Fig. 4. "Epithelium iganientu racknoid train ■ Dura mater. Pia mater. nal nerve. nation. Fig. 5. Plate II. I) lira gamentwtn dent nerve. Fig. 6. Arachnoid. interior median fissure. Dura mater. Pia mater. Ligament . • dent. Fig. 7. ibarachnoid space. Subdural space. Transverse section of lower part of spinal canal. Fig. 8. Interior median fiss. Anterior nerve root. Septum posticum. Posterior median fissure. rang/ion. ; Spina! nerve. Posterior median fins. Posterior nerve root. AND ITS MEMBRANES. 9 III.— THE ARACHNOID. (Figs. 2 to 6.) The Arachnoid is an extremely fine and delicate membrane. It is non-vascular, and thus differs materially from the other two. Many authors deny that it is a special membrane, and consider it to be one of the layers of the pia mater, describing the two together as the pia-arachnoid (see pages 50 and 51). Forming a cylindrical partition between the dura mater and the pia mater, the arachnoid divides the space between them into two — the sub-dural and sub-arachnoid, previously referred to. The sub-dural space is very narrow, for the outer surface of the arachnoid is in more or less close contact with the dura mater. The sub-arachnoid space is much larger and contains the chief part of the cerebro-spinal fluid. It is crossed by the sub-arachnoid trabecular which connect the arachnoid with the pia mater. The arach- noid, moreover, forms tubular prolongations around both the nluni terminale and the teeth-like processes of the ligamentum denticulatum ; and similar cover- ings are furnished to the roots of the spinal nerves. These sheaths enclose the nerve roots as they pass outwards to the dura mater, but, when the roots pierce that membrane, the epithelium of the outer surface of the arachnoid becomes continuous with the epithelium lining the inner surface of the dura mater, whilst the rest of the arachnoidal sheath blends with the perineurium of the nerves (Figs. 2 and 4. epith.). Thus we see that each spinal nerve io THE SPINAL CORD root receives a covering from all tbree membranes of the cord. It is stated, too, that the sub-dural and sub-arachnoid spaces, though they do not directly communicate with one another, are both continuous with the lymphatic plexuses which surround the spinal nerves. It may perhaps be well to mention that the sub-dural space was formerly spoken of as "the cavity of the arachnoid" — the arachnoid membrane itself being then looked upon as a serous membrane, enclosing a serous cavity, and the epithelial lining of tbe deep surface of the dura mater as one of its layers. Minute structure of the arachnoid. — The arachnoid consists of bundles of white fibrous tissue, interlacing with one another, and arranged for the most part longitudinally. Both its surfaces are covered by epithelial cells. The source from which it derives its nerve-supply is still very doubtful; most probably it is the sympatbetic system. As we have said, many authors regard the arachnoid as one of the layers of the pia mater. Cerebro-spinal Fluid. — The cerebro-spinal fluid, about two fluid ounces in quantity, is a clear-looking, alkaline liquid, containing little or no albumen. It chiefly occupies the interstices of the sub-arachnoid trabecular By its means, probably, an equality of pressure is maintained upon the brain and cord ; hence we find that any sudden disturbance of tbe fluid, such as would be caused by pressure on a spina bifida, at once gives rise to serious cerebral symptoms — such as convulsions and loss of consciousness. AND ITS MEMBRANES. n CHAPTER II. SPINAL VESSELS. (Fig. 15, Plate IV., page 22.) Dissection. — Remove the cord with its sheaths from the spinal canal, by cutting through the spinal nerves at their exit through the inter- vertebral foramina, and snipping the bands of connective tissue which attach it to the posterior common ligaments. 1. Arteries. — The arteries on the surface of the spinal cord are the anterior and posterior spinal. The anterior spinal artery, formed above by the union of two branches, arising one from each vertebral artery, runs along the front of the cord underneath the linea splendens. As it passes down- wards it is reinforced by a series of anastomotic branches from arteries in the neck and back. It gives off branches to the pia mater, to the roots of the spinal nerves, to the anterior median fissure (a cleft seen on the front of the cord), and ends below upon the filum terminale. The posterior SPINAL arteries, two in number, run downwards, one on each side, behind the line of origin of the posterior nerve roots. They are derived from the same source as the anterior, and are joined by small branches which enter the inter- vertebral foramina along the roots of the spinal nerves. They anastomose freely and send offsets into the septum at the back of the cord — the posterior median septum. 12 THE SPINAL CORD 2. Yeins. — The veins of the spinal cord lie within the pia mater, have an anterior, a posterior, and right and left lateral trunks, and form a fine plexus over the surface of the corcl. They are larger on the back than on the front of the cord. Laterally, after being joined by veins from the bodies of the verte- brae, branches pass out with the spinal nerves through the inter-vertebral foramina, and open into either the vertebral, inter-costal, or lumbar veins, according to the regions in which they occur. 3. Lymphatics. — There are no lymphatic vessels, properly so-called, in the nervous system, but there are lymph spaces round the nerve cells — peri-cellular, and round the blood-vessels — peri-vascular. CHAPTER III. THE SPINAL CORD. The Spinal Cord is the elongated cylindrical column of nervous substance contained in the vertebral canal. Composed of two kinds of nervous matter — an outer cortical part, consisting princi- pally of white nerve fibres, and an inner grey core or pith, consisting chiefly of nerve cells and their processes — it is invested by three membranes, the meninges ; gives origin to thirty-one pairs of spinal nerves ; and is partially divided by anterior and posterior median fissures into two lateral seg- ments, which are united across the middle line by a band of fibres called a commissure. You have AND ITS MEMBRANES. 13 already seen that the spinal marrow is much smaller than the capacity of its bony case, so much so, that, in the adult, it occupies two-thirds only of the length of the spinal canal. About eighteen inches loner, it reaches from the foramen magnum to the lower border of the first or second lumbar vertebra. Above, it is continuous with the medulla oblongata ; below, it ends in a pointed extremity, the conus medullaris, from the apex of which it is prolonged downwards as a delicate thread-like fila- ment, the filum terminale. According to the regions of the spine in which they are situated, different portions of the cord have received special names ; thus we speak of the cervical, dorsal, and lumbar portions. The spinal cord, moreover, presents two swellings or enlargements — an upper one, the cervical enlargement, extending from about the third cervical to the second dorsal vertebra, and a lower or lumbar enlargement, which, beginning at the tenth dorsal vertebra, is largest opposite the twelfth, and then gradually tapers away to the pointed extremity of the cord. The connection between the increase of nervous substance in these two parts of the cord and the origin of the large nerve bundles given off to the upper and lower limbs respectively is sufficiently obvious. Filum terminale. — This filament is the atrophied remnant of the embryonic spinal cord. In its upper part it is composed of nervous matter surrounding a central canal, but in its lower part it becomes solid and consists of cells (the exact nature of which 14 THE SPINAL CORD is not decided), of blood-vessels, and ultimately of mere prolongations of pia mater and of dura mater (see page 5). We shall now examine (1) the exterior of the cord with its spinal nerves; then (2) the interior. I.— FISSURES OF THE CORD. On both the anterior and posterior aspects of the spinal cord there is seen a median longitudinal cleft or depression, which penetrates some distance into the nervous substance, and partially divides it into two lateral halves. These clefts are called the anterior and the posterior median fissures (Fig. 10, Plate III., page 14). The transverse band which connects the two halves at the bottom of these fissures js called the commissure of the spinal cord. 1. The anterior median fissure is the wider of the two, though in depth it extends through only one-third of the thickness of the cord. It contains a distinct fold of the pia mater, which conveys blood- vessels into the interior of the spinal marrow (Fig. 4, Plate II., page 8; Fig. 15, Plate IV., page 22). At the bottom of this fissure lies the transverse band of nerve fibres, the anterior or white part of the com- missural band (Fig. 14, Plate IV., page 22), which unites together the two lateral halves of the cord. 2. The posterior MEDIAN FISSURE, narrower but deeper than the anterior, is rather a septum than an actual fissure, for it does not contain a fold of the pia mater, but is filled up by blood-vessels Fig. 9 — Cervical Region— Anterior commissure Posterior commissure. Plate III. Anterior cornu. Lateral cornu. Posterior cornu. Clarke's column. (.-_ A nterior mes. fiss. I- Lateral cornu. V f-Posteriov mes. fiss. Anterior group of cells. Anterior cornu. Lateral group of cells. .1^ Posterior group of cells. \ Lateral cornu. 'Posterior ccmu. Fig. 12. —Sacral Region— 'Anterior cornu. Posterior cornu. Sections of the Spinal Cord in the regions of the different groups of nerves AND ITS MEMBRANES. 15 and connective tissue (ependyma and neuroglia cells) attached closely to the deeper layer of the pia mater. The 'posterior or grey commissure lies at the bottom of this septum (Fig. 9, Plate III., page 14). About the middle of this grey band you will find the central canal of the spinal cord. In addition to these median fissures, we have yet to describe, on each side of the cord, two lateral longitudinal depressions, the positions of which are indicated by the lines of origin of the anterior and posterior nerve roots (Figs. 5 and 8, Plate II., page 8). They are called the anterolateral and postero- lateral grooves, though the antero-lateral is scarcely a distinct groove. They mark off the surface of each half of the cord into three columns — an anterior, a lateral, and a posterior. The anterior column lies between the anterior median fissure and the anterior nerve roots ; the lateral column between the an- terior and posterior nerve roots ; and the posterior column between the posterior nerve roots and the posterior median fissure (Fig. 15, Plate IV., page 22). In the cervical region, close to the posterior median fissure, two additional slightly marked clefts can be made out, one on each side, which, with other minor clefts, break up the cord into many tracts or columns, to be described hereafter. Some authors divide the cord thus — (1) A posterior segment — between the posterior median fissure and the posterior nerve roots. (2) An antero-lateral seg- ment — between the anterior median fissure and the posterior nerve roots. The antero-lateral segment is again divided, by the line along which the anterior 1 6 THE SPINAL CORD nerve roots take origin, into (a) an anterior part, between the anterior median fissure and the anterior nerve roots ; and (6) a lateral part, between the anterior and posterior nerve roots. From a develop- mental point of view this sub-division is the more correct. II.— SPINAL NERVES. (Figs. 5 and 8, Plate II., page 8 ; Fig. 15, Plate IV., page 22.) Along the sides of the spinal cord arise, as we have already seen, the several spinal nerves — thirty-one pairs — eight cervical, twelve dorsal, five lumbar, five sacral, and one coccygeal. Each nerve takes origin, called its superficial origin, by two roots — an anterior and a posterior — springing from the antero-lateral and postero-lateral grooves respectively. The roots are enclosed in sheaths similar to those of the cord itself, and they pierce the dura mater by two separate openings, one for each root (Figs. 3 and 6). Sometimes each root, especially the posterior one, consists of two bundles, perforating the dura mater by two separate openings. Except in the case of the first nerve, the posterior roots are the larger, and their fasciculi more numerous and thicker than those of the anterior roots. Thev, moreover, become connected with a ganglion, or collection of nerve cells (spinal ganglion, Fig. 8, page 8 ; Fig. 14, page 22), before they join the anterior roots to form a spinal nerve. The anterior roots are motor, the posterior sensory in function, whereas the spinal nerves themselves are AND ITS MEMBRANES. 17 mixed nerves, carrying both motor and sensory impulses. These nerve roots, with the exception of the first or highest, are not attached to the cord opposite the vertebra below which they leave the vertebral canal, but at a higher level. This differ- ence between the points of origin and exit (a matter of considerable clinical importance),* though slight in the cervical region, increases as we descend the cord, until, at its lower end, the nerve roots form an almost vertical bundle, known as the cauda equina (Fig. 1). In the upper part of the cervical region of the cord, a little in front of the posterior roots of the first four nerves, we see a series of nerve filaments which are the twigs of origin of the spinal part of the spinal accessory nerve. They soon unite to form a single trunk, which ascends to the foramen magnum. III.— WHITE AND GREY MATTER OF THE SPINAL CORD. A transverse section of the spinal cord, such as that represented in the Figures Plate III., page 14, will demonstrate to you the interior of the cord and its sub-division into two lateral halves. It will show you distinctly (1) the existence of the white cortex; (2) the central grey core; and (3) the commissural band, composed of white matter and grey matter, connecting together its lateral segments. * See Hilton's " Rest and Pain/' 5th Ed., 1892, page 86. C iS THE SPINAL CORD 1. THE WHITE MATTER. Taking up such a section of the spinal cord, and examining it with the naked eye, you will see, as we have just said, that the white matter forms the outer or cortical part of the cord, and surrounds the grey centre. This white matter, arranged in a series of columns, gradually increases in amount from below upwards, being specially augmented in the cervical and lumbar enlargements. Compared with the grey matter, it is more abundant in the neck and back, but less so in the loins (Plate III., page 14). 1. White Columns of the Cord, and their Sub-divisions.— Your attention, you will remember, has already been called to the fact that the exit and entrance of the nerve roots sub-divide each lateral half of the cord into three longitudinal white seg- ments or columns — namely, an anterior, a LATERAL, and a posterior; each of which, however, can be again sub-divided into smaller tracts or strands of fibres w T hich have received special names and have special functions (Fig. 13, Plate IV., page 22). Tracts or Strands. — To determine by actual dissection these various tracts or strands would be impossible ; the task has, however, been accomplished by the study of development and by the aid of ex- perimental physiology and pathology; for "we may learn as much of the course of nerve fibres by studying them in their birth as in their death — in their development as in their decay." The development teaches us that different tracts or AND ITS MEMBRANES. 19 bundles of fibres acquire their white substance — medullary sheath — at different periods of their growth (see page 41), so that, in specially prepared specimens, we are enabled to pick out and trace these tracts or systems through successive sections of the cord. On the other hand, the study of phy- siology and the selective action exercised by disease (so analogous to that exercised by certain poisons) affords equally valuable information. For to phy- siology and pathology are we indebted for the knowledge that, when a nerve fibre degenerates in consequence of injury or disease, the proper nerve substance is replaced by connective tissue, which, when treated with staining reagents, behaves differ- ently from the surrounding undegenerated nerves. By the above means, then, we can define with more or less certainty the following tracts or systems in the several white columns of the spinal cord (see Table, page 25). (a) Anterior Column. — In this column have been mapped out a median and a lateral division. 1. The median division — the dieect PYRAMIDAL tract — antero- internal tract— fasciculus of Turck — is a well-marked bundle of fibres, situated close to the anterior median fissure (Fig. 13, page 22). It is a descending or motor tract, and is, as we shall see hereafter, a continuation of that part of the anterior pyramid of the medulla oblongata which does not decussate in the medulla, hence its name, direct pyramidal tract. It decreases in size from above downwards, and can be traced as a distinct 20 THE SPINAL CORD fasciculus to the first lumbar nerve, though isolated fibres have been followed as far as the fourth pair of sacral nerves. Its fibres, as they descend in the cord, cross over at lower levels to the opposite side, and thence through the grey matter of the cord become connected with the anterior nerve roots of that side. 2. The lateral part of the anterior column, antero EXTERNAL TRACT, forming by far the larger part of the column, has also been called the anterior root zone, basis bundle, or anterior ground tract. It is mostly made up of short commissural fibres between different segments of the cord. (6) Lateral Column. — This well-defined column marked off on the surface of the cord by the antero- lateral and postero-lateral grooves, and limited inter- nally by the grey matter, is composed of five distinct strands of fibres (Fig. 13, page 22). 1. Thus occupying the posterior part of the column, at a little distance from the surface of the cord, is a bundle of fibres, known as the crossed pyramidal tract (crossed pyram., Fig. 13). This, as we shall afterwards see, is the continuation downwards of the chief part of the anterior pyramid of the medulla oblongata of the opposite side ; hence the name, crossed pyramidal tract. This crossed pyramidal tract exists in all regions of the cord. It is the principal motor tract, and decreases in sectional area as we descend the cord ; for its fibres constantly turn into the grey matter (Fig. 23, &c, Plate VI., page 32, and Plate XLL, page 184). AND ITS MEMBRANES. 21 2. The DIRECT CEREBELLAR TRACT. — The thin lamina of white matter, which separates the crossed pyramidal tract from the surface of the cord, con- stitutes the direct lateral cerebellar tract, so called from its connection with the cerebellum on the same side (Fig. 13, page 22; and Plate VI., Fig. 23, &c, page 32, lat. cereb.). It is an ascending tract, and extends from the level of the second lumbar nerve upwards to the restiform body. Its fibres are prob- ably connected through a group of cells called Clarke's column, with the posterior nerve roots on the same side. It most likely carries visceral sensa- tions and sensations from tendons, and is concerned with the maintenance of equilibrium. 3. Ascending anterolateral tract (Hadden and Gowers). — This area is situated in the anterior part of the lateral column, in front of the direct cerebellar tract (Fig. 13, page 22). It lies on the surface of the cord and extends into the anterior column nearly as far as the anterior median fissure. It is called by Gowers the antero-lateral ascending tract — comma-shaped tract — not, however, to be confounded with the ill-defined descendinsr comma tracts described in the posterior columns. Gowers' tracts exist most likely in all regions of the cord, but their lowest limits have not yet been defined. Their fibres can be traced to the cerebellum. They are sensory routes from the spinal cord to the brain. 4. Descending antero-lateral tract. — Closely associated with Gowers' tract, and occupying about three-fourths of the antero-lateral column (see Fig. 13) 22 THE SPINAL CORD is the descending antero- lateral tract (Lo wen thai and Marchi). Its fibres mingle with those of Gowers' tract and come from the cerebellum of the same side. Some of them probably pass out by the anterior nerve roots. It may be an indirect motor tract, or may belong to the crossed pyramidal tract. 5. The rest of the lateral column is called the MIXED ZONE or LATERAL GEOUND BUNDLE (Fig. 13), though it is often divided into two parts — anterior and posterior mixed strands — the latter being also called the limiting layer (Fig. 13). It consists of fibres with a short course — commissural fibres. Thus we see that the lateral column contains in all five tracts — the crossed pyramidal tract; the three cerebellar tracts (two ascending and one descending); and the mixed zone. (c) Posterior Column. — The arrangement of the strands in this column somewhat resembles that in the anterior (Fig. 13). Thus, especially in the cervical region, we find, close to the posterior median fissure, (1) a narrow band called the postero - internal tract — posterior median column, fasciculus of Goll ; and (2) an external division named the postero - external tract — cuneate fasciculus, posterior root zone, fasciculus of Burdach. The postero-internal tract, GolVs tract, in- creases in size from below upwards. It is best seen in the dorsal region, for at lower levels it is not distinct from the next fasciculus, the fibres of the two being intermingled. It is a direct tract from Plate IV. Anterior median/. Fig. 13. Direct pyramidal tract. Basis bundle, es. antero-lateral tract Limiting layer. - BB Comma Tract Ascending antero-lateral tract. Crossed pyramidal tract. Lateral cerebellar tract. Lissauer's tract. Postero-external .. , . . ' / Postero-internal tract. Posterior median j. '"\\/ >K^ ( ' : Anterior root. Anterior vesicular column Lateral fillet ... Lateral 1 vesicular column. Clarke's column. Posterior vesicular column \_ Anterior cornu. Anterior commissure. Latend cornu. '^Posterior commissure. U0- Posterior cornu. Ganglion .interior column — -Lateral column -Posterior column Diagram of tht vessels of the cord. J. T MURRAY DELT AND ITS MEMBRANES. 23 the muscles to the brain, its fibres being derived from the posterior roots of the spinal nerves. The POSTERO - EXTERNAL STRAND, fasciculus of Burdach, also increases in size as it ascends. Its fibres, like those of the last bundle, are derived from the posterior nerve roots. We shall presently trace them to their ultimate destination (see " Posterior Nerve Roots"). The distinction between these two parts of the posterior column (Goll and Burdach) is of considerable importance in the pathology of locomotor ataxy, a disease of the posterior nerve roots, in which there is loss of muscular sense — sense of effort — and con- sequently of co-ordination of muscular movements, for in these tracts travel fibres which are closely connected with that sense and with the mechanism of co-ordination. That the columns of Goll and Burdach conduct sensory impulses from the muscles is also proved by the fact that in whales, where the extremities are not developed, these columns — Goll and Burdach — are rudimentary in comparison to what they are in animals with well-developed limbs. Another tract, the postero - marginal, is a small zone which lies in this column round the tip of the posterior horn, separating the horn from the surface. It is often called Lissauer's tract (Fig. 13), and its fibres, which come from the posterior roots, run for a short distance upwards and downwards in the tract, and then enter the grey matter. Lissauer's tract is sometimes described with the lateral column. (See below, " Posterior Roots.") i4 THE SPIN A l. CORD Several other minor strands have been described in the different columns of the cord, but are riot, of sufficient Importance to detain as here, Some of them — Unices tract, the lateral fillet, and the comma tract — are figured in Figs. 13 and 14, Plate IV., page 22. It is well, however, to remember that the several tracts :ire not, so definite as the figure; would indicate for with the ascending fibres we find descending intermingled, and viae vered, To Sinn up, then, the pyramidal t racts, direct and crossed, and part of the anterolateral tract, are descending or motor tracts; the posterior column, the lateral cerebellar tract (spino-cerebellar), and Gowers' tract are, on the other hand, ascending or sensory tracts. The rest of the white matter 1 is chiefly commissural. Of sensory impulses, those of tOUch and of muscular sense travel along the posterior columns, though there are reasons for thinking that common seiisiliilit v not only ascends along I he posl. Fig. 24. Fig. 25. -c. p. G. B. Fig. 26. c. p. ztions of tracts at various levels Fig. 28. Neuroglia cells. *- y <® ■/•to : mm ' Transverse section of white matter of spinal cord. Fig. 29. -; 2$2 •'.•:-'\vr:::.*r-iij*ffl^^> /t7i<«rtor coma of spinal cord. MURRAY D E LT AND ITS MEMBRANES. 2>Z first trophic realm* — situated in certain regions of the brain surface; and (6) a lower stratum — the second trophic realm — the nerve cells of the anterior horn. Now, in any injury to the cells of the upper stratum — -first trophic realm — the muscles are paralysed and rigid, and any atrophy which follows is by no means rapid, being due to disuse; whereas, if the cells of the anterior vesicular column, second stratum — second trophic realm — be injured, the paralysed muscles are flaccid and degenerate rapidly. (b) Posterior vesicular column. — The cells of the posterior horn are for the most part not arranged in distinct groups. At the base of the posterior horn, however, near its inner angle, is a well-marked collection of cells variously known as POSTERIOR vesicular column, Clarke s column, Dorsal nu- cleus (Fig. 10). Though it is found along only the middle region of the cord, from the level of the 7th cervical to that of the 2nd lumbar nerve, it probably has representations in all regions of the cord, even as far as the bulb. As we have seen, its cells are, on the one hand, connected with the posterior or sensory nerve roots, and, on the other, with the lateral cere- bellar tracts, and through them with the cerebellum of the same side. Injury to the other cells of the posterior horn causes sensory and trophic disturbances of the skin. * These well-chosen terms we owe to John Wyllie, M.D., LL.D., late Lecturer on Medicine, School of Medicine, Edinburgh, D 34 THE SPINAL CORD (c) Lateral vesicular column. — A third group of nerve cells — the nucleus of the intermedio- lateral tract, often called the lateral horn (Fig. 14) — lies at the base of the posterior corn a on its outer side, within the column of grey matter of the same name, and, like it, can be distinguished in the dorsal region only. Table op Vesicular Columns. (See Figs. 10, 11, Plate III., page 14.) ( Antero-internal group. 1. Anterior vesicular J Antero-external group. column. ] Posterior group — found in lumbar I region. 2. Lateral vesicular f Lateral horn — found in thoracic column. \ and lumbar regions. ( Clarke's column; dorsal nucleus — 3 ' column VeSiCUlar 1 found from the 7th or 8th cer " v. vical to the 2nd lumbar nerve. Structure of the nerve cells. — The nerve cells of the grey matter of the spinal cord vary considerably, in size, in shape, and in structure. Firstly as to shape and size : — Most of the cells are multipolar and stellate in transverse section, and have two sets of processes — protoplasmic and axis- cylinder processes. The largest cells are found in the anterior vesicular column, especially in its outer group, and similar cells occur in Clarke's column. In the posterior horn most of the cells are small and spindle-shaped, and are called solitary cells ; others are larger, have long curved processes, and are known as comet cells. Fig. 30. Plate VII. Collaterals. J/t.-.c/ .* piii a. Cells of anterior column. Cells of lateral column. Commissural cells. toot cell. cylindd rocess. Fig. 32. ommistural cell Column cell. Iscending branch. Porta >or root. Descending ' branch. Cells of posterior horn. AND ITS MEMBRANES. 35 Secondly as to structure: — Each cell has a distinct oval nucleus, and contains a network of fibrillge and one or more nucleoli. The body of the cell is fibril- lated and granular, and has no distinct nucleated sheath, differing in this respect from the cells found in the various ganglia throughout the body — e.g., sympathetic and spinal ganglia, which have a dis- tinct fibrous capsule, lined by epithelial cells. According to their behaviour with methylene blue, the cells are distinguished into those (a) in which the nucleus only is stained, called Caryochromes, and (b) into those in which both the body of the cell and nucleus stain, Somatochromes. Two chemically distinct substances, chromatic and achromatic, are found in the Somatochromes. The chromatic mat- ter when stained appears as granules, stripes, cones, blocks and networks. (Fig. 4, Plate VIII., page 40.) It is probably the nutritive material of the cell, for it is used up during its activity. The achro- matic matter, on the other hand, is the essential protoplasmic constituent of the cell. On it tbe life and activity of the cell depend — the activity being accompanied by a swelling of the cell. Processes of Nerve Cells are, as we have said, of two kinds — (1) axis-cylinder processes ; (2) proto- plasmic processes. (1) The axis-cylinder processes are branched or unbranched processes of nerve cells, and are con- nected to the nerve cell either directly, as seen in the motor type of cell, or indirectly through a plexus of fibres, as seen in the sensory type. Composed 36 THE SPINAL CORD of many primitive fibrillse they, in many cases, soon acquire a medullary sheath, and become the axis- cylinders of nerve fibres, which, after a longer or shorter course, end in the grey matter in fine brush- lets of branches. They are centrifugal processes, carrying impulses away from the cell. (2) The protoplasmic or grey processes are a series of delicate ramifying branches which cross and recross each other in all directions (Figs. 17 and 19, Plate V., page 28). They are centripetal processes, carrying impulses to the cell, and may, through their relation to neuroglia and to blood-vessels, subserve the nutrition of the nerve elements. At the cells the protoplasmic and axis-cylinder processes are in most cases continuous with each other ; passing through the cells without interruption : their terminal processes, however, do not anastomose with the processes of other nerve cells, but are merely in contact with the nerve elements. Hence it follows (1) that nerve cells, most likely, do not generate nerve motion, but are nutritive only ; and (2) that contact, and not actual continuity, is sufficient for the conduction of motor, sensory, and reflex impulses along the spinal cord. Classification of the Nerve Cells. 1. Cells with short axis-cylinder processes. In these cells the axone soon breaks up into branches which do not become encased in a medullary sheath, but form a fine plexus of fibrillse. They are the sensory type of cell, and are almost entirely confined in their distribution to the posterior horns (Fig. 19, page 28). 2. Cells with long and distinct axis-cylinder processes (Fijr. 22, page 28), which do not branch, or branch but slightly, and which AND ITS MEMBRANES. 37 ultimately become clothed with a medullary sheath. They are regarded as the motor type of cell, and may be sub-divided into two groups — (a) Root Cells which occur in the anterior horn. Their axis- cylinder process is continued into the anterior nerve roots; and their protoplasmic processes pass in various directions through the grey matter (Fig. 31, Plate VII., page 34). (6) Column Cells which are met with in all parts of the grey matter. They are so called because their axis-cylinder processes pass into the antero-lateral white columns of the same or of the opposite side (commissural cells) of the cord (Fig. 32, Plate VII., page 34). IV.— DEEP ORIGINS OF SPINAL NERVES. (Fig. 14, Plate IV., page 22.) The several spinal nerves are, as you know, mixed nerves, for they contain both motor and sensory fibres. They each spring from the spinal cord by two roots, an anterior — motor, coming from the antero-lateral grooves, and a posterior — sensory, from the postero-lateral grooves. These attachments to the cord are called the superficial origins of the nerves. The deep origins are the nerve cells in the grey matter, to or from which the nerve roots can be traced. (1) The Anterior Nerye Roots — motor roots. Most of the fibres of these roots can be traced from the nerve cells of the anterior horn. The cells from which they spring are called root cells, and the several bundles of white fibres which can be seen passing through the peripheral white substance to the antero-lateral grooves — their superficial origin — are the axis-cylinder processes of these cells. Other fibres of the anterior roots come from cells in the opposite anterior and posterior horns (Fig. 14, page 22). 3§ THE SPINAL CORD (2) The Posterior Nerye Roots — sensory roots — before reaching the spinal cord, pass through the spinal ganglion, in which there are unipolar nerve cells. At a little distance from these cells, the pole, or axis-cylinder process, bifurcates in a T-shaped manner, one limb of the T becoming the distal, the other the central end of a nerve fibre of the posterior root. The central processes enter the cord in two sets — a lateral set which goes to the marginal zone (Lissauer's tract), and a mesial set which enters the postero-external strand. On entering the spinal cord, all the fibres, mesial and lateral, at once divide into ascending and descendiDg branches, which give off collaterals to different parts of the grey matter. (Figs. 31 and 32, Plate VII., page 34.) The de- scending branches after a short course turn at right angles into the grey matter and end free in little tufts. Of the ascending fibres, those of Lissauer's tract lose themselves in the substantia gelatinosa of Rolando ; those of the rest of the column after a long, short, or intermediate course enter the grey matter. At first they occupy the postero-external tract, but as they ascend they become more internal and are found in the fasciculus of Goll, their place in the postero-external strand being taken by fibres of the nerve roots which enter the cord at higher levels : hence it follows — 1st, that these two columns of Goll and Burdach are not physiologically distinct ; and, 2nd, that the greater part of the column of Goll consists of fibres, which come from the legs, whereas the fibres that constitute the column of Burdach mostly come from the upper limbs and neck. AND ITS MEMBRANES. 39 The columns of Goll and Burdach ultimately end in nuclei (nucleus gracilis and nucleus cuneatus) in the medulla; from which fibres can be traced to the opposite side of the brain, in what is known as the superior pyramidal or sensory decussation. Besides the fibres of the posterior roots which are thus connected with the nerve cells of the ganglia, there are others having no such connection, but which merely pass through the ganglia on their way to nerve cells in the spinal cord. On entering the cord these fibres pass at once into the grey matter, through which they can be followed in various directions, even as far as the cells of the anterior horn (Fig. 14, Plate IV., page 22). Note. — Gaskell suggests that each spinal segment has two sets of nerve roots : — 1. A somatic set. — The motor and sensory nerves usually described — (a) the former, the motor, being aganglionic and connected with the cells of the anterior horn ; and (b) the latter the sensory, being ganglionic and connected with the cells of the posterior horn. 2. The other set of nerve roots — the splanchnic set, or visceral set — are distributed to blood-vessels, to the heart, and to other hollow viscera. They consist of two groups— (a) A ganglionic set, connected with the cells of Clarke's column — their ganglia being the lateral ganglia of the sym- pathetic trunk. They are the motor group, and pass to the lateral ganglia of the sympathetic, thence to the walls of blood-vessels and viscera. When stimulated, they cause contraction of blood-vessels and viscera, and give rise to increased activity and to increased waste. 4 o THE SPINAL CORD (h) An aganglionic set, connected with the cells of the lateral horn. They are the inhibitory group, and do not pass to the ganglia of the sympathetic, but run on as rnedullated nerve fibres to ganglia situated in the tissues themselves, Stimulation of this set causes dilatation of the vessels and various inhibitory phenomena — such as cardio-inhibition. They diminish activity and tend to promote repair. Summary. — To sum up, then, we have the spinal cord presented to us as an elongated mass of nervous substance, consisting of white and grey matter; in- vested by three membranes ; having two distinct enlargements; giving origin to thirty-one pairs of spinal nerves ; nourished by spinal vessels, and divided by anterior and posterior fissures into two lateral segments, each with three sub -divisions which can be mapped out into numerous tracts, or systems of varying significance. We see, moreover, that our ideas in regard to the constitution of the spinal cord must undergo at least this modification, that whereas formerly the spinal cord was regarded as made up of two distinct ele- ments — nerve fibres and nerve cells — we must now consider that there is only one nerve element — the neurone — the nerve cell with its protoplasmic and axis -cylinder prolongations. These neurones are arranged in tiers, one above the other, and are connected together by their protoplasmic and by their axis-cylinder processes and collaterals. These facts have important bearings on the physiology of the spinal cord. The following Tables will be found useful for reference : — Fig. 1 'erve fibres with collaterals. Plate VIII. Ganglion of Ou posterior root of a spinal nerve. AND ITS MEMBRANES. 41 -Long Tracts of Conduction to and from the Brain. TABLE OF ASCENDING AND DESCENDING DEGENERATIONS. / 1. Lateral cerebellar tract, from visceral tract. 2. Postero- internal strand, Goll's tract, from posterior roots. 3. Antero- lateral ascending tract of Gowers, V from crossed sensory roots. 1. Crossed pyramidal tract—the chief motor tract — contains 90 per cent, of the motor fibres. It is often called the lateral pyramidal tract. 2. Direct pyramidal tract, fasciculus of Tlirck, usually contains only about 10 per cent, of the motor fibres. Tracts which undergo Ascending Defeneration. Tracts which undergo Descending Defeneration, II. — Short Commissural Tracts. These neither undergo ascending nor descending degeneration, or at least for no great distance, and are therefore considered to be commissural fibres between different segments of the cord, 1. The posteroexternal strand — fasciculus of Burdach. 2. Antero-external column — anterior root zone. 3. Part of the lateral column — lateral limiting layer. •4. Lissauer's tract— marginal zone. Table of the Order in which the several Tracts of White Matter receive their Medullary Sheaths. 1. Anterior and posterior root fibres. 2. Ground fibres of the antero-external column. 3. Ground fibres of the postero-external column. i. Lateral mixed zone. 5. Limiting layer and anterolateral ascending tract. 6. Posteromedian column. 7. Direct lateral cerebellar tract. 8. Crossed and direct pyramidal tracts. In man these tracts get their medullary sheath, for the first time, at birth. 42 THE SPINAL CORD. GOWEES' TABLE. SHOWING THE APPROXIMATE RELATION TO THE SPINAL NERVES OF THE VARIOUS MOTOR, SENSORY, AND REFLEX FUNCTIONS OF THE SPINAL CORD. MOTOR. (Nerves. MOTOR. SENSORY. Lower 11-ck muscles, Middle part of Trapezius Lower part of Trapezius J and dorsal muscles Lumbar muscles Peroneus, I. r Flex, of J ankle, Ext. "| of aukle V. J •*■ \ Small rotators of head n i Depressors of liyoid 3 Lev. ang. scapulae J Diaphragm ~\ fcj ISerratus I ~' /Flex, of elbow j o n ~{ Supinators I ^ _ / Ext. wrist and fingers ) Ext. elbow /" Flex, wrist and fingers o J Prouators I -j -j f Muscles of hand 2\ 3 4 5 6 7 8 9 10' 11 12 , 1 J 2 ^ Cremaster o f Flexors of hip I Extensors of knee 4 f Abductors of hip k I Ext. and abduct, of hip Scalp Intercostals .Abdominal muscles S 1 ) Flexors of knee Intrinsic muscles of foot Perineal and muscles Co. 1} Neck and upper part of chest j- Shoulder -Arm, outer side Radial side, fore- arm and hand; thumb Arm, inner side ulnar side of forearm and hand ; tips of fingers Front of thorax > Ensiform area Abdomen (Umbilicus 10th) |_ Buttock, | upper part Groin and Scrotum (front) C outer side a front V. inner 3 \ Thigh [ J pinner side " \ Leg, inner side 1 I C Buttock, lower part I J Back of thigh 2 T | Leg ) except inner I v & foot j i^art 4 V Perinseum and anus 5, (Skin from coccyx to 30 f :U1US REFLEX. *j y Scapular 8 1; 2 3 4" 5 a \. Epigastric 7 Abdominal / 1 Cremasteric Knee-jerk I ure I Gluteal J \ Fcot- ±\ J clonus 2 V Plantar 43 SECTION II. BRAIN AND ITS MEMBRANES. The term Brain, or Encephalon, includes all that part of the great central nervous system which is enclosed within the cavity of the cranium. Invested by three membranes or meninges, it presents, as you will afterwards see, four distinct sub-divisions : — (1) The Cerebrum, a large convoluted mass, or big brain ; (2) the Cerebellum, or little brain ; (3) the Pons Varolii, a white transverse band uniting the two halves of the little brain ; and (4) the Medulla Oblongata, or bulb, the enlarged upper end of the spinal cord (Figs. 33 and 34, page 44). From a developmental point of view, the brain is divided into six parts : Telencephalon, Diencephalon, Mesencephalon, Isthmus Rhombencephali, Meten- cephalon, Myelencephalon ( see " Development, " page 190). We shall describe (1) the membranes and vessels of the brain, and (2) the several parts of the brain ; but before doing so we shall, for convenience, give seriatim the following dissections : — Dissection. — 1. To expose the Membranes of the Brain. — Make an incision along the middle line through the entire thickness of the scalp, from the root of the nose in front to the external occipital protuberance behind. Turn back the pericranium to the level of the ears, and then saw through the outer table of the skull-cap along a line passing round the skull, about half-an-inch above the 44 THE BRAIN external occipital protuberance behind, and about the same distance above the supra-orbital arch in front. Break through the inner table with the chisel and forcibly raise the detached skull-cap, which you will find more or less adherent to the subjacent mem- brane, especially along the lines of the cranial sutures. Branches of the meningeal arteries will be seen ramifying on the outer surface of the exposed membrane, between it and the bone. 2. To Remove the Brain. — With a sharp pair of scissors cut through the dura mater at the same level at which you have sawn the bones, and reflect it upwards towards the top of the head. Examine as far as you can the strong process, falx major, passing down mesially between the cerebral hemispheres. Divide this pro- cess in front where it is attached to the crista galli of the ethmoid, and cut through the veins which enter the sinus contained within its upper border. Eaising the falx out of the longitudinal fissure, turn it back, but do not cut it behind. Now pass the fingers of the left hand beneath the fore-part of the brain, and gently raise it from the anterior cranial fossae, taking care to detach the small white rounded bands, olfactory lobes, 1st pair of nerves, from the cribri- form plate of the ethmoid. The optic nerves, 2nd pair, and the two internal carotids will now be seen close to the anterior clinoid processes, and should be divided. Piercing the dura mater, ex- ternal to the carotid, are the round 3rd nerves, and in the free margin of the tentorium cerebelli, now exposed, are seen the slender lf.th nerves. Cut through these, and through the infundi- bulum which passes down towards the sella turcica of the sphenoid. Then with the point of the knife make an incision through the margin of the tentorium on each side, just behind and parallel to the upper margin of the petrous part of the temporal bone, carry- ing the incision as far back as necessary, but being careful not to injure the parts beneath. You will now see the following nerves, which will require to be divided one after another. Just below the anterior end of the tentorium will be found the large 5th nerves ; nearer the middle line, the slender 6th nerves; below and external to the 5th, the 7th or facial, and 8th or auditory nerves, and the auditory artery ; immediately below the 8th are the 9th, 10th and 11th nerves. Cut the 9th, the glossopharyngeal, and the 10th, the vagus, but leave intact the spinal part of the spinal accessory nerve ; lower down near the middle line is the 12th or hypoglossal nerve, consisting of two bundles, which pierce the dura mater by two Fig. 33. Plate IX. ( 'trrhrum. Pom Varolii, Superior cerebellar p& Middle, cerebellar peduncle. Cerebellum. '^Pf§Ef ' Inferior cerebellar ped'jailr. Fia. 34. Cerebrum, Dura mater. knoid. later. I space N 5th ventricle, 3rd ventricle, Subarachnoid space. 4th ventricle. Medulla. Isthmus Rhombcncep Ti ntorium cerebelli y Cerebellum. 'Foramen Majendie. URRAY OEC AND ITS MEMBRANES. 45 separate openings. Next pass the knife as far down the spinal canal as possible, and divide the spinal cord, the nerve roots attached to its sides, and the vertebral arteries as they wind round from the back. Snip through the veins of Galen, and the brain can now be easily removed from its bed, and should be at once placed in spirit. Leave it there for a few days ; then examine it and carefully remove the pia mater, except at the back of the brain between the cerebrum and cerebellum, in order not to disturb a process of pia mater — the velum interpositum — which passes into the brain at this point. CHAPTER I. MEMBRANES OF THE BRAIN. To examine the first of these membranes — the cranial dura mater — replace, as far as you can, the falx major and the tentorium cerebelli, and fasten them in their places by a few stitches. It will, how- ever, be far more satisfactory for you if you can obtain a specimen specially prepared to show the arrangement of the dura mater (Plate XL, page 52). I.— THE DURA MATER. The Cranial Dura Mater is a dense white fibrous membrane, rough externally, but smooth and polished within. It is lined by a layer of epithelial cells similar to that which lines the spinal dura mater. Composed of two layers (though these are not distinct from each other) — an inner, which sends processes between the parts of the brain, and an outer, which forms the endosteum of the inner table of the cranium — it adheres to the bones of the skull, 46 THE BRAIN in two places especially : (1) along the lines of the cranial sutures, and (2) at the base of the cranium. This latter fact accounts for the rare occurrence of accumulations of pus or blood at the base of the brain between the dura mater and the bone. The dura mater, moreover, sends a tubular process along each of the several cranial nerves as they leave the skull through their various foramina; part of this tubular process forms a sheath to the nerve, part of it becomes attached to the pericranium. The dura mater also passes into the orbital cavities, and there blends with the periosteum. At the lower margin of the foramen magnum the dura mater is closely attached to the bones, and becomes continuous with the spinal dura mater. Along certain lines the two layers, of which the cranial dura mater is composed, separate from each other, leaving variously shaped channels, called VENOUS sinuses, for the passage of the blood from the brain into the venous system. Of the partitions given off from its inner layer, two — the FALX CEREBKI and the FALX CEREBELLI — are vertical ; the third, the TENTORIUM CEREBELLI, is usually said to be horizontal, though it is far more vertical than horizontal, for it slopes downwards and backwards. 1. The Falx Cerebri — so called from its sickle- shaped form (Fig. 37, page 52) — is the vertical mesial process of dura mater which is lodged in the great longitudinal fissure, and separates the inner surfaces of the cerebral hemispheres from each other. AND ITS MEMBRANES. 47 In front it is pointed, and is attached to the apex and to the posterior margin of the crista galli. Behind it widens out, and is fixed along the middle line to the upper surface of the tentorium cerebelli. Its upper convex margin contains the superior longitudinal sinus, and adheres to the ridges seen on each side of the median depression, on the inner aspect of the vault of the skull. The lower margin contains the inferior longitudinal sinus. It is concave and free, and in front is in contact with the upper surface of the corpus callosum, though behind it is some distance from that body. 2. The Falx Cerebelli (Fig. 37) is the small median vertical triangular partition attached behind to the internal occipital crest, and above, at its widest part, to the mid-line of the under surface of the ten- torium cerebelli. Its free concave margin projects forwards and fits into the notch between the halves of the cerebellum. Its posterior attached margin contains the occipital sinus, which is single above but bifid below. 3. The Tentorium Cerebelli (Fig. 37), supporting the posterior part of the cerebrum, slopes forwards and upwards, somewhat horizontally, as an arched tent- like partition between the big brain and the little brain. The posterior convex margin is attached behind to the transverse ridges on the inner aspects of the occipital, parietal, and temporal bones, and further forwards it is fixed to the upper margin of the petrous part of the temporal bones. It ends in front at the posterior clinoid processes. This margin 4 8 THE BRAIN contains the lateral sinuses behind and the superior petrosal sinuses in front. The anterior concave margin, free in the greater part of its extent, is attached on each side by a narrow slip to the anterior clinoid processes. Between this margin and the dorsum sellse of the sphenoid is an oval opening for the passage of the crura cerebri, the superior peduncles of the cerebellum, and the posterior cere- bral arteries. Along the middle line of the upper surface of the tentorium runs the straight sinus, which receives the veins of Galen from the interior of the brain (Fig. 37, page 52). The MINUTE structure of the dura mater of the brain is similar to that of the spinal cord. It consists of layers of fibrous tissue. The larger arteries lie near the outer surface of the membrane, whereas the veins are embedded in its midst or lie between its layers where these exist. There are few capillaries in its substance, but on its inner aspect is a rich network of vessels covered by epithelial cells and surrounded by peri - vascular spaces which by means of stomata communicate with the sub-dural space (Robertson). The Agreements and Differences in the Arrangement of the Cranial and Spinal Dura Mater may be arranged as follows \ (a) Agreements — 1. Both are fibrous membranes. 2. Both invest their respective organs. 3. Both form the outer boundary of the sub- dural space. Fig. 35. Plate X. Dura mater. - Middle mugial artery /'in mater. A rachnoid. Krcvlar sinus, asverse sinus. petrosal sinus *V._ Veins of orbit. S3 CarvenovA sinus. Superior petrosal sinus. Lateral sinus. Occipital sinus. AND ITS MEMBRANES. 49 b) Differences — 1. The cranial dura mater forms the perios- teum of the bones; the spinal does not, though some authors hold that there are two layers in the spinal dura mater — the outer forming the periosteum of the bones. 2. The cranial dura mater sends off processes between the parts of the brain ; not so the spinal dura mater — it gives off no such processes. 3. The cranial dura mater, by the separation of its constituent layers, encloses the various venous sinuses; the spinal dura mater does not form sinuses. The cranial dura mater receives its nerve supply from the 4th and 5th cranial nerves, and from the sympathetic. Pacchionian Bodies (or Glands) are small white elevations, which indent the inner surface of the ver- tex of the skull, along each side of the middle line. They are said to be out-growths of the villi which normally exist in the arachnoid. Their exact nature and use is unknown. Some hold that they are mere excretions, others that they are communications between the sub-arachnoid space and the venous sinuses of the dura mater, others that they are ligaments which fix the pia mater to the dura mater and help to suspend the brain. II.— THE PIA MATER, (Fig. 34, page 44, and Fig. 35, page 48.) The Pia Mater of the brain, like that of the spinal cord, is an extremely fine vascular membrane, com- posed of a plexus of capillary blood-vessels held E 5o THE BRAIN together by delicate connective tissue (Fig. 35, page 48). It is practically one layer with the arachnoid (page 51). Covering the surface of the brain, it dips into the various fissures between the convolutions; and from its inner surface numberless blood-vessels emerge for the nourishment of the substance of the brain. Besides the septa between the convolutions, the pia mater also sends into the brain through a large fissure, called the great transverse fissure, a special prolongation, the velum interpositum, the margins of which are wrinkled and folded, forming a vascular fringe, the choroid plexus ; and a similar vascular process of pia mater, known as the tela choroidea inferior, or choroid plexus of the 4th ventricle, lies in the roof of that cavity (see " Ven- tricles," page 141). In structure the pia mater of the brain corre- sponds to the inner of the two layers of the pia-arach- noid described on the following page. It receives its nerve supply from the 3rd, 5th, 6th, 7th, and 8th nerves, as well as from the sympathetic. III.— THE ARACHNOID. The Arachnoid is a smooth, glistening, trans- parent, colourless membrane, situated between the dura mater and the pia mater (Fig. 35). Formed of a single layer, it is, on the one hand, separated from the dura mater by veins, and, on the other, envelops the brain but does not pass into its fissures. It is closely connected to the pia mater beneath by the sub-arachnoid trabeculse, which are far more numer- ous than those in the spinal cord. At certain spots, AND ITS MEMBRANES. 51 especially at the base of the brain, the pia mater and the arachnoid are more widely separated from each other, forming the SUB-ARACHNOID SPACE, which contains most of the cerebro-spinal fluid. Minute structure. — The arachnoid is not a dis- tinct membrane from the pia mater, and the two are often described together under the name pia-arach- noid. This consists of two layers of epithelium — a superficial and a deep — the latter being in contact with the brain. The two layers are united by numberless trabecular covered by epithelial cells. The nerve supply to the arachnoid is probably the 5th, the facial, and the spinal accessory nerves. Sub-dural and Sub-arachnoid Spaces (Fig. 34). — The space between the dura mater and the arachnoid is called the sub-dural space, and is lined by epi- thelial cells. The space between the arachnoid and pia mater is the sub-arachnoid space, and is crossed by the sub-arachnoid trabecular. It contains cerebro- spinal fluid. Both its inner and outer walls are lined by epithelial cells, which also cover the surfaces of the trabecular. The sub-arachnoid space is most distinct in the following situations — (1) at the great longitudinal fissure ; (2) at the base of the brain, in the triangular interval, called the inter-peduncular or opto-peduncular space; and (3) posteriorly, between the cerebellum and the medulla oblongata. Cerebro-spinal Fluid. — Most of this fluid is contained in the meshes of the sub-arachnoid tra- becule, though a little is found in the sub-dural space. 52 THE BRAIN It communicates with the fluid in the central canal of the spinal cord, and in the ventricles of the braiD, through an opening — the foramen of Majendie — in the roof of the 4th ventricle (Fig. 34, page 44), and through two similar lateral openings in the ven- tricular roof between the medulla and cerebellum. Corresponding communications also exist at the inferior horns of the lateral ventricles. The cerebro- spinal fluid is not merely intended to fill the sub- arachnoid space, but helps to protect the nerve centres from sudden shocks ; acting, in fact, as a water-bed (Hilton) on which the brain rests. According to Foster it is probable that the cerebro- spinal fluid, being of the nature of lymph, also subserves the nutrition of the brain. IV.— VENOUS SINUSES. (Figs. 36, 37, Plates X., XL, pages 48 and 52.) The sinuses of the brain, formed by the separation of the two layers of the dura mater, are lined by epithelial cells continuous with those lining the interior of the veins. These sinuses are fifteen in number — five paired and five single. The single sin- uses are the superior and the inferior longitudinal, the straight, the circular, and the transverse. The paired set includes the two lateral, the two superior, and the two inferior petrosals, the two cavernous and the two occipital. Other smaller sinuses are sometimes described. Plate XI. Fio. 37. Inferior longitudinal sinus. Superior longitudinal sinus. Falx major. rramen caxumJj M-i- Straight sinus. ' -Tentorium, cerebeUi. Lateral sinus. Occipital sinus, petrosal sinus. Superior petrosal sinus. Fig. 38. Parasinoidal- sinus. Superior longitudinal sinus. Pacchionia.n body, Bone. Dura mater. Subarachnoid space. AND ITS MEMBRANES. 53 1. The Superior Longitudinal Sinus (Figs. 37, 38, page 52) begins in front at the foramen caecum as a small vein, which often communicates with the veins of the nose, and thence arches upwards and backwards in the convex margin of the falx cerebri to reach the internal occipital protuberance, where it is joined by several other sinuses at the triangular dilatation, called the torcular Herophili,* or the meet- ing of the sinuses. Cut open the sinus in its entire length and you will see that it is wider above than below, being triangular in section, and that it in- creases in size as it passes backwards, its cavity being crossed by many slender bands — chordaa Willisii. Many veins, from the substance of the brain, pour their blood into this sinus. They run mostly from behind forwards — that is, in the reverse direction to the blood current in the sinus. They pierce the wall of the sinus by slit-like openings, which act as valves, and thus prevent regurgitation of the blood. Before entering the sinus they pass through small spaces, called the para-sinoidal spaces, which lie on each side of the superior longitudinal sinus (Fig. 38, page 52). Other smaller veins enter this sinus from the diploe of the surrounding bones, and it receives an emissary vein through the parietal foramen. 2. The Inferior Longitudinal Sinus (Fig. 37) is really a small vein contained in the free concave border of the falx major. It commences in front * As HoUlen points out, the term torcular is a mistranslation of the original word, (TcoKr]v— a canal or gutter. 54 THE BRAIN by minute venous radicles within the substance of the falx, and ends behind in the straight sinus. 3. The Lateral Sinuses (Fig. 36).— These are two in number, right and left, the right being usually the larger of the two. Through them most of the venous blood from the brain reaches the internal jugular veins. They commence at the internal occipital protuberance, and running out- wards and downwards in the attached margin of the tentorium cerebelli, in the grooves on the inner surface of the occipital, parietal, and temporal bones, finally turn forwards to end in the bulb of the internal jugular vein. Blood enters these sinuses from the superior and inferior petrosal sinuses, from the inferior cerebral and cerebellar veins, from the diploe of the bones, and from the scalp by emissary veins. 4. The Straight Sinus (Fig. 37) lies in the middle line of the upper surface of the tentorium cerebelli, where the falx major is attached. Behind, it ends at the meeting of the sinuses; while in front it receives the inferior longitudinal sinus and the veins of Galen, the latter bringing blood from the third and the lateral ventricles of the brain. Some cerebral and cerebellar veins also open into this sinus. 5. The Cavernous Sinuses (Fig. 36), so called from the spongy appearance of their interior, are placed in the grooves on each side of the body of the sphenoid. They are oval in vertical transverse section, and contain, in their thickened outer wall, AXD ITS MEMBRANES. 55 the third, the fourth, and the ophthalmic division of the fifth nerves ; while internally, and separated from the blood of the sinus by a thin lining- membrane, is the internal carotid artery, with the sixth nerve on its outer side. These sinuses com- municate with the circular sinus, and with the superior and inferior petrosal sinuses; and receive small cerebral veins and the veins from the orbit (Fig. 36, page 48). 6. The Circular Sinus (Fig. 36) surrounds the pituitary body in the sella turcica. At each side it communicates with the cavernous sinuses. 7. The Petrosal Sinuses (Fig. 36) are two on each side — the superior and the inferior. The Superior Petrosal sinus lies along the upper border of the petrous part of the temporal bone, and connects the cavernous sinus with the lateral sinus. The Inferior Petrosal sinus runs in the groove between the occipital bone and the petrous part of the temporal bone. It unites the cavernous sinus to the lateral sinus after the latter has passed out of the skull to become the internal jugular vein. Cerebral, cerebellar, tympanic, and auditory veins pour their blood into one or other of these sinuses. 8. The Transverse or Basilar Sinus (Fig. Bo), sometimes double, is placed across the basilar process of the occipital bone, and connects together the inferior petrosal sinuses. It communicates below with the anterior spinal veins. 56 THE BRAIN 9. The Occipital Sinuses (Fig. 36) commence near the foramen magnum as two small channels which run upwards in the attached margin of the falx cerebelli, and then join above to form a single sinus which opens into the torcular Herophili. Emissary Yeins are small vessels which pass through foramina in the skull, and unite the veins of the scalp with the cranial sinuses. If it were not for these veins injuries to the scalp would lose half their significance (Treves). The Meningeal Arteries, which supply the dura mater with blood, ramify in branching grooves on the inner surface of the cranium, between it and the outer surface of the dura mater (Fig. 35, page 48). From the fossae in which they ramify they are called the anterior, the middle, and the posterior meningeal arteries, and they arise from the ethmoidal, ascend- ing pharyngeal, internal maxillary, occipital, and vertebral arteries. For the description of these vessels see your dissecting manual. We shall now pass on to examine the vessels of the brain. AND ITS MEMBRANES. 57 CHAPTER II. VESSELS OF THE BRAIN. (Figs. 39, 40, Plate XII., page 58.) Dissection. — Remove with great care the pia mater, and in doing so, at the under surface of the brain, dissect out the vessels and the cranial nerves. I.— ARTERIES. The arteries of the brain are derived from the two internal carotids and from the two vertebrals. I. The Internal Carotid Arteries, when they reach the anterior clinoid processes, give off the ophthalmic branches, and then divide into anterior, and middle cerebral, and posterior communicating. 1. The ANTERIOR cerebral arteries run for- wards to the fore part of the great longitudinal fissure, and curving round the anterior end of the corpus callosum, pass backwards on its upper surface, under the name of the arteries of the corpus callosum. At their commencement they are joined by a short transverse branch — the anterior communicating; while behind they anastomose by small branches with the posterior cerebral arteries. They supply blood to the frontal and olfactory lobes, to the optic nerves, to the corpus callosum, and to the anterior perforated spots (Figs. 39, 40). 2. The MIDDLE CEREBRAL or SYLVIAN ARTERIES are the largest branches of the internal carotid, and 58 THE BRAIN run upwards and outwards in the fissure of Sylvius till they reach the surface of the island of Reil, where they ramify in the pia mater, forming part of the cortical system of arteries. They anastomose with the anterior and posterior cerebral arteries, and are distributed by four branches to part of the frontal, parietal, and temporal lobes (Figs. 42, 43, page 60). Other branches of the Sylvian artery are furnished through the anterior perforated spot to the corpus striatum, and are all terminal arteries, belonging to the " ganglionic system " of branches. They are the lenticular, the lenticulo- striate, the lenticulo-optic; and will be again re- ferred to in describing the nuclei of the ventricles of the hemispheres. The middle cerebral artery is the one chiefly concerned in cerebral haemorrhage. 3. The ANTERIOE CHOROIDAL ARTERIES, one Or two in number, are either branches of the internal carotids or of the middle cerebrals. Entering the fissure between the temporal lobe and the cms cerebri (Fig. 40, page 58), they reach the descending cornu of the lateral ventricle, and there form the vascular fringe called the choroid plexus. 4. The POSTERIOR COMMUNICATING ARTERIES mn backwards and join the posterior cerebral arteries (branches of the basilar artery), and thus is established a free anastomosis between the carotids and the vertebral s. IT. The Vertebral Arteries — branches of the sub- clavian — enter the foramen magnum by perforating the posterior occipito-atlantal ligament and the dura Fio. 39. Plate XII. Fori. ,r cen hral. Middle cerebral. Carotid. \-- Corpus callosum. Postei ior cerebral. Superior cerebellar. Basilar. Inferior cerebellar. 'Vertebral. Fio.-40. p?' communiccUiny. Posteriorr ■aiHiiumieating ior cerebellar, averse branch. - Posterior inferior cerebellar, Anterior spinel, A nterior cerebral. Middle cerebral. - Posterior cerebral. - Basilar. A nterior inferior . cerebellar. Vertebral. Arteries of the Brain. AND ITS MEMBRANES. 59 mater. They then curve round to the anterior surface of the medulla between the 12th cranial nerve and the anterior roots of the 1st cervical nerve. At the lower border of the pons they unite to form a single trunk — the basilar artery — which may be seen running in the groove on the front of the pons, till it reaches its upper margin, when it divides into two branches — the posterior cerebral arteries (Fig. 40, page 58). 1. BrAJsX'HES of the vertebral arteries. — Only one branch, the posterior inferior cerebellar, which may, however, sometimes be a branch of the basilar, is given off by the vertebral artery to the brain. It supplies the under surface of the cere- bellum. 2. Branches of the basilar (Fig. 40): — (1) Transverse branches, three or four in number, run transversely outwards on the pons. One of them, the internal auditory branch, enters the inter- nal auditory meatus. (2) The anterior inferior cerebellar arteries are distributed to the anterior part of the lower surface of the cerebellum, and anastomose with the other cerebellar arteries. (3) The superior cerebellar arteries, given off near the termination of the basilar, supply the upper surface of cerebellum, and send branches to the valve of Yieussens, to the pineal gland, and to the velum interpositum. 60 THE BRAIN (4) The posterior cerebral arteries — the terminal branches of the basilar — curve outwards and back- wards, round the crura cerebri, to the under surface of the posterior cerebral lobes, supplying them and anastomosing with the anterior and the middle cere- bral arteries. They send branches to the posterior perforated spot and to the velum interpositum (posterior choroidal), and give three chief cortical branches (Fig. 40). Circle of Willis (Figs. 39, 40, page 58).— This important arterial inosculation takes place at the base of the brain between the internal carotids and the vertebrals. In front the circle is formed by the anterior communicating artery which joins together the two anterior cerebral arteries ; behind by the two posterior cerebral arteries, branches of the basilar; and on each side by the internal carotids, the anterior cerebrals, and the posterior communi- cating. CIRCLE OF WILLIS. In Front : Anterior Communicating. On each Side : Left. Right. Anterior Cerebral. Anterior Cerebral. Internal Carotid. Internal Carotid. Posterior Communicating. Posterior Communicating. Behind : The Two Posterior Cerebrals, branches of the Basilar. Fid. 12, Plate XIII. Area of anterior cerebral. Area, of middle cerebral. Area of poster tor cerebral. Fig. 43. „ A rea of anterior cerebral. X A rea of m id die cerebral. A rea of posterior cerebral. Diagram op Distribution of Arteries of Brain. AND ITS MEMBRANES. 61 II.— VEINS. Cerebral Veins. — The veins of the brain do not accompany the arteries, but open into the various sinuses in the dura mater. In the cerebrum we have two sets of veins — the one superficial, on the surface of the brain, the other deep. Of the super- ficial set those above open into the superior longi- tudinal sinus ; those on the lateral and under aspect of the brain open into the lateral, the cavernous and the superior petrosal sinuses. The deep set of veins, gathering the blood from the interior of the brain, enters the veins of Galen, and thus pours its blood into the straight sinus. Cerebellar Yeins. — The veins on the upper surface of the cerebellum enter the veins of Galen and the straight sinus ; those on the under surface end in the occipital, and in the lateral sinuses. Special Characters of the Cerebral Circulation are — 1. The free anastomosis at the circle of Willis, which provides a ready supply of blood from other vessels in case of the sudden blocking of any of the more direct channels. 2. The tortuous course through bony canals of the arteries as they enter the skull, thus mitigating the force of the heart's beat. 3. Their ramifications in the pia mater before entering the substance of the brain. 62 THE BRAIN 4. The thinness of the arterial walls, and the smallness of the capillaries. 5. Except at the circle of Willis, and at their terminations by capillary vessels, there is little communication between the branches of the cerebral arteries, so that if any artery be obstructed, the nutrition of the area to which it is supplied becomes impaired. 6. The existence of venous sinuses which are without valves, and which do not run with the arteries — the larger arteries, in fact, having no companion veins. Plates XIII., page 60, and XIV., page 62, will give you an idea of the distribution of the chief arterial branches to the cerebral cortex. The numbers 1, 2, 3, and 4 indicate the branches and the parts to which they are distributed. Plate XIV. nterior cerebral. Vac, fotor Pig. 44. II rii i ini . i /•///. Let/. Hearing. irea of posterior cerebral. Area of middle cerebral. -- -Visual \- - Vision. Fig. 45. A rm and shoulder. Trunk, of anterior cerebral. ea of middle cerebral Smell and taste, Touch. Vision. Area of posterior cerebral. Diagram op Arteries and Chief Centres in Brain. AND ITS MEMBRANES. 63 CHAPTER III BRAIN AND ITS SUB-DIVISIONS. (Figs. 33, 34, page 44.) General Outline of the Brain. — Placing the brain before you, you will see that it is an oval-shaped mass of nervous substance, that has not inaptly been likened to the kernel of a walnut ; for, says Wilks, " one cannot but be forcibly struck with the resem- blance between the human head and a walnut. There is, first, the pericranium and the skin, then the bone and the shell ; within a dura mater and a thick membrane lining the shell of the fruit; then the pia mater and the delicate membrane coverinsf the kernel, which is again made up of convolutions into two masses joined together by a commissure or corpus callosum." Viewed from above, the surface of the brain is seen to be arched and convex, and presents many tortuous folds or convolutions of nerve substance, with inter- vening fissures or furrows, the whole giving to the exterior of the organ a most characteristic appearance. Along the middle line of this aspect of the brain runs a deep longitudinal fissure, which, when the brain is in situ, lodges the falx cerebri and divides 64 THE BRAIN the mass into two similar halves. These symmetrical halves are called the cerebral hemispheres, and they together form the first great division of the Encephalon — the CEREBRUM — which is united to the rest of the brain and to the spinal cord by two masses of nerve substance known as the CRURA CEREBRI. On separating the walls of this median longitudinal fissure you will find that, in front and behind, the cleft extends right through to the base of the cerebrum ; but that, in the middle of its extent, it is interrupted below by a transverse band of nerve fibres, the CORPUS CALLOSUM, or great commissure of the brain. The under-surface or base of the brain, convo- luted like the upper surface, is very irregular, for it fits into the corresponding fossae at the l>ase of the skull. On this aspect you will recognise the other sub-divisions of the brain (Fig. 34, page 44), viz. : (1) the cerebellum or little brain, lying behind and below the posterior part of the cerebral hemispheres : (2) the PONS varolii which, when viewed from the front, appears as a broad white band, crossing transversely between the two halves of the cere- bellum ; (3) the MEDULLA OBLONGATA (Fig. 34), or bulb, placed between the pons above and in front, and the cerebellum behind, and serving to connect these several parts of the brain with the spinal marrow below. The sub-divisions of the brain, from a developmental point of view, are given on page 194. We shall describe (1) the MEDULLA; (2) the PONS; (3) the cerebellum ; and (4) the cerebrum. AND ITS MEMBRANES. 65 I.— MEDULLA OBLONGATA. (Figs. 46 to 49, Plate XV., page 66). Dissection. — If you have only one brain to work upon it will be best to pass on at once to the study of the cerebrum (page 113) and afterwards return to the medulla. If you have more than one brain, then cut through the crura cerebri, and begin the examina- tion of the medulla. The Medulla Oblongata — Myelencephalon — the most complicated portion of the central nervous system, is, as you already know, the expanded upper end of the spinal cord, and, like it, is composed of grey and white matter. It has anterior, posterior, and lateral aspects. The anterior surface rests upon the basilar process of the occipital bone ; the posterior is hidden by the cerebellum. In general outline it is more or less conical, with its long axis nearly vertical, its base being directed upwards and forwards towards the Pons Varolii, its apex downwards and backwards, and continuous with the spinal marrow at the lower border of the foramen magnum. In length, the medulla measures about one and-a-quarter inches; in breadth, at its widest part, about one inch ; in thickness about half-an-inch. On its ventral or anterior aspect it is convex, and is limited above by the lower border of the pons ; below, by what is known as the decussation of the pyramids — several bands of fibres passing, at the bottom of the anterior median fissure, from one side of the medulla to the other. Behind, that is on its dorsal aspect, the medulla, in its lower half, is convex, and resembles the spinal cord, but, in its upper half, it expands laterally and becomes flattened from before F 66 THE BRAIN backwards, forming part of the rhomboida] depression called the floor of the 4th ventricle. On this aspect, the upper boundary of the medulla is marked by several transverse lines — the strice acusticce — run- ning across the widest part of the ventricular floor (Fig. 47); the lower boundary is purely artificial, corresponding with the lower margin of the foramen magnum. The lateral aspect of the medulla sup- ports an oval eminence, the olivary body, crossing the lower part of which, as well as the surface of the medulla just below it, you will be able to make out several transverse streaks, the external arciform fibres (Figs. 46, 49), the significance of which you will understand hereafter. 1. FISSURES OF THE MEDULLA. Like the spinal cord the medulla is a symmetrical organ, being divided by superficial median clefts — anterior and posterior — into two similar lateral seg- ments, which are again sub-divided into anterior, posterior, and lateral areas by the continuation up- wards, though not in a direct line, of the place of origin of the anterior and posterior spinal nerve roots (Figs. 46 and 48, page 66). 1. The anterior median fissure of the medulla, a direct continuation upwards of the anterior median fissure of the spinal cord, ends above, at the lower margin of the pons, in a slight recess, needlessly named the foramen caecum. At the lower limit of the medulla, this same fissure is interrupted by the bands of fibres known as the decussation of the pyramids (Fig. 46, page 66). Plate XV. Fig. 46. j)(ic nerve, ic tract. Optic commissure. Crus cerebri. Fig. 47. Nates. Testes. fiddle cerebellar . peduncle. 4th ventricle. External , . . eiform fibres. W"™ f™ rrai area Decussation of pyramids. Fasciculus teres Locus c&ruleus. Superior cerebellar peduncle. Superior- fovea Stria; usticve. —Tuberculum acusticam. Inferior cerebellar peduncle. A la einerea. Anterior median f. Fig. 48. Superior medullary vdmn Fig. 49. \la... t tubercle Claw Calamus criptorius %nd obex. mmedianf. erttl. dorsal s Superior cerebellar ■ peduncle. Pons. "R >st if or >n body. Olivary body -7'i hervk of Rolando. Fv/rwculus of Rolando. -Funiculus cuneatus. Funiculus gracilis. Rostenor median f. Crus cerebri. Corpora geniculata . _ externa. ..-- interna. Rulvinar. \- Corpora quadriyemina. Middle pedunch " of -cerebellum. Olivary body. External arciform fibres. AND ITS MEMBRANES. 67 2. The POSTERIOR MEDTAN FISSURE, continuous with the posterior median fissure of the spinal cord, expands above into the shallow rhomboidal space called the floor of the 4th ventricle, alons - the centre of which runs a mesial groove, iu a line with the median fissure of the cord. External to the pos- terior median fissure of the medulla are two minor sulci — the paramedian and lateral dorsal sulci (Figs. 47, 48). 3. The lateral fissures, continuous with the lateral fissures of the cord, give origin to the roots of the 9th, 10th, 11th and 12th pairs of cranial nerves — the 9th, 10th and 11th pairs springing from the continuation of the postero-lateral groove, and the 12th pair from the continuation of the antero- lateral groove. 2. WHITE MATTER OF THE MEDULLA. (Areas of the Medulla.) By means of the above fissures the surface of each half of the medulla is marked out into three areas, viz.: an anterior area or pyramid, between the anterior median fissure and the line of origin of the 12th nerve ; a lateral area or pyramid, between the 12th nerve in front and the 9th, 10th, and 11th nerves behind ; and a posterior area or pyramid, between the posterior median fissure and the 9th, 10th, and 11th pairs of nerve roots. It will be convenient to begin with the description of the posterior area (Figs. 46 to 49, page 66). 1. Posterior Area. — This area, lying between the posterior median fissure and the line of origin of the 68 THE BRAIN 9th, 10th, and 11th pairs of nerve roots, may be divided into an upper or anterior part, and a lower or posterior part. The upper portion of this area enters into the formation of the floor of the 4th ventricle, a cavity which should naturally be described at this stage, but, owing to the fact that many of its parts are connected with the pons and cerebellum, its description must be deferred until we have treated of that division of the brain. The lower portion of the posterior area, very similar in appearance to the spinal cord, is, for the most part, formed by the upward prolongation of the various white tracts of the posterior column of the cord, the several strands there defined changing, when they reach the medulla, their position, their arrangement, and their names. These changes we shall now describe. In dealing with the white substance of the posterior column of the cord, you will doubtless remember that we called your attention to the existence, in the cervical region, of a strand of fibres close to the posterior median fissure, called the postero-intemal strand — fasciculus of Ooll. Now, traced into the medulla, this strand, which lies between the posterior median and the paramedian fissures, becomes more prominent, and at the point where the central canal of the cord becomes the cavity of the 4th ventricle, the fasciculus enlarges, and is removed a little to one side. It is called the FASCICULUS or FUNICULUS GRACILIS — slender, and its enlarged upper end is known as the clava — a club (Fig. 48, page 66). When followed AND ITS MEMBRANES. 69 upwards it tapers to a point and becomes gradually lost, though we shall afterwards be able to trace its fibres along with those of the next fasciculus, first to collections of grey matter — nucleus gracilis and nucleus cuneatus — in the medulla, and thence to the cerebrum and cerebellum (see " Sensory Decus- sation," page 83). The outer division of the posterior column of the spinal cord, the postero-external strand — fasciculus of Burclach or cuneate fasciculus — passes into the medulla under this latter name. It lies between the paramedian and lateral dorsal sulci, and expand- ing above into a tubercle — the CUNEATE TUBERCLE (Figs. 48, 50) — reaches upwards beyond the clavn, and forms one of the lateral boundaries of the lower part of the 4th ventricle (Fig. 48, page 66). Besides the tracts just described there are on this aspect of the medulla two other strands, known respectively as the funiculus of Rolando and the RESTIFORM BODY. They are not represented on the surface of the posterior column of the spinal cord; for the former is chiefly grey matter, and the latter belongs mainly to the lateral column. (a) The funiculus of Rolando. — The first of these tracts, the funiculus of Rolando, is of a greyish colour, for there is little or no white matter on its surface. It lies outside the funiculus cuneatus, between it and the line of origin of the roots of the 9th, 10th, and 11th pairs of cranial nerves. Like the two tracts previously mentioned, it expands above into a tubercle — the tubercle of Rolando (Figs. 48, 50). 70 THE BRAIN The funiculus of Rolando is often very poorly de- veloped in the adult brain, but in the medulla of children it is always well marked. (6) The RESTIFORM BODY. — The remaining prom- inence, the largest and most conspicuous on this surface of the medulla, is called the INFERIOR CERE- BELLAR PEDUNCLE Or RESTIFORM BODY — Testis, a rope (Figs. 47, 48). Placed behind, and to the outer side of the lateral column of the cord, it lies above the level of the clava, the cuneate tubercle and the tubercle of Rolando, and, when looked at from the surface, seems to be incorporated with these fasciculi, though not so in reality; for, as we shall presently see, it is composed of fibres derived, for the most part, from the lateral columns of the spinal cord (Fig. 50, page 70). On this area of the medulla, then, we note four strands : (a) the funiculus gracilis and its clava ; (6) the funiculus cuneatus and its tubercle; (c) the funiculus of Rolando ; and (d) the restiform body. 2. The Lateral Area of the medulla, continuous with the lateral column of the cord, lies between the roots of the hypoglossal nerve in front and those of the 9th, 10th, and 11th nerves behind. It is wider below than above, where it is partially hidden from view by the oval eminence — the olivary body. To determine the origin of its various strands we must refer back to the constitution of the corresponding column of the spinal cord. You will recollect that, in that column, we traced five tracts — the crossed pyramidal, the (ascending and descending) antero- Fiu. 50. To cerebellum. Restiform body. -Mf Olivary body. Plate XVI. To motor areas. / / < Tubercle .— /---/-- -y""\ liculus of Rolando. \- - fasciculus of Goll. - mculus of Burdock. ... Mixed zone of lateral column kral cerebellar tract. \- - - prior median fissure Posterior columns. External rciform fibres. - Mixed zone. Crossed ill pyramidal tract. Direct pyramidal tract. Anterior median fissure. Lateral column. A nterior columns. AND ITS MEMBRANES. 71 lateral tracts, the direct lateral cerebellar tract, and the mixed zone. The crossed pyramidal tract can be followed to the anterior pyramid of the opposite side of the medulla, The direct lettered cerebellar tract, on passing up into the medulla, will be seen as a superficial band of fibres (Figs. 49, 50), running upwards and backwards across the line of origin of the 9th, 10th, and 11th pairs of nerve roots, over the funiculus of Rolando, above its tubercle and across the cuneate funiculus. Then, turning sharply upwards, it is joined by a set of fibres — the external arciform fibres (Fig. 50, page 70), which, together with it, form the main mass of the restiform body or inferior peduncle of the cerebellum (Figs. 48, 50). (For a full account of the constitution of the restiform body or inferior cerebellar peduncle, see page 99.) The mixed zone of the lateral column, when traced upwards to the bulb, is seen to dip under the olivary body, so that only a small part of it is visible on the surface of the medulla as a narrow white strand, between the olivary body and the roots of the 9th. 10th, and 11th pairs of nerves. Most of the fibres of this strand go to form part of a network of fibres that will become familiar to us under the name formatio reticularis. Some of them are, however, commissural between different segments of the cord. Of the antero-lateral ascending and descending tracts, the former is probably a crossed sensory tract from the posterior roots, and passes most likely to 72 THE BRAIN the cerebellum ; the latter, the descending tract, starts in the cerebellum, being part of the indirect or secondary motor tract. Thus, then, the fibres of the lateral column of the spinal cord, on reaching the medulla, are disposed of in three ways (see Fig. 50): (1) Some — crossed pyramidal tract — go to the opposite anterior pyramid ; (2) others — direct lateral cerebellar tract — to the restiform body, and thence to the cerebellum of the same side ; (3) others — antero-lateral tracts — to or from the cerebellum ; (4) others — the mixed zone — pass partly behind, and partly beneath the olive to the formatio reticularis. The constitution and ulti- mate destination of these strands of fibres will be seen hereafter. They are figured on Plates XL., XLL, XLIL, pages 182, 184, and 186. Olivary Body. — The olivary body is an oval prominence, about half-an-inch long, placed at the upper end of the lateral area of the medulla. It is bounded in front by the roots of the 12th nerve, but is separated behind from those of the 9th, 10th and 11th nerves by a narrow white tract, which, as we have already seen, is part of the mixed zone of the lateral column of the cord. Above, the olive almost touches the lower border of the pons; below, it is crossed transversely by delicate fibres— the external arciform fibres — already referred to. If you cut into it you will see that it contains a grey core called the inferior olivary nucleus (see page 81). 3. The Anterior Area, also called the anterior pyramid of the medulla (Fig. 46, page 66), lies AND ITS MEMBRANES. 73 between the anterior median fissure and the roots of the hypoglossal nerve, which serve to separate it from the olivary body. It is a pear-shaped prominence, which is broader above than below, though it be- comes slightly constricted before disappearing beneath the transverse fibres of the pons. The pyramids of the two sides will afterwards be traced through the pons to the cerebral peduncles, and thence to the cerebral cortex. To their constitution you will re- quire to give your closest attention, for it is some- what complicated and of the greatest importance (Fig. 50, page 70). Note that although we are tracing these pyramidal tracts upwards, they are in reality descending or motor strands. The anterior column of the spinal cord, you will remember, had a median division, direct pyramidal tract, and a lateral division, basis bundle or anterior root zone. Now, the pyramids of the medulla are, to a small extent only, made up of fibres derived from this inner division — direct pyramidal tract — of their own side of the cord; by far their greater part is composed of fibres — crossed pyramidal tract — which come from the opposite lateral column of the spinal cord. This crossed tract, after leaving the opposite lateral column, passes upwards and inwards through the anterior commissure, across the bottom of the anterior median fissure, where it decussates in a series of bundles, with a similar set of fibres from the other side, constituting together the decussation of the 'pyramids. Turning upwards this crossed tract then forms the inner and by far the larger part of the opposite anterior pyramid (Fig. 50); the outer 74 THE BRAIN and smaller part of the same pyramid being formed by the continuation upwards of the direct pyramidal tract of the same side. Thus, for example, the LEFT anterior pyramid of the medulla is chiefly made up of the crossed pyramidal tract of the right lateral COLUMN, and, to a much smaller extent only, by the direct pyramidal tract of its own side. We must note, however, that although the fibres of the direct pyramidal tract do not decussate in the medulla, they do decussate as they pass down the spinal cord, and ultimately, through the nerve cells of the anterior born, become connected, like the crossed tracts, with the anterior nerve roots of the opposite side to that at which they left the brain. The pyramidal tracts are the chief motor tracts, and their decussation explains the phenomena of certain forms of paralysis, in which when one side of the brain, say the left, is injured, loss of motion ensues, not on that side but on the opposite side of the body. " In cases, then, in which we find total or partial paralysis of the muscles on one side, with increased excitability to mechanical stimuli and a tendency to contracture, we can justly conclude that the opposite pyramidal tracts are affected " (Edinger). The outer division of the anterior column of the spinal cord — antero-extemal tract — will afterwards be traced through the upper part of the medulla (see "White Matter," floor of 4th ventricle, page 113) to the following destinations — (1) to the formatio reticularis, (2) to the posterior longitudinal bundle, and (3) to the tract of the fillet. AND ITS MEMBRANES. 75 External arcifoem fibres (Fig. 49, page 66). — These are a set of fibres which, as at present seen, emerge from the anterior median fissure of the medulla, cross over the surface of the anterior pyramids, over the lower part of and below the olives, and finally turn upwards, along with the lateral cerebellar tract, to form, with it, part of the restiform body. They will be again referred to in treating of the grey matter of the medulla. Table of Objects seen on Surface of Medulla. Fissures. -> Areas. t Anterior median, with decussation of the pyramids. | Posterior median. I Two lateral, with nerve roots — the 12th from antero- *■ lateral ; 9th, 10th, 11th from postero-lateral groove. f Anterior pyramids. \ Decussation of pyramids. ( 1. Anterior area, 2. Lateral area. ( Olive. ( Lower part. Posterior area. >per f art. \ Upper par \ External arciform fibres. (_ Tract of white matter. Restiform body. Funiculus of Eolando and its tubercle. Funiculus cuneatus and its tubercle. Funiculus gracilis and its clava. Lower half of the floor of 4th ventricle. Summary. — Thus, then, we see that the surface of the medulla oblongata presents four fissures — an anterior, two lateral, and a posterior: three areas — an anterior, with its pyramids and their decussation ; a lateral, with its olive ; a posterior, with its funiculus 7 6 THE BRAIN gracilis, funiculus cuneatus, and funiculus of Rolando, and their respective enlargements; and, finally, the restiform body, or inferior cerebellar peduncle. Table of Relations between White Tracts of Cord and their representatives in the medulla. Anterior column. Lateral column. Posterior column. Cord. 1. Direct pyramidal tract 2. Mixed zone 1. Crossed pyramidal tract 2. Direct lateral cere- bellar tract 3. Mixed zone 4. A ntero- lateral ascending tract 5. Antero-lateral de- scending tract [ 1. Postero-internal strand (Goll) 2. Posteroexternal strand (Burdach) 3. Lissauer's tract Medulla. to outer part of the an- terior pyramid of the medulla of the same side. to posterior longitudinal bundle. to olivary peduncle and fillet. to formatio reticularis. to the inner part of the pyramid of opposite side; a few fibres go to the pyramid of the same side. to restiform body and thence to the cere- bellum of same side. to olivary fillet. to formatio reticularis. to the cerebellum — prob- ably crossed sensory tract. from the cerebellum of same side. to funiculus gracilis, and to its nucleus. to funiculus cuneatus, and to its nucleus. to formatio reticularis. to the grey matter of the posterior horn. AND ITS MEMBRANES. 77 3. GREY MATTER OF THE MEDULLA. From the white strands of the medulla, which have so far occupied our attention, we next turn to the consideration of the arrangement of its grey substance. This is far more irregular than that of the spinal cord, and for its satisfactory study you will require a special series of preparations, though tlie accompanying figures will no doubt give you material assistance (Fig. 51, &c, Plate XVII., page 78). The grey matter of the medulla oblongata may be described under two heads — (1) that represented in the spinal cord, and derived from one or other of its grey crescents ; (2) that not so represented, but forming isolated collections or nuclei not obviously connected with the grey matter of the spinal cord. 1. Grey Matter derived from the Grey Cres- cents. — Transverse sections of the lower part of the medulla will show you that the grey matter of the medulla has an arrangement very similar to that of the grey matter of the spinal cord; but that higher up in the medulla the appearance of the grey crescents becomes much changed. (Compare Figs., Plate XVII, page 78, with Figs., Plate III., page 14.) (a) The anterior cornu of the spinal cord, as we trace it upwards, loses its characteristic shape, for, owing to the fact that the fibres of the crossed pyramidal tract cut their way through the neck of the anterior horn, the head becomes separated from the base, and the neck is replaced by a reticulum of fibres, part of the formatio reticularis. The head of the anterior horn, thus detached, enlarges, and, 78 THE BRAIN PLATE XVII. Fia:. 53. Fig. 56. a. in. f. Anterior median fissure. a, m.f. Anterior median fissure. a. re. Arciform nucleus. a. n. Arciform nucleus. as. gloss. Ascending root of the glosso- a. 0. Accessory olive. pharyngeal nerve. c. r. Corpus restiforme. e. a. f. External arciform fibres. e. a. f. External arciform fibres. h. n. Hypoglossal nerve. ij. n. Glossopharyngeal nerve. n. c. Nucleus cuneatus. l.orf. Lemniscus or Fillet. n. g. Nucleus gracilis. n. a. Arciform nucleus. n. 1. Nucleus lateralis. n. am. Nucleus ambiguus. n. v. b. Nucleus of olivary body. n.f. t. Nucleus of fasciculus teres. p. m. f. Posterior median fissure. n. o. b. Nucleus of olivary body. p.t. Pyramidal tract. p. 1. b. Posterior longitudinal bundle. s. g. R. Gelatinous substance of Rolando. p. t. Pyramidal tract. 5th as. Ascending root of 5th nerve. v. Vagus. 5th as. Ascending root of the 5th nerve. Fig. 52. Fig. 55. a. m. f. Anterior median fissure. a. m. f. Anterior median fissure. f.c. Fasciculus cuneatus. a. n. Arciform nucleus. f.'J. Fasciculus gracilis. a. o. Accessory olive. h. n. Hypoglossal nerve. e. a. f. External arciform fibres. ft. c. Nucleus cuneatus. /• r. Formatio reticularis. n. g. Nucleus gracilis. /. 8. Fasciculus solitarius. v. m. f. Posterior median fissure. h. n. Hypoglossal nerve. p. t. Pyramidal tract. n. am. Nucleus ambignus. s. g. R. Substantia gelatinosa Rolando. ft. c. Nucleus cuneatus. s. p. d. Superior pyramidal decussation. n. f. t. Nucleus of fasciculus teres. 5th as. Ascending root of the 5th nerve. it. a. Nucleus gracilis. s. d. Sensory decussation. ft. h. Nucleus of the hypoglossal nerve ft. o. b. Nucleus of olivary body. p. t. Pyramidal tract. r. Raphe. v. Vagus nerve. v. re. Vagus nucleus. 4th vent. 4th ventricle. 5th as. Ascending root of the 5th nerve. Fig. 51 Fig. 54. a. c. Anterior cornu. a. m. f. a-, m. f. Anterior median fissure. a. n. d. p. Decussation of the pyramids. a. o. d. p. t. Direct pyramidal tract. •:■ '••• / f.B. Fasciculus of Burdach. f.c. f.G. Fasciculus of Goll. f.Q. m. z. Mixed zone. f. r. p. c. Posterior cornu. i. a./. p. m /■ Posterior median fissure. n. c. n. q. n. 1. ft. o. b. p. m. /. P . t. s. (j. R. t. R 5th as. Anterior median fissure. Arciform nucleus. Accessory olive. External arciform fibres. Fasciculus cuneatus. Fasciculus gracilis. Formatio reticularis. Internal arciform fibres. Nucleus cuneatus. Nucleus gracilis. Nucleus lateralis. Nucleus of olivary body. Posterior median fissure. Pyramidal tract. Gelatinous substance of Rolando. Tubercle of Rolando. Ascending root of the 5th nerve. a.m./. Plate XVII. Fig. 53. Pio. 56. '■<(.?'. *.g.R. 5th as. M^/^WL^.'^^ p.vKf. ' as. gloss, -n.y. R^gp^ / // ^v<-v ■■'■-■■Mm r iz.9"- ) f c.r. & a. it. j / n./.t. p.l.b. 5th{ n.ai p.m./, v.n. f.G. p.m./ p-m./. AND ITS MEMBRANES. 79 by the interposition of the anterior pyramids between it and the anterior median fissure, is pushed from the antero-lateral aspect of the cord to the lateral aspect of the medulla, where, in sections of the lower part of the medulla, it may be seen as a distinct grey collection, the nucleus lateralis (n. L, Fig. 54, page 78). Again, when the central canal of the cord opens out into the floor of the 4th ventricle, the grey matter which surrounds that canal is brought to the surface, and hence the base of the anterior horn appears on the ventricular floor as a narrow grey tract close to the median furrow (Fig. 57, page 82). It is known as the nucleus of the hypoglossal nerve ; and in sections of the medulla at this level the nerve may be seen as a band of fibres running from the nucleus towards the anterior surface of the medulla (n. h., Fig. 55, page 78). Close to this nucleus is another small collection of nerve cells — the nucleus of the fasciculus teres (n.f. t., Fig. 55, page 78). The greater part of the anterior horn — the neck — is replaced, as above said, by a reticulum of cells and fibres, part of these latter being the fibres of the crossed tract on their way to the opposite side of the medulla. This network constitutes the anterior part of the formatio reticularis or grey reticulum (Figs. 54, 55, /. v.). (b) The grey matter of the posterior horn, on reaching the medulla, also takes up a lateral position and increases in amount. The head of the horn, much enlarged, comes nearer to the surface, and appears at about the middle of the medulla as a 5c THE BRAIX well-marked grey nucleus, the tubercle of Rolando (t. JR.. Fig. 54). beneath the surface prominence of the same name. To its outer side will be seen a band of white fibres, the ASCENDING ROOT of the FIFTH CRANIAL NERVE {5th OS., Fig. 54). The grey matter of the base of this horn is also much increased in amount, and, between the median line and the grey nucleus of Rolando, grows out as two superficial aggregations — the inner one into the fasciculus gracilis to form the NUCLEUS of bbe FASCICULUS gracilis, the outer into the fasciculus cuneatus to form the nucleus cuneatus [Fig. 54). Each nucleus thus lies beneath the corresponding white tract on the surface, and causes the elevations called the clava and the cuneate tubercle. To these nuclei, as we have seen, can be traced most of the fibres of the posterior column of the spinal cord. The nuclei themselves may be followed as far as the pons. From them start a set of fibres (internal arciform fibres) which cross the middle line and go t . form the sensory decussation (see page 83). The neck of the posterior cornu, like that of the anterior, is replaced by a network of fibres, the white reticulum, which becomes continuous with - e srey reticulum, constituting together the FOR- MATio reticularis (Figs. 54 and 55), a network of longitudinal, oblique, and transverse fibres (arciform fibres), with nerve cells and neuroglia cells embedded amongst them. The fibres of this reticulum are chieflv commissural in nature, though there are rea- - for thinking that sensory impulses, and im- pulses wmich inhibit spinal reflexes, travel through AND ITS MEMBRANES. 81 it to the cerebrum (see "Sensory Decussation/' 8. d., Fig. 52, page 78). Some of its longitudinal fibres are derived from the mixed zone — anterior root zone — of the anterior column of the spinal cord. 2. Isolated Grey Masses or Nuclei of the Medulla. — The chief of these nuclei are — (1) The corpus dentatum or olivary nucleus. (2) The accessory olives. (3) The arciform nucleus. (a) The corpus dentatum or olivary nucleus (Fig. 55) is contained in the centre of the olivary body, which must be cut into in order to see it. It is covered superficially by the external arciform fibres (Fig. 55). This ganglion consists of "flask - shaped, multipolar nerve cells, and neuroglia cells arranged as a zig-zag lamina or crumpled sheet of grey matter, concave and open on its inner aspect. Through this opening, called the hilum, enters the olivary peduncle, a bundle of nerve fibres, which passes into the centre of the olivary nucleus, to be there distributed in different directions. The corpus dentatum or olivary ganglion is closely connected with a grey mass in the cerebellum — the corpus dentatum of the cerebellum — for any injury to this latter nucleus causes atrophy of the opposite olivary ganglion (Fig. 99, Plate XXXVII., page 168). (b) Two other isolated nuclei — the ACCESSORY olives (Fig. 55) — will be found, the one on the inner G 82 THE BRAIN side of, the other behind, the corpus dentatum. They are linear in shape, and are closely connected with the olivary nucleus. (c) The third nucleus — arciform nucleus ; nucleus of the external arciform fibres (Fig. 54) — is placed amongst these fibres as they cross over the anterior pyramids of the medulla. The nucleus itself is continuous above with the nucleus pontis (See " Pons "). The remainder of the grey matter — the nuclei of the several cranial nerves — will be studied in re- lation to the floor of the fourth ventricle (page 111). Table of Grey Matter of the Medulla. Cord. Grey Matter of the Spinal Cord. Anterior Cornu. Posterior Cornu. Medulla, Head — Nucleus lateralis. Neck— Anterior part of the formatio reticularis. , Base — Nucleus of the hypoglossal nerve. ^ Head— Nucleus of Rolando. Neck — Posterior part of the formatio reticularis. < Base- J Nucleus gracilis. , Nucleus cuneatus. I Nuclei on floor of 4th ventricle. T Nucleus of the olivary body. (^ Nucleus of the external arciform fibres. Isolated grey nuclei _, Accessory olivary nuclei in the Medulla. Raphe. — A transverse section of the medulla will show you that above the level of the pyramidal decussa- tionHheJmedulla is partially divided into two lateral segments by a central median raphe* or partition, which forms a thin membranous septum of nerve substance, extending' from the anterior median fissure to beneath Fig. 57. pora Qnadrigeinina ~ 3rd. Plate XVIII. Descending root of 5th nerve. Locus cxruleus. ■'.fth nerve. - St/i nerve Nucleus of 6th nerve. Facial nerve. Auditory nerve. - - - hiferiov fovea. --Vagus nerve Ascending root of 5th nerve. Ascending root of glnxsop- pharyngeal Nuclei in 4th Ventricle fcending root of 5th. nerve. 12th. 10th. -9th Fig. 58. Uth. Olivary body A ud.itory. $ tfl - Nuclei of the cranial nerves. AND ITS MEMBRANES. 83 the central groove on the floor of the 4th ventricle. It consists of numerous fibres running in various directions, and interspersed with small collections of multipolar nerve cells. Sensory Decussation. — The superior pyramidal decussation, which has so often been referred to, must not be confused with the inferior or motor decussation (page 73). You will remember that fibres can be traced from the posterior spinal nerve roots through the posterior columns of the spinal cord to their primary stations, the nucleus gracilis and cuneatus of the medulla. From the cells of these nuclei, fibres travel towards the anterior aspect of the medulla until they come to lie in front of the central canal, and there decussating in the raphe, above and behind the pyramidal decussation, cross to the opposite side of the bulb. Becoming longitudinal, they form a considerable bundle of fibres just behind the pyramids (motor fibres), and ascend to the cerebrum in what is known as the tract of the fillet, or lemniscus. They are crossed sensory tracts to the brain (Fig. 96, Plate XXXVIL, page 168, fillet). Thus we see that we have in the medulla two sets of decussating fibres : — (1) The inferior pyramidal or motor decussation ; and (2), behind and above this, the superior pyramidal or sensory decussation. The Arciform Fibres have already been several times alluded to, so that w T e shall now merely collect together, for the sake of clearness, the several state- ments previously made. These fibres are divided 84 THE BRAIN into a superficial or external set, and a deep or internal set. 1. The SUPERFICIAL ARCIFORM FIBRES form two groups : — (a) The anterior external arciform fibres, which spring, you will remember, from the anterior median fissure, through which they can be traced to the central raphe, where they probably cross over to the posterior column of the opposite side of the medulla, to the nucleus gracilis and cuneatus. Emerging from the fissure, they wind round the anterior pyramids below, and over the outer surface of the olives, across the line of origin of the 9th, 10th, and 11th pairs of nerves. Turning upwards, they then blend with the fibres of the direct lateral cerebellar tract, and with them form the chief part of the restiform body (Fig. 50, page 70). (6) The other group — the posterior external arci- form fibres — passes from the posterior column of one side to the restiform body of the same side (Fig. 99, Plate XXXVII., page 168). 2. The DEEP arciform FIBRES are a delicate net- work of fibres found between the olives and behind the pyramids. The exact course and origin of these fibres are not yet known with certainty. Many of them are derived from the nuclei of the pos- terior columns before mentioned (superior pyramidal decussation). Others enter the centre of the olivary nucleus through its hilum, and either join the cells of that nucleus or pass through the nucleus to the restiform body directly, or first make for the AND ITS MEMBRANES. 85 surface, and then, along with the external arcifonn ribres, go to the restiform body. Recapitulation. — Since the constitution of the medulla is so complicated, it will be well to sum- marise the above facts in a somewhat different order. Commencing at the anterior median fissure, we meet on each side of that fissure — 1. The ANTERIOR PYRAMIDS OF THE MEDULLA, composed of longitudinal fibres, derived chiefly from the crossed pyramidal tracts of the opposite lateral columns, and to a small extent from the direct pyramidal tracts of the same side. Most of the motor fibres coming from the brain are contained in these pyramids, though others, as we shall afterwards see, first go to the pons, thence to the cerebellum, thence to spinal cord (secondary motor tract). 2. Superficial to the anterior pyramids are a set of fibres, which, having their origin in the nucleus gracilis and nucleus cuneatus of the one side of the medulla, and having decussated in the middle line, emerge from the anterior median fissure, cross over the surface of the anterior pyramids and the olives of the opposite side to that from which they started, and go to join the restiform body. They are the superficial arcifonn fibres (Figs. 55, 56,e.«./.), and amongst them is a mass of grey matter, the ARCIFORM NUCLEUS. 3. In the middle line behind the pyramids, between them and the central canal, are thick white bundles of fibres arranged in concentric curves. They spring from the region of the posterior columns, 86 THE BRAIN nucleus gracilis and cuneatus, and form the SUPERIOR PYRAMIDAL DECUSSATION, or decussation of the fillet. Higher up in the medulla this same region is occupied by fibres which traverse the medulla in all directions — formatio reticularis. Neuroglia cells, and a few nerve cells are scattered amongst the fibres, many of which are known as the deep arciform fibres. 4. Outside the formatio reticularis, between it and the surface, but behind the pyramids, is the olivary nucleus (n.o.K, Fig. 56) with the olivary PEDUNCLES. This nucleus is covered superficially by the external arciform fibres (Fig. 56). Close to it are the accessory olives (Figs. 55, 56, a.o., page 78). 5. Behind the olives you will see the nucleus LATERALIS (n.L, Fig. 54), the upward continuation of the anterior cornu of the spinal cord. 6. Posterior to the nucleus lateralis appears the grey tubercle of Rolando (Figs. 53, 54, t.R.) — the enlarged head of the posterior horn of the spinal cord. On its outer side will be seen the ascending root of the 5th nerve (5th as., Fig. 54). Superficial to the tubercle of Rolando are the fibres of the DIRECT LATERAL CEREBELLAR TRACT, which are on their way, along with the arciform fibres, to the inferior cerebellar peduncles or restiform body. 7. Posterior and internal to the nucleus of Rolando is a mass of grey matter— the NUCLEUS CUNEATUS (n.c, Fig. 54) — lying beneath the cuneate tubercle ; and still nearer the middle line is another grey collection, the nucleus of the funiculus gracilis (Figs. 54, 55, n.g.). AND ITS MEMBRANES. 87 In front of the nucleus cuneatus will be seen a special white rounded fasciculus, known as the FASCICULUS SOLITARIUS {f.s., Fig. 55), the ascending root of the glosso-pharyngeal nerve. Close to the middle line, internal to the nucleus cuneatus, lies the nucleus of the hypoglossal nerve, from which the nerve itself may be seen running forwards to its superficial origin. The rest of the grey matter, internal to the nucleus of the funiculus gracilis, and on each side of the posterior median groove, belongs to the floor of the 4th ventricle (Fig. 57, page 82), and will be fully described with that cavity. The next division of the brain to be considered is the metencephalon, comprising the pons and cerebellum. II.— PONS VAROLII. General Outline. — The Pons Varolii — part of the metencephalon, the other part being the cerebel- lum — is a broad white band which crosses transversely between the two halves of the cerebellum above the level of the anterior aspect of the medulla oblongata. It rests on the body of the sphenoid. Composed of grey and white matter, it presents an upper and a lower border — an anterior and a posterior surface (Fig. 46, page 66, and Figs. 59, 60, Plate XIX., page 90). The UPPER BORDER is arched, being higher in the middle than at the sides, and from it spring the crura cerebri, or cerebral peduncles. Two cranial nerves, the 3rd and 4th, are seen at this border, the former being near the middle line. 88 THE BRAIN The LOWER BORDER, which marks the upper limit of the medulla in front, is horizontal, and is almost in contact with the upper margin of the pyramids and the olives (Figs. 46, 49, page 66). At this border are seen, from within out, the 6th, 7th, and 8th cranial nerves. The anterior SURFACE of the pons is convex, and rests on the dorsum sellse of the sphenoid bone. Along the middle line it presents a shallow groove, which lodges the basilar artery. On each side of this groove is a longitudinal elevation caused by the continuation upwards of the pyramids of the medulla. At the sides, the pons becomes narrowed, and passes as two rounded bundles, one on each side, into the cerebellum, forming its "middle 'peduncles. From the lateral part of this aspect of the pons springs the 5th cranial nerve. The posterior surface of the pons has ill-defined limits, being continuous below with the medulla, above with the cerebral peduncles. Flattened from before backwards it forms the upper part of the floor of the 4th ventricle, and will be described with that cavity (page 106). When we examine transverse sections of the pons, we find that it can be divided into two portions, a ventral portion and a dorsal portion. The latter, called the tegmentum, is a continuation upwards of the constituents of the spinal cord and medulla, with the exception of the pyramidal tracts. The former, the ventral part, contains, besides the proper fibres of the pons, the upward continuation of the pyramids of the medulla. AND ITS MEMBRANES. 89 It is interesting to note that in mammalia the size of the pons bears a direct relation to the size of the lateral lobes of the cerebellum; and that in birds, reptiles, and fishes, where we have no cerebellar hemispheres, the pons also is wanting. We shall consider the constitution of the Pons according to the folio wins: table: — Table of the Structure of the Pons Varolii. I. WHITE MATTER OF THE PONS. 1. Transverse fibres. (1) Superficial. (2) Deep. 2. Longitudinal fibres. (1) Superficial. Anterior pyramids. (2) Deep. (a) Lemniscus, or fillet. (6) Posterior longitudinal bundle. (c) Fasciculus teres. 3. Fibres of the 5th, 6th, 7th and Sth cranial nerves. 4. Formatio reticularis. 5. Raphe. II. GREY MATTER OF THE PONS. 1. Nucleus pontis. 2. Superior olive. 3. Nuclei of origin of 5th, 6th, 7th and 8th cranial nerves. 4. Locus caeruleus. 1. WHITE MATTER OF THE PONS. Dissection. — To see the arrangement of the white fibres of the pons, you will require to dissect down on each side of the middle line, through the superficial transverse fibres, until you reach a longitudinal set passing upwards from the medulla. Reflect the longitudinal fibres, when a still deeper transverse group will come into view. 9 o THE BRAIN The white or medullated nerve fibres of the pons are arranged in two sets, a transverse and a longitudinal, each being again divisible into a superficial and a deep group. 1. TransYerse fibres. — (1) The superficial transverse fibres appear on the surface of the pons, and (2) the deep transverse fibres lie behind the superficial longitudinal ones (Figs. 59, 60, page 90). At the lower part of the pons, near the medulla, the deep set of transverse fibres forms a special collection, called, from its peculiar arrangement, the trapezium (Fig. 59). Traced laterally, all the transverse fibres of the pons pass into the middle peduncle of the cerebellum. Some of these fibres are commissural between the two halves of the cerebellum; others are connected with the nucleus pontis of the same side or of the opposite side. 2. Longitudinal fibres. — (1) The superficial LONGITUDINAL fibres (Figs. 59, 60) are mostly the upward continuation of the anterior pyramids of the medulla, and in transverse sections are seen as two rounded bundles behind the superficial trans- verse fibres, though many of them are intersected by these latter. (2) The deep longitudinal bundles (Figs. 59, 60) are placed near the dorsal aspect of the pons, which is chiefly made up of the formatio retic- ularis, and of a prolongation of the grey matter of the medulla. Three distinct sets of longitudinal fibres can, however, be recognised — viz., (a) the lemniscus, or fillet, the continuation of the sensory decussation ; (b) the posterior longitudinal bundle, Fig. 59. Plate XIX. . Nucleus of Nucleus of Pars intermedia. '■■ ''" • <-*^L 8 ffinerv *- 6th, /nerve. ..--' v s--- 7