PART 1.

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‘pigment of the skin€’

TRANSLATIONS

OF

FOREIGN BIOLOGICAL MEMOIRS

II

London

HENRY FROWDE

[Illustration]

Oxford University Press Warehouse

Amen Corner, E.C.

THE

ANATOMY OF THE FROG

BY

Dr. ALEXANDER ECKER

PROFESSOR OF HUMAN AND COMPARATIVE ANATOMY

IN THE UNIVERSITY OF FREIBURG

Translated, with numerous Annotations and Additions

BY

GEORGE HASLAM, M.D.

SCIENTIFIC ASSISTANT IN THE MEDICAL DEPARTMENT IN THE UNIVERSITY

OF ZÜRICH; FORMERLY ASSISTANT LECTURER IN PHYSIOLOGY IN THE

OWENS COLLEGE, VICTORIA UNIVERSITY, MANCHESTER

_ILLUSTRATED WITH MANY WOOD ENGRAVINGS_

_AND TWO COLOURED PLATES EXECUTED BY HOFMANN, WÜRTZBURG, BAVARIA_

Oxford

AT THE CLARENDON PRESS

1889

[_All rights reserved_]

TRANSLATOR’S PREFACE.

I undertook the publication of a translation of Ecker’s ‘Anatomie des

Frosches’ at the suggestion of Professor A. Gamgee while I was working

under his superintendence in the physiological laboratory of the Owens

College. The work was subsequently accepted by the Delegates of the

Clarendon Press, as one of the series of ‘Foreign Biological Memoirs,’

published by them. Early in the progress of the work it became evident

that a mere translation would be unsatisfactory, and that it would

be desirable to recast and modify several portions of the book. It

was deemed advisable to give greater completeness to the work by

descriptions of the minute structure of the several organs. For these

purposes the appearance of the work has been unavoidably delayed.

I have done my best to bring the book up to date by including the

results of recent researches, to which I have added many facts derived

from my own personal investigations. All such additions are enclosed

within square brackets [ ]. More than a hundred new figures, of which

one-third are original, have been added; and copious, though it is

feared still incomplete, lists of references to frog-literature have

been drawn up. By these additions the size of the book has been

considerably increased.

In the several sections into which the book is divided the following

points may be more particularly noticed:--

Sect. I. The Bones and Joints. The nomenclature of Parker and Bettany

has been adopted throughout.

Sect. II. The Muscles. This section remains in its original form.

Sect. III. The Nervous System. The chapters on the central

nervous system and the sympathetic system have been rewritten.

The description of the arteries of the brain is entirely new,

while the chapters on cranial and spinal nerves have received

many smaller additions, and have been rearranged to facilitate

reference.

Sect. IV. The Vascular System. The chapter on the heart is

practically new, and many additions and alterations have been made

in the descriptions of the blood-vessels and lymphatics.

Sect. V. The Alimentary Canal, with its appendages, the Spleen and

the Peritoneum. In this section much new material has been added:

the descriptions of the blood-vessels of the liver, the ducts

of the liver and pancreas, etc., being the results of original

investigation.

Sect. VI. The Respiratory Organs, the Thymus and Thyroid Glands.

These organs have been carefully studied and numerous new details

are noted. The lymphatic glands of the hyoid region have, after

some hesitation, been designated tonsils.

Sect. VII. The Urino-Genital Organs. A very large number of

preparations have been made to investigate the vessels and

uriniferous tubes of the kidneys; and the descriptions of the

remaining organs of this section have received large additions

from recent publications.

Sect. VIII. The Skin and the Sense-Organs. This section has, with the

exception of very small portions, been re-written and very much

enlarged.

Before concluding this preface, I must thank my friend Professor A.

Milnes Marshall, of the Owens College, for all the help and kindness

he has extended to me before and during the time this work has been

in hand; to him I am indebted not alone for the loan of books,

pamphlets, etc., and for much useful information, but also for the care

and patience with which he has read and corrected the whole of the

proof-sheets.

To Professor G. Lunge, of Zürich, I am indebted for the use of the

library of the Gesellschaft der Naturforscher of Zürich; and to my

friend Mr. C. Herbert Hurst, of the Owens College, for the drawings

for figures 132, 133, 134, and 136; also to Dr. Max Köppen, of

Strasburg, for the proof-sheets of his valuable paper, ‘Zur Anatomie

des Froschgehirns’: to these gentlemen I beg to express herewith my

heartiest thanks. Lastly, I must express my sense of indebtedness to

the Delegates of the Clarendon Press, who have kindly allowed me to

alter the original plan of the book, and to make extensive additions

far beyond the limits originally intended.

A second edition of the original German work is in course of

publication. The first part, on the bones and muscles, has already

appeared.

GEO. HASLAM.

Zürich, 1888.

PREFACE TO THE FIRST PART.

The idea of this manual on the anatomy of the frog, of which I now

offer the first part to physiologists and to those who would become

such, occurred to me during the preparation of the plates for my

‘Icones Physiologicae.’ I was then convinced of the necessity of such

a book. I regret that many direct and indirect causes have hindered

its earlier completion; fortunately, however, its appearance is still

opportune, as the need for the book has not diminished. I am conscious

that the book requires a recommendation to the indulgent judgment of

my fellow-workers, since almost every one has studied the frog for one

purpose or another, and each will closely criticize in that department

with which he is most familiar. Although I shall not be able to satisfy

all, still I hope that my work may serve as a useful basis for further

investigations, and I would apply to it the words with which Sömmering

prefaced his anatomy: ‘Ich wünschte ein Handbuch zu liefern und seine

Einrichtung so zu treffen, dass man künftig an ihm als einer Basis nach

Erforderniss leicht ändern, wegnehmen und zusetzen könnte[1].’

[Footnote 1: ‘I wished to furnish a manual so arranged that it might

serve as a basis easily altered, pruned, or enlarged as the future

might need.’]

Lest more be anticipated from the book than it is intended to supply,

I would observe that I have throughout had in mind only a descriptive

anatomy of the indigenous (German) frog; a comparative anatomy of

Batrachians was as foreign to my intention as were developmental

or histological questions: hence morphological details must not be

expected. Any hope of formulating a systematic nomenclature of the

muscles has been abandoned; as neither one based upon their mode of

action, of which we know so little, nor one based upon their origins

and insertions, as demonstrated by the unpronounceable names of

Chaussier and Dugès, is really practicable. I have therefore preferred

to avail myself, as far as possible, of the received names, which have

been chosen partly according to mode of action, partly according to

origin and insertion, and partly according to position and form; while

in the choice of new names I have given preference to the simplest.

The figures are, with few exceptions, original, and drawn by myself.

Their careful execution in woodcut has added a very necessary neatness

to that correctness, which alone I claim as mine.

ALEXANDER ECKER.

Freiburg,

_February, 1864_.

PREFACE TO THE SECOND PART.

Sixteen years have elapsed since the first portion of this anatomy of

the frog appeared; this second portion, therefore, requires a somewhat

apologetic introduction.

The nervous and vascular systems have, in substance, been known for

some years; still, certain points required a thorough revision: this

seemed especially necessary with regard to the cranial nerves. In

consequence of my anthropological investigations, and particularly

through undertaking the editorship of the ‘Archiv für Anthropologie,’

my attention was drawn into another channel, and I found it impossible

to work out this chapter: consequently the whole was deferred, and

would have been still longer delayed had I not received assistance.

At my request Professor Wiedersheim undertook to investigate afresh the

cranial nerves, the brain, the spinal cord, and the sympathetic system;

and the descriptions of these parts are the result of his work alone. I

regard it as most advantageous to this second part that so experienced

an investigator in the anatomy of Amphibia should have given me his

help.

The remaining portions appear almost unaltered as written several

years ago; and the majority of the illustrations date from the same

period. I had neither the time nor the zeal necessary to re-examine the

whole; besides, it is doubtful whether eyes some twenty years older

would improve matters.

This somewhat neglected book is therefore commended to the indulgence

of my fellow-workers, with the hope that it may at least form a basis

upon which further work may easily be done; to proffer more than this,

as I stated, with a quotation from Sömmering, in the preface to the

first part, I have never even hoped.

The final part of the work, on the viscera and sense-organs, has been

undertaken by Professor Wiedersheim, and will appear in the Spring of

1882.

ALEXANDER ECKER.

Freiburg,

_August, 1881_.

CONTENTS.

PAGE

INTRODUCTION 1

SECTION I. THE BONES AND JOINTS 11

" II. THE MUSCLES 53

" III. THE NERVOUS SYSTEM 121

" IV. THE VASCULAR SYSTEM 203

" V. THE ALIMENTARY TRACT WITH ITS APPENDAGES, THE

SPLEEN, AND THE PERITONEUM 267

" VI. THE LARYNX, LUNGS, VOCAL SACS, THYMUS AND THYROID

GLANDS, AND THE LYMPHATIC GLANDS (TONSILS?) OF

THE HYOID REGION 307

" VII. THE URINO-GENITAL SYSTEM, THE ADRENALS, AND THE

FAT-BODIES 325

" VIII. THE SKIN AND THE SENSE-ORGANS 351

ADDENDA, ETC. 425

INDEX 441

ILLUSTRATIONS.

FIGURE PAGE

1. The Green water-frog, _Rana esculenta_, L. 4

2. The Brown grass-frog, _Rana temporaria_, L. 8

3. Femur of _Rana esculenta_ 16

4. Vertebrae of do. 17

5. Vertebral column of do. 18

6, 7. Section through a vertebra of _Rana esculenta_ 20

8, 9. Urostyle of _Rana esculenta_ 21

10–14. Skull of do. 22, 23, 25, 28

15. Nasal Cartilages of frog 29

16, 17. Skull of _Rana esculenta_ 29, 30

18. Origin of suspensory cartilage from the skull 32

19, 20. Skull of _Rana esculenta_ 32, 33

21. Mandible of _Rana esculenta_ 34

22. Hyoid of _Rana esculenta_ 35

23. Omosternum of _Rana esculenta_ 36

24. Shoulder-girdle and sternum of do. 36

25. Shoulder-girdle of the frog 37

26, 27. Suprascapula of _Rana esculenta_ 38

28. Left scapula of _Rana esculenta_ 38

29. Scapula seen from behind 38

30. Left coracoid 39

31. Clavicle of the left side 39

32. Right shoulder-girdle of _Rana esculenta_ 40

33. Hinder border of the scapula and coracoid 40

34. Clavicular cartilage of _Rana esculenta_ 40

35. Humerus of _Rana esculenta_ (female) 41

36. Do. (male) 41

37. Do. (female) 41

38. Radio-ulnar of _Rana esculenta_ 43

39, 40. Bones of the forearm and hand of _Rana esculenta_ 44, 46

41, 42. Pelvis of _Rana esculenta_ 48

43. Horizontal section through the iliac bones, etc. 49

44. Femur of _Rana esculenta_ 49

45. Tibio-fibula of do. 50

46. Section of the tibio-fibula 50

47. Right foot of _Rana esculenta_ 51

48–50. Eye-muscles of do. 55, 56

51. Skull and orbital cavities of _Rana esculenta_ 57

52. M. levator bulbi of _Rana esculenta_ 57

53. Eye-muscles of _Rana esculenta_ 58

54. Facial muscles of do. 59

55. Muscles of the back and shoulder 60

56, 57. Muscles of the lower jaw of _Rana esculenta_ 61, 62

58. Muscles of the throat, chest, and abdomen of do. 63

59. Muscles of the hyoid bone and the tongue of do. 64

60. Do. (from below) 65

61. Do. (from right side) 66

62. Muscles of the throat, chest, and belly of _Rana esculenta_ 68

63. Muscles of trunk of _Rana esculenta_ (from the right side) 69

64. Second layer of abdominal muscles of _Rana esculenta_,

from right side and below 70

65. M. obliquus internus 71

66. Muscles of the back and shoulder blade 72

67. Muscles of the back and pelvic girdle of _Rana esculenta_ 74

68. Muscles of the shoulder, from below 77

69. Muscles of the right shoulder and upper arm 78

70. Right shoulder, from below 79

71. Muscles of the back and shoulder 79

72. Muscles of the chest, throat, and belly of _Rana esculenta_ 81

73. Muscle of right shoulder and upper arm 83

74. Muscles of the right arm of _Rana esculenta_ 84

75. Do. (deep layer) 85

76. Muscles of forearm of _Rana esculenta_ 86

77. Muscles of hand of _Rana esculenta_, volar surface 88

78. Second layer of muscles on volar surface of hand of

_Rana esculenta_ 88

79. Muscles of hand of _Rana esculenta_ 92

80. Muscles of left thigh of do. 95

81, 82. Do. (ventral surface) 98, 99

83. Deep muscles of left thigh of _Rana esculenta_ 100

84. Do. (Dorsal view) 101

85. Left half of pelvis of _Rana esculenta_ 101

86. Muscles of the right leg and foot of _Rana esculenta_

(Dorsal view) 103

87. Do. (seen from below) 105

88. Do. (Dorsal view) 106

89–91. Muscles of the plantar surface of foot of

_Rana esculenta_ 107, 111, 112

92–94. Dorsal view of muscles of foot of

_Rana esculenta_ 115, 117, 118

95. Pectoral region of _Rana esculenta_ 119

96. Hind portion of back and thigh of _Rana esculenta_ 120

97. The nervous system of _Rana esculenta_, from the

ventral surface 136

98. Dorsal view of brain of _Rana esculenta_ 143

99. Transverse section through hinder end of Medulla

oblongata 144

100. Do. at the point of origin of the abducens nerve 145

101. Do. of the auditory nerve 146

102. Ventral view of brain of _Rana esculenta_ 149

103. Lateral do. 150

104. Transverse section through the anterior portion of

the optic lobes opposite the origin of the

motor-oculi nerve. 151

105. Horizontal section through the brain to show the

ventricles 153

106. Section through the lower division of the pituitary body 157

107. Transverse section through the hinder portion of the

cerebral hemispheres 158

108. Transverse section near the middle of the cerebral

hemispheres 158

109. From a transverse section through one of the cerebral

hemispheres 159

110. Diagram to show the Vena spinalis posterior, etc. 164

111. Dorsal view of the orbit, etc. (deep dissection)

(_coloured_) Plate I.

112. Do. (superficial do.) (_coloured_) Plate I.

113. View of roof of mouth; mucous membrane, etc.

(_coloured_) Plate I.

114. Lateral dissection of head, etc. (_coloured_) Plate I.

115. Dissection of the floor of the mouth (_coloured_) Plate I.

116. Right half of skull of _Rana esculenta_ 174

117. The nervous system of _Rana esculenta_, from the

ventral surface 176

118. Ventral view of the brain and spinal cord, to show the

points of exit of the spinal nerves 178

119. Ventral view of the spinal ganglia 179

120. Schema of spinal ganglion 179

121. Dorsal branches of the spinal nerves 181

122. The brachial plexus 184

123, 124. Nerves of the ventral surface of the arm 185, 186

125. The N. radialis 186

126. Ventral view of the brain and spinal cord 188

127. The sciatic plexus 190

128. Distribution of the sciatic nerve 193

129. Nerves of the leg and sole of the foot 194

130. Distribution of the N. peroneus 196

131. Sympathetic cord 198

132. The heart and blood-vessels, seen from the ventral

surface 213

133. The heart, seen from above 214

134. The frog’s heart, seen from the ventral surface 215

135. Dissection of a case in which the auricular septum

is placed more to the left than is normal 215

136. Dissection of the heart from the left side 216

137 I. Transverse section through the junction of the hinder

and middle thirds of the ventricle of _R. temporaria_ 217

137 II. Transverse section through junction of the middle

and anterior thirds of the same heart 217

138. Portion of a transverse section through the middle

of the ventricle of _R. temporaria_ 218

139. Course of the cardiac nerves in the auricular septum 219

140. Group of nerve-cells on the cardiac nerve, from the

auricular septum 220

141_a_. Small group of nerve-cells from the auricular septum 221

141_b_. Isolated nerve-cells from frog’s heart 221

142. Arteries and veins of the Truncus arteriosus of _Bufo

vulgaris_ 222

143. Schema of the arterial system of _Rana esculenta_ 223

144. Right carotid gland 224

145. Arterial system of Rana esculenta 225

146. Transverse section at level of the larynx 226

147. Dissection to show the occipito-vertebral and the

cutaneous arteries 227

148. Branches of the occipito-vertebral and cutaneous

arteries in the head 228

149. Dissection to show the occipito-vertebral and the

cutaneous arteries 229

150. Subclavian artery of the left side 231

151. Arteries of the palmar surface of the hand 232

152. Arteries of the dorsal surface of the hand 232

153. Arterial system of _Rana esculenta_ 234

154. The urinogenital arteries 235

155. Bifurcation of the aorta and the iliac arteries 236

156. Arteries of the hinder extremity 237

157. Arteries of the dorsal surface of the foot 239

158. Arteries of the sole of the foot 240

159. Schema of the veins of _Rana esculenta_ 242

160. Distribution of the internal jugular vein and the

anterior portion of the cutaneous vein 243

161. The anterior caval vein and its branches 244

162. Course of the cutaneous vein as seen from the side 245

163. Veins in the region of the kidney 246

164. Veins of the liver 248

165. Veins of the hinder extremity 250

166. Transverse section of a septum with the attached skin 252

167. The sinus abdominalis lateralis 252

168. Sinus thoracicus transversus 253

169. The lymph-sacs of _Rana esculenta_ (seen from the

dorsal surface) 254

170. Do. (seen from the ventral do.) 256

171. Do. (seen from the side) 257

172. Transverse section through the trunk in the region

of the iliac lymph-sac 258

173. Dissection to show the iliac lymph-sac 259

174. Plan of attachments of the inferior femoral etc. septa 259

175. Transverse section of the thigh 260

176. The anterior lymph-hearts 261

177. The posterior lymph-hearts 261

178. The roof of the mouth 276

179. The floor of the mouth 277

180. The capillaries of the submucous layer (_coloured_) Plate II.

181 I. Transverse section of the premaxillary bone, to

show attachment of the teeth 279

181 II. Dentine and enamel 279

181 III. Enamel 279

182, 183. Muscles of the tongue 281, 282

184. The alimentary canal 283

185. The abdominal viscera of _Rana esculenta_ 284

186. Longitudinal folds of stomach of _Rana temporaria_ 285

187. The cells at the mouth of the gland of the fundus

of the stomach. (_coloured_) Plate II.

188. The mucous membrane of the pyloric end of the stomach

of _Rana esculenta_ 286

189. Mucous membrane of the pyloric end of stomach and

duodenum 288

190. Isolated fold of mucous membrane of small intestine of

_Rana temporaria_ 291

191. Fold of mucous membrane of _Rana temporaria_ 291

192. The large intestine of _Rana temporaria_ 292

193. Large intestine of _Rana esculenta_ 293

194. The liver 295

195. The pancreas and bile-canals 296

196 I. }

} The hepatic veins (_coloured_) Plate II.

196 II. }

196 III. The hepatic arteries (_coloured_) Plate II.

197. Liver-cells 299

198. The bile-capillaries 299

199. The pancreas and bile-canals 300

201. The peritoneum of _Rana esculenta_ 305

202. The position and relations of the larynx 311

203. The cartilaginous skeleton of the larynx 312

204. The larynx and surrounding parts 313

205. The muscles of the larynx 314

206. Three sections through the larynx of _Rana esculenta_ 316

207. The Rima glottidis 317

208. The lung of _Rana temporaria_ (_coloured_) Plate II.

209. The vocal sac of the right side 320

210. The thymus gland 321

211. The thymus gland of _Rana esculenta_ 321

212. The thyroid gland of _Rana esculenta_ 323

213. The lymphatic gland of _Rana esculenta_ 324

214. The male urino-genital organs 331

215. The right kidney 332

216. The blood vessels and lymphatics of the kidney 333

217. Vertical sections through the kidney (_coloured_) Plate II.

218. The uriniferous tubes 335

219. A gold preparation of the kidney of _Rana esculenta_

(_coloured_) Plate II.

220. Transverse section of the kidney 338

221. The bladder 339

222. The male reproductive organs 341

223. Various preparations from the testis 342

224. The female reproductive organs 344

225. Preparations from ovary and oviduct 346

226. The male urino-genital organs 348

227. The fat-body of _Rana esculenta_ 349

228. The epidermis from the head of _Rana esculenta_ 367

229. Vertical section through the skin of the back 368

230. Surface view of epidermis of _Rana temporaria_ 368

231. Nerve terminations of the branched pigment-cells of

the cutis 368

232. The temporary papillae in _Rana temporaria_ 371

233. The epidermis of the supplemental toe of

_Rana esculenta_ 375

234 I. Fore-foot of a male frog 375

234 II. The swelling on the supplemental toe of a male frog 375

235. The blood-vessels and lymphatics of the skin 376

236. Lateral sense-organ of tadpole of frog 378

237. Various parts from the fungiform papillae 381

238, 239. Frontal sections through the nose of two tadpoles 384

240 I A. Bowman’s glands _in situ_ from _Rana temporaria_ 386

240 I B. Section of Bowman’s gland 386

240 II. Vessels of nasal mucous membrane of _Rana esculenta_ 386

241. Separations from the olfactory mucous membrane of _Rana

temporaria_ 388

242. The tympanic membrane of _Rana esculenta_ 389

243. The columella 391

244. Antero-posterior section through the capsule of the

right labyrinth of _Rana esculenta_ 392

245. The membranous labyrinth of _Rana esculenta_ 394

246. Part of the outer wall of the perilymphatic space 395

247, 248. The right membranous labyrinth of _Rana esculenta_ 397

249, 250. The membranous labyrinth of _Rana esculenta_ 399, 401

251. Preparations from the ear of _Rana esculenta_ 402

252. The nerve-terminations in the membranous labyrinth of

_Rana esculenta_ 404

253. Endothelium from the inner surface of the sclerotic coat 406

254, 255. Preparation from cornea of _Rana esculenta_ 407, 408

256. The vessels of the choroid and iris 410

257. Fibres from the lens of the frog 414

258. Vertical section through retina of frog 415

259. Various preparations from the eye of the frog 417

260. The vessels of the vitreous body 421

261. Preparations from the nictitating membrane of _Rana

esculenta_ 423

INTRODUCTION.

There is no occasion, now-a-days, to offer a lengthened apology for

devoting a treatise solely to the anatomy of the frog, which enjoys

the doubtful honour of being, κατ' εξοχήν, the physiological domestic

animal. It is kept in every physiological laboratory, and is daily

sacrificed in numbers upon the altar of science. The physiologist has

recourse to it, not only to obtain answers to new questions, but for

the sake of demonstrating easily and quickly the most important known

facts of the science. These unlucky batrachians are to be had in any

number, and are specially adapted for experimental investigation: they

have consequently fallen under a harsher tyrant than the stork in the

fable, and their prophetic outcry in the frog-chorus of Aristophanes,

δεινὰ πεισόμεσbθα, has been literally fulfilled.

As the history of the most important physiological discoveries is

closely related with the employment of the frog in physiological

research, it will not be without interest to review briefly the history

of its use in scientific, especially in physiological, investigations,

and to record the services which it has already rendered to science.

Swammerdam (1637–1685), as du Bois-Reymond justly remarks, was the

first to make known the frog as an important means of research; he

says concerning it:--‘An den Thieren, die das heisseste Blut haben,

ist die Bewegung der Muskeln nicht so merklich oder hält vielmehr

nicht so lange an, als an Thieren die mit kälterem Blute begabt

sind. Dergleichen sind die Fische und viele andere Wasserthiere, wie

auch solche, die so wohl im Wasser als auf dem trocknen Lande leben

können. Deswegen habe ich insonderheit mit dem Frosch meine Versuche

angestellt. Denn an diesem Thiere sind die Sehnen sehr sichtbar und

lassen sich leicht entdecken und entblössen[2].’ Swammerdam made the

earliest experiments on the contraction of muscle by means of chemical

and mechanical stimulation of its nerves; thus laying the basis of our

present nerve and muscle physiology, which has been built up within

rather less than two hundred years; though during the first half of

this period but little advance was made.

[Footnote 2: ‘In animals with warm blood the action of the muscles is

neither so apparent nor so long continued as in those animals which

have cold blood, such as fishes and many other aquatic animals, and

those also which live both in water and on dry land. On this account I

have made my investigations chiefly on frogs, for in them the nerves

are very distinct, and are easily found and exposed.’ Buch der Natur,

Leipzig, 1752, p. 330.]

From the famous September evening of the year 1786, on which Galvani

first observed the twitchings of a frog’s leg suspended by a metallic

hook to an iron balcony, the frog has, down to the present time,

afforded almost the only material for the investigation of the

excitability of nerve and its associated electromotive changes,

and also no inconsiderable part of the remaining nerve and muscle

physiology. It was not until Müller devised the method of operating

on the frog that Bell’s law became capable of easy proof; and much

of our knowledge of the functions of the spinal cord is derived from

experiment upon it. Again, the muscles of frogs served, from the time

of Swammerdam to that of Eduard Weber and his followers, for the

investigation of the phenomena and the conditions of contraction;

and in almost all other branches of physiology there are important

doctrines which were first definitely established by experiment upon

the frog. But for the web of the foot of this animal (and the gills and

tail of its tadpole, in which Leeuwenhoek[3] describes the phenomena

most clearly) we should not, perhaps for a long time, have arrived

at a satisfactory knowledge of the existence and the conditions of

the capillary circulation. As is well known, an accurate acquaintance

with the constituents of the blood directly concerned in nutrition

has been obtained by observation on the frog, as well as important

facts in the physiology of the blood and lymph, such as the intimate

knowledge of the corpuscles of both fluids, and the coagulability

of the plasma; while in no less degree have experiments on these

animals served to establish the laws of the heart’s action. Moreover,

physiology is not the only science indebted to the frog: in histology

many important results have been obtained from observations on it, and

for histological instruction it is now indispensable. To it we owe much

of our knowledge of the structure of nerve fibres, their origin and

termination, especially in muscle, their relations within the ganglia,

and even the structure of muscular fibre itself. For the study of

reproduction and development the frog has, next to the chick, afforded

the most important material: one need but refer to the investigations

on impregnation from the time of Spallanzani to that of Newport[4], the

phenomena of cleavage, and many others.

[Footnote 3: Leeuwenhoek, Arcana Naturæ III, epist. 65 ad Reg. Soc.

Lond., p. 158.]

[Footnote 4: It may be well to remind the reader that this introduction

was written in 1864.]

Thus with progress of time the field in which the frog has been

submitted to observation and experiment, whether for the demonstration

of established facts to students or for the solving of new problems,

has vastly increased, and this batrachian has indeed become, as we have

stated, the physiologist’s domestic animal.

That, for these manifold uses, a more exact anatomical knowledge of

the frog is very necessary is self-evident. The majority of students

commencing the study of physiology have little more than a superficial

knowledge of the sciatic nerve and the leg-muscles; at most, of the

spinal cord and its nerve-roots; and only acquire any further knowledge

in a disconnected manner. For this they can scarcely be reproached,

the literature of the anatomy of the frog being so widely scattered in

monographs and journals that reference to it involves the expenditure

of much time. This attempt, therefore, to produce a complete anatomy

of the frog, based throughout upon my own observations, cannot be

considered superfluous; it is rather to be feared it may be thought

insufficient.

The European frogs[5] alone are treated of in the following

description, _i.e._ the two species, _Rana esculenta_, L., and _R.

temporaria_, L., the former being more particularly described, though

such differences in structure as occur are noted. This is not the place

to discuss the exact systematic characters of the two species, yet they

cannot be ignored entirely. The species were, from their habitats,

long ago distinguished by C. Gessner[6], and named _Rana rubeta_, _s.

gibbosa_, the garden or grass-frog, and _Rana aquatica_, _s. innoxia_,

the water-frog; at least, from his figure, the former can be no other

than _R. temporaria_, though Gessner, probably expecting to find in it

the _rubeta_ of older writers, adds that it ‘ist für giftig zu halten.’

[Footnote 5: For purposes of comparison other than European frogs

were examined, especially American species of _Rana_, for which I am

indebted to the kindness of my much esteemed friend Agassiz. These were

_Rana Catesbyana_, Shaw (_R. mugiens_, Catesby--_R. pipiens_, Harlan);

_R. sylvatica_, Leconte; _R. clamitans_, Daudin; _R. palustris_,

Leconte; _R. halecina_, Leconte. To these, however, no further

reference will be made in the text.]

[Footnote 6: C. Gessner, Thierbuch. Zürich. Fol., p. 157.]

Leeuwenhoek[7] also correctly distinguished between them, but it is to

Rösel[8] that we are chiefly indebted for a careful discrimination and

an accurate knowledge of the life-histories of the two species.

[Footnote 7: Leeuwenhoek, Arcana Naturæ, Vol. III of Ecker’s edition.

Leyden, 1722. Epist. 65 ad Reg. Soc. Lond., p. 154.]

[Footnote 8: Rösel, Naturgeschichte der Frösche Deutschlands, ed.

Schreber. Nürnberg, 1815, p. 36.]

[Illustration: Fig. 1.

The green water-frog, _Rana esculenta_ L.]

*Rana esculenta*, L. The green water-frog, Fig. 1, usually attains a

larger size[9] and is more active than the other species, and for this

reason is better adapted to the purposes of the physiologist; hence I

have chosen it for description.

[Footnote 9: The extreme sizes are much more pronounced in this

species. I have never found _R. temporaria_ of the size attained by

large specimens of the water-frog.]

The head is flat, as broad as it is long, and triangular with an

obtuse snout in front. The upper surface of the head, _i.e._ the space

between the eyes, is slightly concave, grooved, and narrower than in

_R. temporaria_. The tympanic membrane is circular, and relatively to

the eye is larger. The upper eyelids have several transverse folds in

their hinder part. The pupil is oval, with the long axis horizontal.

The vomerine teeth are arranged in two clusters, which are relatively

larger than in _R. temporaria_ and lie exactly between the posterior

nares, without however touching them. The openings of the Eustachian

tubes do not exceed in size the posterior nares to so great an extent

as they do in _R. temporaria_. The male possesses a vocal sac on

either side, which reaches the surface beneath the tympanic membrane

through a cleft placed behind the angle of the mouth, and is, in

well-developed specimens, about the size of a cherry. The hind limbs

are relatively longer. The toes are long, and taper towards their tips:

the webs between the toes are cut out semicircularly, and that of the

longest or fourth toe is continued to the tip of the last phalanx. The

supplemental toe is an oval prominence of cartilaginous hardness. The

skin of the back has wart-like tubercles arranged longitudinally in

raised lines; one of these lines runs on each side from the posterior

canthus as far as the thigh, and is very constant: in the male a second

line surrounds the posterior margin of the vocal sac; a corresponding

line exists in the female.

The skin of the belly is quite smooth, the colour presenting many

variations which appear to depend upon very diverse circumstances. It

varies with changes in the physiological condition of the animal. Von

Wittich[10] has shown that a bright green specimen changes to a dark

leafy green colour on exclusion of light; also, that dark specimens

become almost a lemon-yellow colour on exposure to bright sunlight;

and he has pointed out that this brightening of the skin is an active

condition dependent upon contraction of the stellate pigment-cells. It

is therefore not surprising, as the same inquirer observes, that one

should sometimes find specimens of _R. esculenta_ in which the ground

colour is almost a greenish yellow (as in Rösel’s figure, Pl. XIII),

whilst in others it can only be distinguished from the dorsal black

patches by a faint greenish shade. There is no doubt that difference

of habitat influences the colour; but this may again be modified by

light[11], as has been established in the case of fish by direct

observation[12]. Apparent varieties may this occur.

[Footnote 10: Von Wittich, Müller’s Archiv, 1854, p. 41.]

[Footnote 11: Lister, On the Cutaneous Pigmentary System of the Frog.

Phil. Trans., 1857, p. 627.]

[Footnote 12: Agassiz et Vogt, Histoire naturelle des poissons d’eau

douce (Neuchatel, 1839), Pl. IV, mention that the colour of trout is

very variable and that in shaded and deep-lying brooks and rivers a

variety is found which is black.]

In frog-tanks such diversities of colour may not unfrequently be

observed in the same individual, as for example when the lower part

of the body immersed in muddy water is dark, while the part above the

water is bright. That the process of casting the skin exercises an

influence on the brightness of the colouring is certain, yet there are,

as von Wittich has correctly remarked, other alterations of colour

which are in no way connected with this process, and are evidently

more of a pathological nature; such as when the frog assumes a dirty

green spotted appearance, the green fading more and more, until all

the patches which are usually green appear of a dirty greyish-brown

with a faint bronze shimmer. According to this author these changes are

most readily brought about by starvation. The dark colour which frogs

exhibit after hibernation is perhaps to be ascribed to the co-operation

of several of the causes mentioned above.

The usual colouring of healthy animals is as follows: the back is

bright green with three golden yellow longitudinal stripes, one median

and two lateral, and a number of irregular brown or black stripes of

approximately uniform width: on the head are a pair of black stripes

which pass from the angles of the eyes across the nares to the tip of

the nose; now and then the tympanic membrane and surrounding parts

have also a black patch, as in _R. temporaria_: another black stripe

is found on the anterior surface of the arm, in the region of the

shoulder: and on the thighs are black, yellow, and white mottlings.

The whole of the under-surface is white or yellowish. At times the

yellow stripes of the back are wanting or are indistinct. It has

already been mentioned that many varieties may occur; and these have

in all probability given rise to the descriptions of reputed new

species, such as _R. maritima_, Risso, found in South Europe; _R.

alpina_, Risso, found in the high-lying Alpine lakes; _R. hispanica_ of

Fitzinger and Bonaparte, and _R. calcarata_ of Michahelles, the last

three of which certainly cannot be retained. It is not improbable

that the water-frog, which Spallanzani[13] used in his experiments on

impregnation, was the _R. maritima_ of Risso. He says, one must not

confound his frog with that which Rösel calls the green water-frog;

the former being much smaller, without the three dorsal golden-yellow

stripes, and the spawning season (in Lombardy) occurring during April

and May. Rusconi[14] also describes two varieties in Northern Italy.

[Footnote 13: Spallanzani, Versuche über Erzeugung der Thiere and

Pflanzen. Leipzig, 1786, p. 5.]

[Footnote 14: Rusconi, Développ. de la Grenouille. Milan, 1826, p. 6.]

*Rana temporaria*, L., the brown or grass-frog, is so named from the

large black patch in the temporal region, _i.e._ between the eye and

the shoulder. While the separation of the preceding species into

several varieties does not seem to be well founded, it appears that two

distinct species have been included under the name of _R. temporaria_.

Millet of Angers[15] first described, in his Fauna du département de

Maine-et-Loire, as ‘grenouille rousse,’ a species differing from _R.

temporaria_, and gave the species previously known as _R. temporaria_

the name of _R. flaviventris_, ‘grenouille à ventre jaune.’ No further

notice, however, was taken of this observation, not even by Duméril and

Bibron in their ‘Erpétologie.’ Quite independently Steenstrup[16], in

the year 1846, pointed out that two frogs, differing in structure and

habits, had been confounded under the name _R. temporaria_; these he

distinguished as _R. platyrhinus_ and _R. oxyrhinus_. Von Siebold[17],

and also Schiff[18] in part, have confirmed these statements. My

own observations lead me to a like conclusion; I shall therefore

distinguish two species, viz.:--(1) _Rana temporaria_, L., _Rana

platyrhinus_, Steenstrup; (2) _Rana oxyrhinus_, Steenstrup.

[Footnote 15: Annales des Sciences naturelles. Zoologie, IV Série,

Vol. IV, 1855, p. 368.]

[Footnote 16: Amtl. Bericht über die 24. Versammlung deutscher

Naturforscher in Kiel, 1846, p. 141; Wiegmann’s Archiv, 1847, Vol. II,

p. 341; Steenstrup, Oversigt K. Danske Selsk. Forhandlgr., 1846, p. 92.]

[Footnote 17: Wiegmann’s Archiv, 1852, Vol. I, p. 14.]

[Footnote 18: Annales des Sciences naturelles. Zoologie, IV Série,

Vol. IV, 1885, p. 368.]

[Illustration: Fig. 2.

The brown grass-frog, _Rana temporaria_, L.]

*Rana temporaria*, L.; _Rana platyrhinus_, Steenstrup. The brown

grass-frog, Fig. 2, does not attain the dimensions of _R. esculenta_,

L., but is, however, always larger than _R. oxyrhinus_. The head is

somewhat broader than long, and the upper surface of the skull is not

grooved, as in _R. esculenta_, but is flat. The space between the

eyes is wider (according to Duméril, equal to the width of the upper

eyelid, whereas in _R. esculenta_ it is just two-thirds this width):

the fronto-parietal bones are wide and flat. The tympanic membrane,

in comparison with the eye, is smaller than in _R. esculenta_, and is

usually less distinguishable from the surrounding parts as regards

colour and transparency. The apertures of the Eustachian tubes are,

relatively to the posterior nares, larger than in the water-frog. The

vomerine teeth are comparatively small and lie in two groups placed

obliquely to each other, their anterior ends diverging from each other

and being prolonged as ridges to the anterior margins of the posterior

nares. The two groups do not lie between the nasal apertures, but

behind a line drawn transversely through their posterior margins. Vocal

sacs are absent in both sexes. The hind legs are relatively shorter:

the toes are not so evenly tapered off, indeed they are slightly

swollen: the fourth toe, as compared with the third and fifth, is

somewhat longer than in _R. esculenta_; the web of this toe does not

extend to the tip of the toe, but terminates in both sexes at the last

phalanx but one; the web on the third toe is less developed on the

thumb side than on the other: on the remaining toes also the margins

of the web are less developed than in _R. esculenta_, so that the

free borders appear more crescentic. The supplemental toe forms only

a soft and inconspicuous prominence. The back is mostly smooth; the

raised glandular ridge, which extends along each side from the eye to

the thigh, is present, but is much narrower and less prominent than in

_R. esculenta_; another ridge passes from the angle of the mouth to the

shoulder. The colouring in general, and especially the ground colour

of the dorsal surface, varies from the brightest tints to the darkest

brown-black; the conditions causing these variations being, no doubt,

the same as those described above in _R. esculenta_. A dark-brown

specimen taken from a dark frog-tank is usually yellowish red on the

following day. The black patch between the angle of the mouth and the

shoulder has given this species the name of _R. temporaria_, and is

constant. A black stripe passes from the eye across the nostril to

the tip of the snout, and a similar one is found upon the anterior

surface of the upper arm. On the hind legs the bands are chiefly

transverse. The ventral surface is yellowish, and sometimes spotted.

The thighs have a granular appearance, and these as well as the belly

and the neighbourhood of the anus have frequently a reddish coloration

presenting the appearance of an irritated surface.

*Rana oxyrhinus*, Steenstrup. This species is always smaller and more

elegant in shape than the preceding one. The head is conical, with

the pointed snout projecting beyond the lower jaw; a feature which is

especially evident on looking from below. The space between the eyes

is narrower than in _R. temporaria_, and is not grooved, but convex;

the fronto-parietal bones are narrow and arched. With respect to the

arrangement of the vomerine teeth and the sizes of the apertures of the

Eustachian tubes, this species holds an intermediate position between

the other two. Next to the pointed snout, the greatest difference

between this species and _R. temporaria_ is the presence of a much

larger supplemental toe, which is of cartilaginous hardness, compressed

from side to side, and contains a larger bone[19]. The vocal sacs

are absent. In the males the web of the longest toe reaches to the

last phalanx but one; in the females, on the contrary, the last three

phalanges project freely beyond the web. The extremities of the toes

are more pointed than in _R. temporaria_, in which respect, as also

in several others, it approaches _R. esculenta_. In colouring, _R.

oxyrhinus_ resembles _R. temporaria_; the throat, however, is usually

pure white, at least in the males, the breast dusky white and spotted,

while in _R. temporaria_ the throat and breast are more uniformly

coloured and yellowish. V. Siebold has remarked that, during the

pairing-season, the males are covered with a bluish bloom[20]; and, the

whole ground colour being bright at this period, very beautiful tints

result. V. Siebold[21] moreover states that the note which the males

produce during the pairing-season is different in the two species.

On the whole, _R. oxyrhinus_ appears to stand midway between _R.

esculenta_ and _R. temporaria_.

[Footnote 19: According to Steenstrup, _l. c._, the size of the

supplemental toe is in _R. temporaria_ one-half and in _R. oxyrhinus_

two-thirds of that of the next toe.]

[Footnote 20: I do not find, however, that this disappears when the

animals are on land; in fact I have animals before me in a glass in

which it is plainly seen.]

[Footnote 21: _l. c._, p. 15.]

Thomas[22], in addition, distinguishes another species, _R.

agilis_, which however may be the ‘grenouille rousse’ of Millet.

Schlotthauber[23] has described a frog which, in marking and colouring,

might hold a middle place between _R. esculenta_ and _R. temporaria_;

in my opinion this is probably a cross between the two. That attempts

at copulation are made, despite the difference of the pairing-season,

is well known; Pontallié[24] mentions this, and I have myself often

found males of _R. temporaria_ in conjunction with females of _R.

esculenta_.

[Footnote 22: Annales des Sciences naturelles. Zoologie, IV Série,

Vol. IV, 1855.]

[Footnote 23: Wiegmann’s Archiv, Vol. I, 1844, p. 255.]

[Footnote 24: Annales des Sciences naturelles. Zoologie, III Série,

Vol. XVIII, 1852, p. 243.]

* * * * *

I use the following terminology. I suppose the animal to be in its

natural position, the belly towards the ground, the back upwards; a

horizontal plane passing from the snout to the anus divides the body

into a superior or dorsal half and an inferior or ventral half. The

terms superior and inferior, dorsal and ventral, indicate positions

with relation to this plane. I call that part anterior which looks

towards the head, and that posterior which looks towards the anus. A

vertical plane at right angles to the middle of the longitudinal axis

of the body, divides it into an anterior or cephalic and a posterior

or caudal half. All sections and planes which lie parallel to this, as

well as this itself, are frontal. Lastly, by a perpendicular section

along the middle line of the body the animal is divided into right and

left halves; this plane is the median plane; and the position relative

to this plane is expressed by the terms median or lateral. Planes

parallel to the median plane are termed sagittal.

SECTION I.

THE BONES AND JOINTS.

THE BONES AND JOINTS.

LITERATURE.

*van Altena*, Commentatio ad quæst. zoologicam in academia

Lugduno-Batav. a. MDCCCXXVIII propositam, qua desideratur ut

systematice enumerentur species indigenæ reptilium ex ordine

batrachiorum addita unius saltem speciei anatomia et præsertim

osteographia accurata. Lugd. Bat. 1829. 4^o. With 4 Plates.

*Ange, Martin St.*, Recherches sur les organes transitoires des

batraciens. Annales des Sciences naturelles. 1^{re} Série.

Vol. XXIV. 1831.

*Bell*, Article Amphibia, in Todd’s Cyclopaedia of Anatomy and

Physiology. Vol. I, p. 90. 1835–1836.

*Born, Dr. Gustav*, Ue.d. Nasenholen u.d. Thränennasengang der

Amphibien. Leipzig, 1877.

*Bruch, G.*, Beiträge zur Naturgeschichte und Klassification der

nackten Amphibien. Würzburger Naturzeitschrift, 1862.

*Bruch, G.*, Neue Beobachtungen zur Naturgeschichte der einheimischen

Batrachier. Würzburger Naturzeitschrift, 1863.

*Cuvier*, Recherches sur les ossements fossiles. Vol. V. Pt. II.

Paris, 1825.

*Cuvier*, Leçons d’anatomie comparée. Paris, 1835. Vol. I.

*Cuvier*, Ueber die Rückenwirbel der Reptilien und Amphibien,

Froriep’s Notizen. Vol. XIII, p. 74. 1826.

*Daudin*, Histoire naturelle des Rainettes, Grenouilles et des

Crapauds. Paris, 1802.

*Ducrotay de Blainville*, Ostéographie ou description iconographique

comparée du squelette et du système dentaire des cinq classes

d’animaux vertébrés. Paris, 1841.

*Dugès*, Recherches sur l’ostéologie et la myologie des batraciens à

leurs différents âges. Paris, 1834. 4^o. With 20 Plates.

*Duméril et Bibron*, Erpétologie générale ou Histoire complète des

Reptiles. 1836.

*Gegenbaur*: 1. Ueber Bau und Entwicklung der Wirbelsäule bei

Amphibien überhaupt und beim Frosche insbesondere. Abhandlungen

der naturforschenden Gesellschaft zu Halle, Vol. VI. Halle, 1861.

2. Untersuchungen zur vergl. Anatomie der Wirbelsäule bei

Reptilien und Amphibien. Pt. I. Leipzig, 1862. (Carpus and

Tarsus.) With 4 Plates. 4^o.

*Gegenbaur*, Untersuchungen zur vergl. Anatomie der Wirbelthiere.

Pt. II. Schultergürtel. 1865.

*Günther*, Ueber geschlechtliche Differenzen in Knochen von lebenden

und fossilen Fröschen und Fischen. Annals of Natural History.

1859. Vol. III.

*Hallmann*, Die vergleichende Osteologie des Schläfenbeins, etc.

Hannover, 1837. 4^o. With 3 Plates.

*Hoffmann, C. K.*, Beiträge zur Erkenntniss des Beckens der Amphibien

und Reptilien. Leyden, 1876.

*Hoffmann, C. K.*, Bronn’s Klassen und Ordnungen des Thierreichs,

Vol. VI. Amphibien. Leipzig, 1873–8.

*Huxley*, On the Theory of the Vertebrate Skull; Croonian Lecture,

Proc. Royal Society, p. 381. 1858.

*Huxley*, Article Amphibia, Encyclopædia Britannica, IXth Edition.

1875.

*Huxley*, Lectures on the Elements of Comparative Anatomy.

*Huxley*, Handbuch der Anatomie der Wirbelthiere. Deutsche Ausg. von

T. Ratzel. 1873.

*Kehrer, G.*, Beiträge zur Kenntniss d. Carpus und Tarsus d.

Amphibien, Reptilien, und Säuger. Berichte d. naturf. Gesell. z.

Freiburg. 1886.

*v. Klein*, Beiträge zur Anatomie der ungeschwänzten Batrachier.

Jahres-Heft. Würtemberg, 1850.

*Köstlin*, Der Bau des knöchernen Kopfs. Stuttgart, 1844. 8^o.

*Leukart*, Zwischenkiefer. Valentins Repertoire. 1841, p. 155.

*Marshall, A. M.*, The Frog. Manchester and London. 2nd Edit., 1885,

pp. 45–59.

*Mayer, A. F.*, Beiträge zu einer anatomischen Monographie der Rana

pipa. Acad. Caes. Leop. Nov. Acta. 1825. Vol. XII, p. 527; and

Isis v. L. Oken. 1825. col. 317.

*Meckel*, System der vergleichenden Anatomie. II. Thl. I. Abthlg.

Halle, 1824. 8^o.

*Meckel*, Ueber das Zungenbein der Amphibien. Meckel’s Arch. f.

Physik. 1818. Vol. IV, p. 60.

*Mertens*, Anatomiæ batrachiorum prodromus sistens observationes

nonnullas in osteologiam batrachiorum nostratium. Halæ, 1820. 8^o.

*Mivart*, On the Classification of the Anurous Batrachians. Proc.

Zool. Soc. 1869.

*Morren*, Observations ostéologiques sur l’appareil costal des

batraciens. Bulletins de l’Acad. de Bruxelles, 1835, II.--Mémoires

de l’Académie, 1837. Tome X.

*Müller*, Beitrag zur Anat. d. Amph. Zeitschrift f. wissenschaftliche

Zoologie. Vol. IX. 1858, p. 178.

*Parker, W. K.*, Structure and Development of the skull of the common

frog. Phil. Trans. 1871, p. 137.

*Parker, W. K.*, Skull of Batrachia. Phil. Trans. 1876, p. 601.

*Parker, W. K.*, and *Bettany, G. T.*, Morphology of the Skull.

London, 1877.

*Pouchet*, Note sur les différences que le sexe imprime au squelette

des grenouilles. Comptes rendus. Vol. XXV, p. 761. 1847.

*Reichert, K. B.*, Vergleichende Entwicklungsgeschichte des Kopfs der

nackten Amphibien nebst den Bildungsgesetzen des Wirbelthierkopfs

im Allgemeinen und seinen hauptsächlichen Variationen durch die

einzelne Wirbelthier-Classe. Königsberg, 1838.

*Remak*, Untersuchungen über die Entwicklung der Wirbelthiere.

Berlin, 1855.

*Rösel, von Rosenhof*, Historia naturalis ranarum nostratium.

Nörnberg, 1758.

*Rudolphi and Breyer*, Observationes anatomicae circa fabricam Ranae

pipae. Berolini, 1811.

*Rusconi*, Développement de la grenouille commune. Milan, 1826.

*Rusconi*, Sulle metamorfosi delle osse della testa della rana.

Annali di Bologna. 1^{re} Série, Vol. II, p. 357.

*Schneider*, Historia amphibiorum. Jenae, 1799.

*Shaw*, General Zoology. London. Vol. II, Pt. I, p. 167.

*Stannius*, Zootomie der Amphibien (Handb. der Zootomie der

Wirbelthiere, 2. Buch). 2nd Edit. Berlin, 1856. 8^o.

*Stricker*, Untersuchungen über die Entwicklung des Kopfes der

Batrachier. Arch. f. Anat. u. Physiol. 1864, pp. 52–76.

*Stricker*, Beiträge zur Biologie der Batrachier. Verhandl. der

Wiener Akademie. 1866. Vol. XVI, pp. 451–456.

*Townson, R.*, Facts and Observations in Natural History. London,

1799.

*Troya*, Mémoire sur la structure singulière du tibia et du cubitus

des grenouilles et des crapauds. Mémoires de mathématique et de

physique présentées à l’acad. de Paris. Vol. IX. 1780.

*Wagner*, Icones Zootomicae. Leipzig, 1841.

*Wagner*, Lehrbuch der vergleichenden Anatomie. Leipzig, 1834–1835.

*Wiedersheim, R.*, Lehrbuch d. vergleichenden Anatomie der

Wirbelthiere auf Grundlage d. Entwickelungsgeschichte. Jena, 1886.

2nd Edit.

*Wiedersheim, R.*, Elements of Comparative Anatomy of Vertebrates,

translated by W. Newton Parker. London, 1886.

THE BONES AND JOINTS.

The consideration of the differences in form, number, and histological

structure, which the parts of the skeleton present during the various

stages of development does not fall within the scope of this book: we

have here but to deal with the adult frog.

The skeleton is made up of histologically different materials;

these are:--(1) bone, (2) hyaline cartilage, and (3) so-called

calcified cartilage. Concerning the last it is necessary to make some

observations. I have chosen for it the name calcified cartilage in

place of the more usual names ‘cartilaginous bone’ or ‘primordial

ossification,’ as by this term its nature appears to be expressed

without any ambiguity[25]: it is hyaline cartilage in which calcareous

particles have been deposited to a greater or less extent: in the fresh

state it has the appearance of moderately firm cartilage; when dry it

becomes opaque and white, like the calcareous crusts on the cartilages

of the Plagiostomata. The calcareous material is deposited in the

cartilage in finer or coarser granules; after removal of the lime by

means of acids, the cartilaginous structure becomes apparent although

not so perfectly as in unchanged cartilage.

[Footnote 25: Compare Müller, Zeit. f. wissen. Zoolog., Vol. IX.]

This calcified cartilage is widely distributed in the frog’s skeleton:

very many parts, which in higher animals consist only temporarily of

this substance during the transition from cartilage to bone, are in

the frog formed of it throughout life. It is especially well-marked

in the epiphyses of the long bones in the hand and foot, in the

bones of the shoulder-girdle, etc. To avoid repetition later on I

will briefly describe it as found in the first-mentioned situation.

Dugès[26] has described its external appearance, while Bruch[27] has

made us acquainted with its histological peculiarities. If a long

bone of the frog be dried, the femur for example, the middle part is

found to differ considerably from the epiphyses in colour and in other

particulars. The shaft alone has the appearance of bone, the epiphysis

consisting of a white, opaque, firm substance, resembling plaster of

Paris or lime, but which in the fresh moist state is exactly like

cartilage. The epiphyses, which are fitted to the ends of the diaphysis

like the cap of a stick-handle, have sharply defined margins (Fig. 3),

as is well seen in Figs. 36, 39, 45, and 46. If a section be made

through the epiphysis and part of the diaphysis, the long tube of true

bone is seen to cease abruptly above _o_, Fig. 3, and over the end of

it the epiphysis _E_ is fixed. This epiphysis consists almost entirely

of calcified cartilage _c′_, and has merely a superficial layer of

hyaline cartilage _A_. The bony cylinder of the diaphysis _o_, which

contains the marrow _M_ in its interior, is shut off from the epiphysis

by hyaline cartilage, the cells of which are arranged in transverse

layers, _o_.

[Footnote 26: Dugès, Recherches sur l’ostéologie et la myologie des

batraciens à leurs différents âges, p. 116.]

[Footnote 27: Bruch, Beiträge zur Entwicklungsgeschichte des

Knochensystems. Schweiz. Denkschriften, p. 118.]

[Illustration: Fig. 3.

Longitudinal section through the upper extremity of the femur of _Rana

esculenta_, magnified.

_A_ Hyaline cartilage (articular cartilage).

_c_ Hyaline cartilage closing end of bony cylinder.

_c′_ Calcified cartilage of epiphysis.

_D_ Bony cylinder of diaphysis.

_E_ Epiphysis.

_M_ Marrow cavity.

_o_ End of diaphysis.

]

I. THE VERTEBRAL COLUMN.

The _vertebral column_ of the frog consists of ten bones, _viz._ nine

true vertebrae, and the rod-shaped urostyle, which alone is almost as

long as all the remaining vertebrae.

I. GENERAL DESCRIPTION OF THE NINE VERTEBRAE.

*a.* The _bodies_ of the vertebrae are compressed from above downwards;

the posterior surface of each body, with the exception of the eighth,

presents an articular head covered with cartilage; the anterior

surfaces, with the exception of the ninth, present corresponding

articular depressions, covered with cartilage.

*b.* The _arches_, which have somewhat sharp margins both before and

behind, bear the following processes:-- *1.* The *articular processes*

(Figs. 4 and 5 _o o_) are similarly placed to those of the dorsal vertebrae

of man: they project horizontally, the cartilaginous articular surfaces

on the posterior processes being directed downwards, those on the

anterior upwards.

[Illustration: Fig. 4.

Vertebrae of _Rana esculenta_, seen from below, twice the natural size.

1 to 9 First to ninth vertebræ.

_c_ Urostyle.

_o o_ Articular processes.

_sc_ The two facets for articulation with the urostyle.

]

*2.* The *transverse processes* (Figs. 4 and 5 _t t′_) are strong,

flat, and of very varying size and direction. The transverse processes

of the fourth vertebra are the longest, those of the third only a

little shorter; the shortest are those of the seventh and eighth.

The atlas has no transverse processes. Those of the second and third

vertebrae project directly outwards and slightly downwards; those of

the fourth, fifth, and sixth upwards and backwards. The seventh and

eighth project more directly outwards and at the same time backwards;

the ninth upwards and markedly backwards. All the transverse processes

have cartilaginous epiphyses; the largest are those of the second,

third, fourth, and ninth vertebrae.

*3.* The *spinous processes* are generally small, but individually of

varying size, appearance, and direction. The longest are those of the

third, fourth, and fifth vertebrae; these are, in transverse section,

of a three-sided prismatic form, as in the dorsal vertebrae of man;

they are directed backwards and provided with cartilaginous epiphyses.

The spinous processes of the sixth and seventh are shorter, compressed

from side to side, project directly upwards, and are usually without

cartilaginous epiphyses; that of the eighth is still shorter. As

regards the spinous processes, those of the third, fourth, and fifth

vertebrae resemble those of the dorsal vertebrae in man; those of the

sixth, seventh, and eighth, lumbar vertebrae. The ninth has either

no spinous process or only a rudimentary one. The first and second

vertebrae may be looked upon as cervical vertebrae: the second has a

short spinous process with a cartilaginous epiphysis. In the first,

the cartilage which unites the two halves of the arch represents the

rudiment of a spinous process.

[Illustration: Fig. 5.

Vertebral column of _Rana esculenta_, from above, twice nat. size.

1 to 9 First to ninth vertebræ.

_c_ Urostyle.

_o o_ Articular processes.

_sc_ Facets for articulation with the urostyle.

_t t′_ Transverse processes.

]

II. DESCRIPTION OF PARTICULAR VERTEBRAE.

*1.* The *atlas* or *first vertebra* has a thin body, compressed from

above downwards, and an arch. The body has posteriorly a slightly

raised, cartilaginous, articular head, which is broader transversely:

in front it has two oval articular facets, which are separated from

each other by a median projection. Each facet is concave, and directed

forwards, outwards, and slightly upwards. The arch is completed above

by cartilage, which projects slightly to form the rudiment of a spinous

process. The hinder margin of the arch bears two articular processes.

Transverse processes are wanting.

*2.* The *second vertebra* presents all the general characters of an

ordinary vertebra, except that the transverse processes are directed

somewhat downwards.

*3.* The transverse processes of the *third vertebra* are longer than

those of the second: each is directed downwards, is broader at its

extremity than at its base, and bears a hammer-shaped cartilaginous

epiphysis larger than those of the remaining transverse processes.

*4.* The transverse processes of the *fourth vertebra* are the longest:

each is broader at its free end than at its base, is directed upwards

and backwards, and provided with a cartilaginous epiphysis.

*5*, *6*, *7*. The transverse processes of the *fifth*, *sixth*,

and *seventh vertebrae* are smaller, contracted towards their free

extremities, and directed upwards.

*8.* The *eighth vertebra* is distinguished from the rest by its body

possessing no articular head. It presents, at each end, a concave

articular depression. The transverse processes resemble those of the

seventh.

*9.* The *ninth vertebra* unites the vertebral column with the

hip-bones, and is hence to be regarded as a *sacrum*. The body bears on

its anterior surface an articular head for articulation with the eighth

vertebra: on its posterior surface are two small rounded and closely

approximated processes (Figs. 4 and 5 _sc_) for articulation with the

urostyle. The transverse processes are strong, broader at the free ends

than at their origin, directed upwards and backwards, and provided with

cartilaginous epiphyses.

III. ARTICULATIONS OF THE VERTEBRAE.

The articular heads and depressions of the vertebral bodies, together

with the joint surfaces of the articular processes, are covered with

hyaline cartilage. The periosteum of the bodies, as also that of

the articular processes, forms true capsular joint ligaments. The

articulations of the vertebrae are still further strengthened by

longitudinal fibres, which extend along the anterior and posterior

surfaces of the vertebrae, and correspond to the _ligamentum vertebrale

commune anticum et posticum_ of man. Between the vertebral arches are

membranes which represent the _ligamenta intercruralia_. Between the

spinous processes are bands of connective tissue which form _ligamenta

interspinalia_. (For the articulation of the atlas with the occiput,

see page 24.)

IV. STRUCTURE OF THE VERTEBRAE[28] (Figs. 6 and 7).

[Footnote 28: Cf. Gegenbaur, (1) Über Bau u. Entwicklung der

Wirbelsäule bei Amphibien überhaupt u. beim Frosch insbesondere.

Abhand. d. Naturforsch. Gesell. zu Halle, vol. vi, 1861; (2)

Untersuch. zur vergleich. Anat. d. Wirbelsäule bei Amphibien u. Rept.,

Leipzig, 1862. As regards the adult animal I can bear out Gegenbaur’s

observations. The scope and limits of this book forbid me to go further

into the developmental history.]

[Illustration: Fig. 6.

Transverse section through a vertebra of _Rana esculenta_, magnified.

_c_ Cancellous bone.

_Ch_ Chorda dorsalis.

_Ch′_ Sheath of chorda dorsalis.

_o_ Compact bone on the upper and lower surfaces of the body.

]

Each vertebral body consists of a cylinder of compact bone, which is

directly continued into the bony substance of the arch. In the interior

of the cylinder is found an isolated persistent vestige of the _chorda

dorsalis_ (_Ch_): this is surrounded by cancellous bone (_c_), which

extends backwards towards the articular head and forwards directly into

the articular cartilage, compact bone being absent in these parts. In

a transverse section of a vertebral body the following parts are seen

(Fig. 6):--*a.* An outer layer of compact bone (_o_) (the transverse

section of the above-mentioned cylinder), which is formed of parallel

lamellae of varying thickness. These, according to Gegenbaur, and as

I can confirm, are arranged in well-defined groups, each of five to

eight lamellae. The number of the secondary lamellae increases with the

age of the animal. *b.* In the interior, in the form of a cylinder, is

the remnant of the _chorda dorsalis_. It consists of a double sheath

(_Ch′_) and contents (_Ch_) composed of chorda-cells. *c.* Immediately

around the persistent portion of the chorda lies the central part of

the vertebral body, formed by transformation of the vertebral cartilage

and of the bases of the original cartilaginous arches. At each side of

the _chorda_ are large marrow-spaces (_c_), filled with cells, from

which proceed narrower canals, winding in various directions, and

anastomosing freely with one another both before and behind. Their

walls are constituted partly of true bone, partly of cartilage.

[Illustration: Fig. 7.

Longitudinal section through the posterior half of the body of a

vertebra of _Rana esculenta_.

_a_ Cartilage of the head.

_c_ Cancellous bone.

_o_ Shell of compact bone.

]

V. THE UROSTYLE (Figs. 8 and 9).

The *urostyle* is a long, median, rod-like bone, which projects

backwards, midway between the two hip-bones, and terminates over the

anus. The anterior end (Fig. 8 _a_) is the thicker and broader part of

the bone, and has two articular depressions (Fig. 9) for articulation

with the two facets of the ninth vertebra. The hinder end is pointed

and cylindrical, and terminates in a cartilage, which is fixed in the

tubular end of the bone. The middle portion is almost cylindrical, and

has a groove along the ventral surface which gradually becomes less

marked behind. The dorsal surface bears a ridge (Figs. 8 and 9 _s_),

which is high and thick in front, becomes sharper and less prominent

as it proceeds backwards, and gradually disappears towards the

hinder third of the bone, so that in transverse section the anterior

two-thirds of the bone appear triangular, with a ventral and two

lateral surfaces: while the hinder third is cylindrical. The anterior

portion of the bone contains a canal, _canalis vertebralis_ (Fig. 9

_c.v._), which is a continuation of the vertebral canal, along which

the hindermost spinal nerves pass. On each side of the anterior portion

of the urostyle are small apertures (Fig. 8 _c.c._), which lead into

canals (_canales coccygei_), which open into the vertebral canal, and

through which the coccygeal nerves pass. In front of these openings

and partly overhanging them are small triangular projections (Fig. 8)

(_processus transversarii_): these, however, are not constant, and are

more often found in _R. esculenta_ than in _R. temporaria_, in which

latter species the openings are smaller.

[Illustration: Fig. 9.

Urostyle of _Rana esculenta_, seen from the side, twice nat. size.

_c_ Ventral border.

_c.v._ Canal. vertebralis.

_s_ Dorsal ridge (pr. spinos.).

]

[Illustration: Fig. 8.

Urostyle of _Rana esculenta_, seen from the side, twice nat. size.

A bristle is passed through the _canal. vert._ and out through the

_canal. coccyg._ of the right side.

_a_ Anterior extremity.

_c.c._ Canal. coccyg.

_s_ Dorsal ridge (pr. spinos.).

]

II. THE SKULL.

The flat form of the frog’s head, as in batrachians generally, depends

upon the wide separation of the jaw-bones of the two sides, and on the

large size of the orbital cavities and the horizontal direction of

their floors. The outer circumference of the head forms a parabolic

frame (Figs. 10 and 11), composed of the maxillary (_m_), premaxillary

(_i_), and quadratojugal bones (_j_). In the middle of this curved

framework lies the elongated prismatic cranium. Anteriorly, this is

attached to the fore-part of the frame by means of the cartilaginous

skeleton of the organs of smell (Fig. 11 _e′_); posteriorly, it

widens out into two transverse arms (_p_), which contain the organs

of hearing. From this base, on either side, a bony strut, composed

of the posterior arms of the squamosal (_t′_) and of the pterygoid

bones, passes backwards to the hinder end of the frame. The anterior

arm of the squamosal bone (_t_) does not quite reach the framework,

but is attached to it by ligament alone. Between the last-named

arm posteriorly, the cranium on the inner side, and the maxillary

frame-work laterally, is a large space representing the orbital and

temporal fossae of human anatomy.

[Illustration: Fig. 10.

Skull of _Rana esculenta_, seen from above, twice natural size.

_e_ Sphenethmoid.

_fn_ Nasal.

_f.p._ Fronto-parietal.

_i_ Premaxillary.

_j_ Quadrato-jugal.

_m_ Maxillary.

_o_ Exoccipital.

_op_ Opisthotic.

_p_ Prootic.

_pt_ Pterygoid.

_pt′_ Posterior limb of pterygoid.

_t_ Squamosal.

_t′_ Posterior arm of the same.

]

[Illustration: Fig. 11.

Skull of _Rana esculenta_, seen from below, twice natural size.

_c_ Cartilaginous wall of skull.

_e_ Sphenethmoid.

_e′_ Cartilaginous skeleton of nose.

_h′_ Stylo-hyoid.

_i_ Premaxillary.

_m_ Maxillary.

_m′_ Quadrate tract.

_o_ Exoccipital.

_p_ Prootic.

_p′_ Anterior arm of prootic (_ala magna_ autt.).

_p″_ Trigeminal foramen.

_pl_ Palatine.

_pt_ Pterygoid.

_pt′_ Posterior arm of pterygoid.

_s_ Parasphenoid.

_v_ Vomer.

]

A. THE CRANIUM.

The cranium of the frog is a prismatic tube, wide behind, narrow in

front, and formed in great part of cartilage (Figs. 15 and 17). Our

indigenous species are characteristically distinguished from one

another by peculiarities in the form of the cranium. In _R. esculenta_

it is long and narrow, in _R. temporaria_ short and wide. The superior

surface in the former is markedly concave, while in the latter it is

flat, and in _R. oxyrhinus_ arched. These differences are readily

recognised in the living animal.

*The Bones of the Cranium.*

*1.* The *exoccipital bones*, _ossa occipitalia lateralia_, Cuvier

(Figs. 10, 11, 12, 14, 16 _o_).

*Cuvier*, _l. c._, p. 387, Pl. XXIV, bb.--*Dugès*, _l. c._, n.

14.--*Parker* and *Bettany*, _l. c._, p. 166, exoccipitals.

[Illustration: Fig. 12.

Skull of _Rana esculenta_, seen from behind, twice natural size.

_a_ Stapes.

_a′_ Columella auris.

_a″_ Extrastapedial.

_c_ Condyles of exoccipitals.

_h_ Stylo-hyoid.

_o_ Exoccipitals.

_p_ Prootic.

_p′_ Process to which the jaw is attached.

_t_ Squamosal.

]

These paired bones form the hinder part of the cranium; they bound the

_foramen magnum_, and articulate with the vertebral column. They are

imbedded in the cartilaginous matrix of the skull, and are separated

above by an unossified part of this matrix (_occipitale superius_,

Dugès), which represents the tabular portion of human anatomy: below

they are separated by a similar part (_occipitale basilare_, Dugès)

which represents the body of the occipital bone. They therefore

properly represent only the condylar portions (_partes condyloideae_)

of the human occipital bone. Each possesses a cartilaginous articular

head, for articulation with the first vertebra: these converge below,

and surround the lower half of the circumference of the _foramen

magnum_. This latter has, in _R. esculenta_, a transversely oval

outline; in _R. temporaria_, a somewhat heart-shaped outline, with

the apex directed upwards: in accordance with this the whole bone is

wider than high in the first species; and in the latter it is higher

than it is wide. From the upper and outer border of the _foramen

magnum_ on each side a ridge runs obliquely outwards and downwards,

in which lies the suture between this bone and the prootic bones. This

bony ridge (_processus mastoideus_, autt.) is usually cartilaginous

in _R. esculenta_, even in old animals; in _R. temporaria_, even in

young specimens, it is bony. In the latter species the bones unite

very early, while in the former they remain separated by the primitive

cartilage. Between this crest and the _processus condyloideus_

there is a depression (_fossa condyloidea_), with a hole (_foramen

condyloideum_) through which the vagus nerve leaves the cranium. The

exoccipital take part in the formation of the labyrinth of the ear, as

will be noticed later on.

_Articulation of the Exoccipital Bones with the Atlas._ From the middle

of the anterior surface of the body of the atlas a ligament arises,

representing to a certain extent the _lig. suspensorium dentis_, and

attached to the basal portions of the exoccipital bones.

*2.* The *prootic bones*, _ossa petrosa_, Cuvier (Figs. 10, 11, and

12 _p_).

*Cuvier*, rocher, _l. c._, p. 388, Pl. XXIV, ee.--*Dugès*, n. 12,

rupéo-ptéréal.--*Stannius*, ala temporalis.--*Meckel*, Schädelstück

des Schläfenbeins.--*Parker* and *Bettany*, _l. c._, prootic.

These paired bones lie at the sides and in front of the exoccipital

bones. As already explained, they remain in _R. esculenta_ separated

from these by cartilage, while in _R. temporaria_ they early enter

into bony union with them; this is due to the complete ossification

of the _processus mastoideus_ in the latter species, as stated above.

The prootics form the lateral expansions of the posterior part of the

skull in which the organs of hearing are placed. The large cavity which

contains the ear labyrinth is completed by the exoccipital: internally

it opens freely into the skull, and externally on the posterior wall

of the skull through the _foramen ovale_, which is formed by both

these bones. The postero-lateral part of the prootic usually remains

cartilaginous: at the side and in front of the _foramen ovale_ this

cartilage is pierced by a small opening, through which passes the

_nervus facialis_ or _ramus tympanicus n. vagi_ (Volkmann). At the

side there is a process to which the suspensorium of the lower jaw is

attached (Fig. 12 _p_): behind this is a hollow in which the auditory

ossicles lie, and which may be designated _fossa tympanica_ (Fig. 12

_t_). The anterior border of the bone forms the hinder and inner

walls of the orbit. Here also is the trigeminal foramen (Fig. 11

_p″_) through which the _N. trigeminus_ and the several nerves for

the muscles of the eye pass; it represents the _foramen ovale_, _for.

rotundum_, and the _fissura orbitalis superior_ (sphenoidal fissure) of

the human sphenoid bone. The foramen is sometimes, especially in young

animals, only a notch, which is completed by cartilage. On account of

the relation of this part (Fig. 11 _p′_) of the bone to the nerves

which pierce it, the whole bone has been named by Stannius the _ala

magna_ or _temporalis_ of the sphenoid; it has been also looked upon as

a bone which contains these elements, as by Dugès, who on this account

calls it rupéo-ptéréal.

[Illustration: Fig. 13.

Skull of _Rana esculenta_, seen from below, twice natural size.

_c_ Cartilaginous wall of skull.

_e_ Sphenethmoid.

_e′_ Cartilaginous nasal skeleton.

_h′_ Stylo-hyoid.

_i_ Premaxillary.

_m_ Maxillary.

_m′_ Quadrate tract.

_o_ Exoccipital.

_p_ Prootic.

_p′_ Anterior arm of prootic.

_p″_ Trigeminal foramen.

_pl_ Palatine.

_pt_ Anterior arm of pterygoid.

_pt′_ Posterior arm of pterygoid.

_s_ Parasphenoid.

_v_ Vomer.

]

_Appendages of the Prootic Bones._

1. The *styloid cartilage*. From the cartilaginous portion of the

prootic the styloid cartilage runs downwards, backwards, and inwards,

and is continued directly into the anterior cornu of the hyoid bone

(Figs. 11 _h′_ and 12 _h_).

2. The *auditory ossicles*.

_a._ A thick cartilaginous disc, the _operculum_ (Fig. 12 _a_),

closes the _foramen ovale_.

_b._ To the operculum is attached a bony, club-shaped piece, the

_columella auris_ (Fig. 12 _a′_), which has at its inner, thicker end

a cartilaginous epiphysis, the *interstapedial*; it lies transversely

with the apex directed outwards, and this longer portion is the

*mediostapedial*.

_c._ To the apex of the mediostapedial is attached, at an obtuse

angle, the third cartilaginous piece, the *extrastapedial* (Fig. 12

_a″_). It is attached to the tympanic membrane, and by its upper

portion is fastened to the cartilaginous tympanic ring by a smaller

piece, the *suprastapedial*.

3. The *tympanic ring* (_annulus tympanicus_) is an annular

cartilaginous frame; or more exactly, has the shape of a short,

truncated cone, as it narrows towards the middle line: it is attached

to the squamosal bone. (See Organ of hearing.)

*3.* The *parasphenoid*, _os sphenoideum_, Cuvier (Figs. 11 and 16

_s_).

*Cuvier*, _l. c._, p. 388, Pl. XXIV, d.--*Dugès*, n. 8.--*Meckel*,

Theil des Grundbeins.--*Parker* and *Bettany*, _l. c._,

parasphenoid.

A large portion of the base of the cranium is taken up by this

cruciform bone. Of the two longitudinal median processes, the

posterior is by far the shorter, and lies in front of and partly

below the cartilaginous _os occipitale basilare_. The anterior longer

longitudinal arm closes in the greater part of the cranium from below,

and articulates by its outer edges with that part of the prootic bones

often described as the _alae magnae_, and also with the cartilage

lying in front, which forms the greater part of the lateral walls of

the cranium. The anterior extremity of the bone articulates with the

palatine bones. The transverse arms lie on the under surface of the

exoccipitals and of the prootics.

The greater width of the cranium in _R. temporaria_ is associated with

the greater relative width of the anterior arm of this bone.

4. The *fronto-parietal bones*, _ossa fronto-parietalia_, Cuvier

(Figs. 10 and 14 _fp_).

*Cuvier*, _l. c._, p. 387, Pl. XXIV, c. c.--*Dugès*, n.

1.--*Parker* and *Bettany*, _l. c._, fronto-parietal.

These are a pair of somewhat long, flat bones, which form the principal

part of the upper wall or roof of the cranium, and cover in superiorly

the cartilaginous cranium, which is here, in great part, persistent.

They are united in the middle line by the sagittal suture; posteriorly

they articulate with the exoccipital and prootic bones; anteriorly with

the sphenethmoid, which they overlap like tiles. The outer margin of

each bone is bent somewhat downwards (Fig. 16 _fp_), and between it and

the parasphenoid there is a space in the wall of the cranium which is

closed in by cartilage and connective tissue only.

These bones are narrower in _R. esculenta_, and along the sagittal

suture are depressed into a groove: where the superior surface bends

down to become lateral the edges are much more prominent. In _R.

temporaria_ the bones are broader and flat or even somewhat arched. The

latter condition is still more marked in _R. oxyrhinus_.

*5.* The *sphenethmoid*, _os ethmoideum_ (Figs. 10, 11, 14, and 16

_e_).

*Cuvier*, os en ceinture, _l. c._, p. 387, Pl. XXIV, a.--*Dugès*,

n. 15.--*Rathke*, anterior or sphenoidal wing (Vortr. z. vergl.

Anat. d. Wirbelthiere, Leipzig, 1862, p. 42).--*Meckel*, Riechbein,

_l. c._, p. 502.--*Parker* and *Bettany*, _l. c._, ethmoid.

The long tubular cranium is completed anteriorly by a single bone,

which forms at once the roof, floor, and lateral walls. It is

consequently more or less ring-shaped, on which account it has been

named ‘os en ceinture’ by Cuvier. Only the posterior portion is

annular, however: the anterior portion forms a double canal, with a

median partition, for the passage of the nerves of smell, and as these

canals are widened out anteriorly, this part of the bone helps to

complete the nasal cavities, which, however, are bounded for the most

part by cartilage, as described below. In some species of frogs (as for

example _R. occellata_, Rathke) this cartilage is partly ossified.

The sphenethmoid has on each side a small bony canal, running forwards

and inwards, through which the _ramus nasalis_ of the first division of

the trigeminal nerve passes.

The *cartilaginous skeleton of the nose* (Figs. 14 and 16 _n_, _n″_,

_n‴_, _n⁗_). The anterior borders of the funnel-shaped cavities of

the sphenethmoid pass into cartilage, which forms two capsules,

separated from each other by a median cartilaginous septum, and opening

laterally. We can distinguish, (_a_) a cartilaginous septum, forming

a continuation of the bony one; (_b_) the floor of the nasal cavity,

narrower behind, wider in front; (_c_) a roof somewhat narrower

than the floor. The floor and roof are united in front by an arched

surface. From this cartilaginous capsule various processes project,

which unite it to other portions of the facial skeleton: firstly, from

the most posterior portion of the capsule there passes transversely

outwards a bar of cartilage (_sn_), which, widening, becomes continuous

with the cartilage (_sp″_) forming the basis of the anterior arm of

the pterygoid bone. From the anterior angle a cartilaginous process

passes outwards (Figs. 14, 16 _n″_), which is attached to a projection

on the anterior end of the maxillary bone; from the same spot a

horn-shaped cartilaginous process (_n″_, _n‴_) curves round backwards

and towards the middle line. This projection bounds the nasal cavity

externally, sending off a free process (_n‴_) on the way, and ends with

a double point on the roof of the cartilaginous nasal cavity (_n⁗_).

On the floor of the nasal capsule, on either side, is a knob-shaped

cartilaginous eminence, running from behind, forwards and outwards;

this may be regarded as an indication of a turbinated bone. From the

anterior wall there passes into each nasal cavity an almost horizontal,

partly ossified plate (_c.n._), ending posteriorly in a free pointed

border. These are the _cornets_ of Dugès (_l. c._, p. 12, Pl. I,

Figs. 1, 5), who correctly regarded them as turbinated bones. Cuvier

described them as rudimentary nasal bones[29].

[Footnote 29: Cuvier (Ossem. fossil., V, 2. 388) says that these little

bones are fixed outside the nasal cavity, so that we may suppose that

he confounded them with the nasal cartilages found in this situation.

Dugès correctly indicates their position to be inside of the nasal

capsule. Meckel (Vergl. Anat. II, 504) also regards them as nasal

bones. Bruch (Würzb. naturwiss. Zeitschrift, vol. II, 1861, p. 213)

could not find them in any frog-skull, doubtless because he did not

look inside the nasal capsules.]

[Illustration: Fig. 14.

Cartilaginous basis of the skull of _Rana esculenta_, from above, twice

natural size. Cartilage shown by stippling.

_a.n._ Wings of nasal cartilage.

_c.n._ Concha narium.

_e_ Sphenethmoid.

_ff′_ Foramina of cranium.

_fn_ Nasal.

_fp_ Fronto-parietal.

_i_ Premaxillary.

_j_ Quadrato-jugal.

_m_ Maxillary.

_n_, _sn_ Cartilaginous nasal skeleton.

_n′_, _n″_, _n‴_ Processes of the cartilage.

_o_ Exoccipital bones.

_p_. Prootic.

_pt_ Anterior arm of pterygoid.

_pt′_ Posterior arm of pterygoid.

_s′_ Cartilaginous cranium.

_sp_ Cartilage of suspensorium.

_sp′_ Cartilage under the squamosal.

_sp″_ Cartilage under the pterygoid.

_t_^1 Anterior }

_t_^2 Middle } arm of the squamosal.

_t_^3 Posterior }

]

The alar cartilages of the nose have still to be described. They are

(_a.n._) shell-shaped cartilages, hollowed out on the inner sides and

decreasing in width from before backwards. They are movably attached

by their anterior broader ends to the tips of the projecting portions

of the premaxillary bones, and are so arranged as to bound the lateral

margins of the nasal apertures, which they overlap.

[Illustration: Fig. 15.

Nasal cartilages of frog, front view.

_an_ Concha narium.

_i_ Premaxillary bones.

_i′_ Ascending process of same.

_m_ Upper jaw.

_m′_ Lower jaw.

_n_ Olfactory capsule.

]

*6.* The *cartilaginous basis of the skull*, _Primordial cranium_

(Figs. 14, 16).

*a.* If the fronto-parietals be removed, (Figs. 14, 16), a cartilage

(_s′_) is found which partially closes the cranial cavity, and which is

usually perforated on either side by a foramen (_f′_). More anteriorly

in the median line is found a space (_f_) which extends to the

sphenethmoid bone, and is only closed by connective tissue: posteriorly

the cartilage extends between the exoccipitals as far as the _foramen

magnum_, representing the _os occipitale superius_.

*b.* The base of the cranium is (Fig. 16) also partly cartilaginous. If

the parasphenoid (_s_) be removed we find above it a cartilage (_s′_)

which passes backwards between the exoccipitals, and extends as far

back as the _foramen magnum_: it corresponds to the _os occipitale

basilare_.

[Illustration: Fig. 16

Cartilaginous skull of _Rana esculenta_, from below, twice natural

size. Cartilage shown by stippling.

_c_ Cartilage of prootic.

_e_ Sphenethmoid.

_fp_ Fronto-parietal.

_i_ Premaxillary.

_j_ Quadrato-jugal.

_m_ Maxillary.

_n_, _sn_ Cartilaginous nasal skeleton.

_n′_, _n″_, _n‴_, _n⁗_ Processes of nasal cartilage.

_o_ Exoccipital.

_p_ Prootic.

_pl_ Palatine.

_pt_ Pterygoid.

_pt′_ Articular surface for pterygoid.

_r_ Optic foramen.

_r′_ Foramen for N. abducens.

_s_ Parasphenoid.

_s′_ Cartilaginous cranium.

_sp_ Cartilage of suspensorium.

_sp′_ Cartilage under squamosal.

_sp″_ Cartilage under pterygoid.

_v_ Vomer.

]

*c.* The sides of the cranium are formed almost entirely of cartilage.

This cartilage fills the space bounded by the nasals above, the

parasphenoid below, the so-called _ala magna_ of the prootic bone

behind, and the sphenethmoid in front; and has in its posterior part

an aperture (_r_) through which passes the _nervus opticus_, and below

this a smaller one (_r′_) for the _nervus abducens_. The extent of

the cartilage in the antero-posterior direction is greater in _R.

esculenta_ than in _R. temporaria_, or in other words the sphenethmoid

stretches further back in the latter than in the former.

B. THE BONES OF THE FACE.

[Illustration: Fig. 17.

Cartilaginous basis of the skull of _Rana esculenta_, from above, twice

natural size. Cartilage shown by stippling.

_a.n._ Wings of nasal cartilage.

_c.n._ Concha narium.

_e_ Sphenethmoid.

_ff′_ Foramina of cranium.

_fn_ Nasal.

_fp_ Fronto-parietal.

_i_ Premaxillary.

_j_ Quadrato-jugal.

_m_ Maxillary.

_n_, _sn_ Cartilaginous nasal skeleton.

_n′_, _n″_, _n‴_ Processes of the cartilage.

_o_ Exoccipital bones.

_p._ Prootic.

_pt_ Anterior arm of pterygoid.

_pt′_ Posterior arm of pterygoid.

_s′_ Cartilaginous cranium.

_sp_ Cartilage of suspensorium.

_sp′_ Cartilage under the squamosal.

_sp″_ Cartilage under the pterygoid.

_t^1_ Anterior }

_t^2_ Middle } arm of the squamosal.

_t^3_ Posterior}

]

The *Suspensorium*.

7. The *squamosal bones*, _ossa tympanica_, Cuvier (Figs. 10, 14 _t_).

*Cuvier*, _oss. foss._, V, 2. 390, Pl. XXIV, Figs. 1, 2

n.--*Dugès*, _temporo-mastoidien_, n. 10.--*Meckel*, Articular part

of temporal.--*Hallmann* and others, quadrate-bone.--*Parker* and

*Bettany*, _l. c._, squamosal.

The suspensorium, which forms the articulation between the cranium and

the lower jaw, is T shaped; and consists of cartilage covered by bone.

Of the three arms of the T the anterior has a free pointed extremity

(Fig. 14 _t^1_) at the postero-lateral margin of the temporo-orbital

fossa: it forms a true _processus zygomaticus_, and is bound to the

upper jaw by ligaments. In _R. temporaria_ it is comparatively much

shorter than in _R. esculenta_. The posterior upper arm (_t^2_)

articulates with the prootic bone: the third or lower arm (_t_^3),

supported by a cartilage[30], runs backwards and downwards to unite

with the side of the broad hinder extremity of the quadrato-jugal bone.

[Footnote 30: This cartilage is the quadrate cartilage of Parker;

others regard it as the upper end of the cartilaginous mandibular arch,

of which Meckel’s cartilage is the lower distal part.]

The cartilaginous basis[30] of the suspensorium (Fig. 17 _sp_, _sp′_)

is a direct continuation of the cartilaginous basis of the cranium,

and proceeds from it to the outer extremity of the prootic; from this

point the cartilage runs backwards and divides into two branches; of

these, one (_sp′_) retains the original course backwards and outwards

towards the articulation of the lower jaw, and in conjunction with the

quadrato-jugal (_j_) forms this joint. The other (_sp″_) is directed

forwards along the anterior arm of the pterygoid bone; it widens in

front (_sn_), and becomes directly continuous with the cartilaginous

strut described above as projecting from the hinder part of the

cartilaginous olfactory capsule.

*8.* The *quadrato-jugal bones*, _ossa jugalia_, Cuvier (Figs. 10,

17, 19 _j_).

*Dugès*, _tympano-malléal_, n. 11.--*Cuvier*, _l. c._, p. 399,

Pl. XXIV, Figs. 1 2 o.--*Hallmann*, quadrato-jugal bone, _l. c._,

p. 39.--*Parker* and *Bettany*, _l. c._, quadrato-jugal.

This is a small process of bone, likened by Dugès to a comma, which

rests by its posterior broader extremity on the cartilage of the

mandibular arch[1], and forms with it a portion of the articular

head for the lower jaw. The anterior extremity is pointed, directed

forwards, and attached by ligament to the maxillary bone.

*9.* The *pterygoid bones*, _ossa pterygoidea_ (Figs. 10, 11, 17, 19

_pt_).

*Cuvier*, _l. c._, p. 389, Pl. XXIV, Figs. 1, 2 m m.--*Dugès*, n.

9.--*Parker* and *Bettany*, _l. c._, pterygoid bones.

These bones, shaped like the Greek letter λ, possess each three arms:

of these the middle one is attached to the under surface of the prootic

bone by a cartilaginous process (Fig. 18 _sp_); the hinder grooved arm

lies upon the under surface of the cartilaginous suspensorial arch

(Fig. 18 _sp′_); and the anterior arm (_pt_) runs forwards along the

inner border of the anterior ramus of the suspensorium (_sp″_), in

contact with the maxillary, palatine, and nasal bones.

The cartilage just mentioned, by means of which the middle arm of the

pterygoid is connected with the cranium (compare Fig. 18), proceeds

from the cranial origin of the cartilaginous arch (_sp_), turns

downwards and then forms a sort of articular head (_cpt_), which is

articulated to an oval flat elevation of the prootic cartilage by a

movable joint. Externally the articular head curves somewhat abruptly

into the bony substance (Fig. 18 _pt_) of the pterygoid bone.

[Illustration: Fig. 18.

Origin of the suspensory cartilage (_sp_) from the skull, with

attachment of the pterygoid; front view. Cartilage shown by stippling.

_cpt_ Cartilaginous epiphyses of the pterygoid bone, from the cartilaginous suspensorium.

_j_ Quadrato-jugal bone.

_pt_ Pterygoid bone.

_sp_ Origin of suspensorial cartilage from the skull.

_sp′_ Hinder arm of same.

_sp″_ Anterior arm of same.

]

[Illustration: Fig. 19.

Cartilaginous skull of _Rana esculenta_, from below, twice natural

size. Cartilage shown by stippling.

_c_ Cartilage of prootic.

_e_ Sphenethmoid.

_fp_ Fronto-parietal.

_i_ Premaxillary.

_j_ Quadrato-jugal.

_m_ Maxillary.

_n_, _sn_ Cartilaginous nasal skeleton.

_n′_, _n″_, _n‴_, _n⁗_ Processes of nasal cartilage.

_o_ Exoccipital.

_p_ Prootic.

_pl_ Palatine.

_pt_ Pterygoid.

_pt′_ Articular surface for pterygoid.

_r_ Optic foramen.

_r′_ Foramen for N. abducens.

_s_ Parasphenoid.

_s′_ Cartilaginous cranium.

_sp_ Cartilage of suspensorium.

_sp′_ Cartilage under squamosal.

_sp″_ Cartilage under pterygoid.

_v_ Vomer.

]

*10.* The *maxillary bones*, _ossa maxillaria superiora_, Cuvier

(Figs. 10, 11, 17, 19 _m_).

*Cuvier*, _l. c._, p. 389, Pl. XXIV, Figs. 1, 2 k k.--*Dugès*, n.

4, _maxillo-jugal_.--*Parker* and *Bettany*, _l. c._, maxillary

bones.

These are elongated, curved bones, narrower behind, broader in front,

and forming with the quadrato-jugals the bony support of the face. The

superior surface is convex and directed outwards, the inferior surface

is grooved and bears teeth along the outer edge of this groove. From

the upper and inner border of the anterior half of this bone a process

(_processus frontalis_) passes upwards, which articulates with the

nasal bone. The anterior extremity of the bone articulates with the

premaxillary bone, the posterior with the quadrato-jugal.

The maxillary bone articulates at its anterior end (Fig. 19 _n″_) with

a process of the cartilaginous capsule of the nose. The broad plate of

cartilage (Fig. 19 _n_), under cover of the _processus frontalis_,

connects the hindermost transverse portion of the nasal cartilage

with the cartilage which runs forwards from the suspensorium upon the

anterior arm of the pterygoid.

*11.* The *nasal bones*, _ossa fronto-nasalia_, Dugès (Figs. 10, 17

_fn_).

*Cuvier*, _frontale anterius_, _l. c._, h.--*Dugès*, n.

2.--*Meckel*, nasal bone.--*Parker* and *Bettany*, _l. c._, nasal

bones.

These flat, triangular bones, which rest upon the nasal cartilages

(_n_), assist to bound the orbital cavities in front, and are connected

by their external angles with the maxillary bone on either side[31].

[Footnote 31: For _os nasale_, Cuvier (_l. c._, g), _cornet_

(_concha_), Dugès, see cartilaginous nasal capsule, p. 27.]

*12.* The *premaxillary bones*, _ossa intermaxillaria_ (Figs. 10, 11,

17, 19 _i_).

*Cuvier*, _l. c._, p. 388, Pl. XXIV, Figs. 1, 2 f f.--*Dugès*, n.

3.--*Parker* and *Bettany*, _l. c._, premaxillary bones.

These two bones are united in the middle line: each consists of a

tooth-supporting portion, which completes the maxillary arch, and

an ascending process, to the apex of which the nasal cartilage is

articulated by a movable joint (_an_). As muscles are attached to this

process, it becomes a lever, by which movements are effected.

[Illustration: Fig. 20.

Skull of _Rana esculenta_, seen from below, twice natural size.

_c_ Cartilaginous wall of skull.

_e_ Sphenethmoid.

_e′_ Cartilaginous nasal skeleton.

_h′_ Stylo-hyoid.

_i_ Premaxillary.

_m_ Maxillary.

_m′_ Quadrate tract.

_o_ Exoccipital.

_p_ Prootic.

_p′_ Anterior arm of prootic.

_p″_ Trigeminal foramen.

_pl_ Palatine.

_pt_ Anterior arm of pterygoid.

_pt′_ Posterior arm of pterygoid.

_s_ Parasphenoid.

_v_ Vomer.

]

*13.* The *palatine bones*, _ossa palatina_ (Figs. 19, 20 _pl_).

*Cuvier*, _l. c._, p. 389, Pl. XXIV, Figs. 1, 2 i i.--*Dugès*, n.

7.--*Parker* and *Bettany*, _l. c._, palatine bones.

Are two transverse bony bars, situated under the anterior portion of

the sphenethmoid bone on either side: each extends outwards from the

apex of the parasphenoid to the maxillary bone, where this gives off

its ascending process to the nasal bone.

*14.* The *vomers*, _vomer_ (Figs. 19, 20 _v_).

*Cuvier*, _l.c._, p. 389, Pl. XXIV, Figs. 1, 2 l l.--*Dugès*, n.

6.--*Parker* and *Bettany*, _l.c._, vomer.

A pair of flat bones, placed horizontally on the under surface of

the cartilaginous nasal capsule in the triangular space between the

palatine bone and the anterior portion of the arch of the jaw. They

limit the outer border of the floor of this capsule, which lies in a

groove on the dorsal surface of the bone. The outer border of each bone

has three processes, and between them two notches; the hinder of these

notches bounding the posterior nares. The inner borders of the two

bones touch, and the under surface of each bears a transverse row of

small, pointed teeth.

*15.* The *mandible*, _maxilla inferior_ (Fig. 21).

*Cuvier*, _l.c._, p. 398, Pl. XXIV, Figs. 1, 2 _rst_.--*Dugès*, n.

16, 17, 18, 19.--*Parker* and *Bettany*, _l.c._, mandibular arch.

[Illustration: Fig. 21.

_A_ Mandible of _Rana esculenta_, from above, nat. size.

_B_ External view of left side, } twice nat. size.

_C_ Angulosplenial, }

1. Angulosplenial.

2. Dentary.

3. Mentomeckelian.

4, 4′ Meckel’s cartilage.

_a._ Rudimentary _processus coronoideus_.

_b._ Groove for Meckel’s cartilage.

]

The mandible consists of two bony arches united by ligament in the

middle line, and each formed of four pieces. These are, (1) the

*angulosplenial*, a strong, bony piece (Fig. 21 _C_, _B_1, _A_1),

which forms the inferior and internal part of the lower jaw; below,

it extends posteriorly, as far as the articulation; in front, it is

attached to the dentary. In the greater part of its length it has

a groove directed upwards and outwards (_C_, _b_), and above has a

process which represents the _processus coronoideus_: (2) the *dentary*

(_A_ 2, _B_ 2), a bony lamella which rests on the outer surface of

the anterior half of the angulosplenial, and articulates in front with

*Meckel’s cartilage*; it is a thin, flat lamella of bone; (3) in the

groove of the angulosplenial is found a cartilage occupying its entire

length (_A_ 4, _B_ 4), *Meckel’s cartilage*; it is broader behind,

and here forms the articular fossa of the lower jaw; anteriorly it is

covered by the dentary, and articulates with the (4) *mentomeckelian*

(_A_ 3, _B_ 3) at its anterior end, which is bound to its fellow of the

opposite side by ligaments, and unites the two halves of the lower jaw.

The upper part of Meckel’s cartilage, which forms the joint with the

suspensorium, must be regarded as the _articulare_; to this also

belongs the anterior, ossified portion of the mentomeckelian[32];

the dentary, which covers Meckel’s cartilage, must represent the _os

dentale_[33]; the first and largest bone, the angulosplenial, which

lies on the inner side of the cartilage, must be considered as the _os

operculo-angulare_, Dugès, for it contains the elements of these bones.

[Footnote 32: Dugès, _os dentale_.]

[Footnote 33: Dugès, _surangulaire_.]

[Illustration: Fig. 22.

Hyoid of _Rana esculenta_, twice nat. size.

_H_ Body; _h h′_ processes.

_H′_ Anterior horn.

_H″_ Posterior horn.

]

*16.* The *hyoid bone*, _os hyoideum_ (Fig. 22).

*Parker* and *Bettany*, _l.c._, hyoid.

This consists of a broad, thin, cartilaginous plate, with processes

passing from it as represented in the figure. We distinguish in

it:--(*a*) the body (_H_), somewhat oblong, and wider in front than

behind. From the anterior angles there projects on either side a blunt

rounded process (_h_); from either posterior angle a more pointed

process (_h′_). (*b*) The anterior cornua (_H′_) (_cornua styloidea_)

project from the anterior border of the body, first in a forward

direction, then in a curve backwards and upwards (compare Fig. 12),

to unite with the cartilage of the prootic bone. Where the direction

of these _cornua_ is changed each sends off a short blunt process.

(*c*) The posterior cornua (_cornua thyroidea_) (_H″_) are bony rods

which run from the hinder margin of the body, diverge posteriorly, and

enclose the larynx.

III. THE STERNUM (Fig. 24 _es_, _hs_).

*Cuvier*, _l. c._, p. 401, Pl. XXIV, Fig. 31 _e f_.--*Dugès*, n.

27, 28.--*Parker* and *Bettany*, _l. c._, sternum.

The bones of the sternum are in very close relationship with the

shoulder-girdle, and some difference of opinion exists as to the

relations of the several parts.

[Illustration: Fig. 23.

Omosternum of _Rana esculenta_, twice nat. size.

_a_ Anterior,

_p_ Posterior extremity.

]

The following parts must be regarded as undoubtedly belonging to the

sternum:--

*1.* An anterior part, consisting of a bony and a cartilaginous portion.

_a._ The *omosternum* (Figs. 23, 24 _e.s._) is a bony rod, wider

behind, narrower in front, and supports

_b._ The *episternum* (Fig. 24 _es′_), a semicircular plate of

calcified cartilage, with a thin free border of unchanged hyaline

cartilage.

*2.* A posterior part, which also consists of a bony and a

cartilaginous portion.

[Illustration: Fig. 24.

Shoulder-girdle and sternum of _Rana esculenta_, twice nat. size. The

scapula and suprascapula are turned outwards.

_c._ Connecting cartilage.

_cl._ Clavicles and precoracoids.

_co_ Coracoid.

_es._ Omosternum.

_es′_ Episternum.

_h_ Hyaline cartilage.

_hs._ Sternum proper.

_hs′_ Xiphisternum.

_o_ Bone.

_o′_ Calcified cartilage.

_sc._ Scapula.

_sc′_ Suprascapula.

]

_a._ The *sternum proper* (Fig. 24 _h.s._), a rod of cartilage

ensheathed in bone, broader in front and narrower behind. To its

posterior extremity is attached

_b._ The cartilaginous *xiphisternum* (Fig. 24 _hs′_), a plate, the

shape of which is best understood by referring to Fig. 24, and the

structure of which is similar to that of the episternum.

*3.* The *epicoracoids*[34] lie between the episternum and the sternum

proper: they are a pair of narrow cartilages, closely applied to each

other and placed between the ventral ends of the precoracoids and

coracoids, see p. 40.

[Footnote 34: Dugès, _l.c._, p. 61, regards the cartilaginous laminae

as _os episternale_ and _proc. xiphoideus_, and the two bones _e.s._

and _h.s._ (Fig. 24) as sternum.]

IV. THE BONES OF THE ANTERIOR EXTREMITY.

A. BONES OF THE SHOULDER-GIRDLE.

The shoulder-girdle of the frog (Fig. 25) consists of four bones, two

in the upper or *scapular portion*, and two in the lower or *coracoid

portion*.

*1.* The *scapular portion*.

*a.* The *suprascapula*, _pars suprascapularis scapulae_, _scapula

superior s. omolita_ (Figs. 25 _s′_, 26, 27). This forms the upper,

thinner portion. It is trapezoid in shape, its narrowest, thickest,

and lowest border articulating with the scapula. The remaining

three borders are free; the posterior and superior borders are very

considerably thinned out, the inferior surface is curved towards the

ventral surface. Only the thin upper and hinder borders (_hh_) are of

hyaline cartilage; the middle portion (_co_) consists of calcified

cartilage; the lateral part (_o_) and the anterior border (_o′_) of

true bone.

[Illustration: Fig. 25.

Shoulder-girdle of the frog in transverse section, diagrammatic.

_c_ Connecting cartilage.

_c′_ Epicoracoids.

_cl_ Clavicles.

_h_ Humerus.

_s_ Scapula.

_s′_ Suprascapula.

_ss_ Articulation of these two.

_v_ Vertebra.

]

The bony part forms a thin plate striated both radially and

concentrically with the margin. The anterior border (_o′_) is bent on

itself so as to form a groove (Fig. 27). The dorsal surface of this

bony plate is quite smooth, and covered only by periosteum; the ventral

surface, on the contrary, is for the most part covered by an even layer

of calcified cartilage (Fig. 27 _co′_). This passes directly into the

second portion of calcified cartilage (_co_); so that if the whole be

dried, the thin hard lamina of bone can easily be split away from the

underlying parchment-like cartilage. The layer of calcified cartilage

is, however, not so complete that the _suprascapula_ can be said to

consist of calcified cartilage.

[Illustration: Fig. 26.

Suprascapula of _Rana esculenta_, from above, twice nat. size.

_co_ Calcified cartilage.

_h_ Hyaline cartilage.

_o_ Bony portion.

_o′_ Anterior curved border.

]

[Illustration: Fig. 27.

Suprascapula of _Rana esculenta_, from below, twice nat. size.

_co_ Calcified cartilage.

_co′_ Layer of calcified cartilage on the under surface of the bone.

_h_ Hyaline cartilage.

_o_ Bony portion.

_o′_ Anterior curved border.

]

[Illustration: Fig. 28.

Left scapula of _Rana esculenta_, from below, twice nat. size.

_a_ Anterior border.

_c.a._ _Cavitas articularis._

_l_ Outer border.

_m_ Inner border.

_p_ Posterior border.

]

[Illustration: Fig. 29.

Scapula seen from behind, twice nat. size.

_c.a._ _Cav. articularis._

_d_ Dorsal process.

_v_ Ventral process.

]

*b.* The *scapula* (Figs. 25 _s_, 28, 29) is a long, flat,

four-cornered plate of compact bone, contracted in the middle, and

wider at either end. From above (Fig. 28 _l_) it descends to the

shoulder-joint with a slight curvature, with the convexity directed

outwards. The inferior or ventral border (Fig. 28 _m_), which assists

in the formation of the shoulder-joint, is cleft into two processes,

a ventral and a dorsal. This may be best seen by looking at the

border of the bone from behind or before (as in Fig. 29). The lower

ventral process (Fig. 29 _v_) may be regarded as the _acromion_, or

at least as a rudiment of the acromion, _i. e._, _spina scapulae_.

The upper dorsal process evidently represents a rudimentary _processus

coracoideus_. The anterior border (Fig. 28 _a_) is free, somewhat

sharp, and slightly concave; the posterior (Fig. 28 _p_), also concave,

is free externally; while internally, where the division of the ventral

border commences, it takes part in the formation of the articular

cavity of the shoulder (Figs. 28 and 29 _c.a._).

_Articulation with the suprascapula._ The scapula consists of a shell

of compact bony tissue and an inner portion of cancellous tissue; this

latter passes, at the boundary between this bone and the suprascapula,

through an intermediate layer of calcified cartilage, which is directly

continuous with the cartilage of the suprascapula. The compact

superficial layer ceases where the calcified cartilage begins, and the

periosteum of the scapula passes directly into the perichondrium of the

suprascapula.

*2.* The *ventral portion* of the shoulder-girdle consists of three

parts, which lie one behind the other.

*a.* The larger and posterior bone, the *coracoid* (Figs. 24 _co_,

30), is contracted in the middle, and expanded at either end,

especially at the sternal end. The inner half is flattened from above

downwards; externally it becomes more cylindrical. The anterior and

posterior borders are markedly concave; the outer border (Fig. 30 _l_)

articulates by means of a cartilage with the _proc. coracoideus_ of

the scapula; the inner border (_m_) is separated from its fellow of

the opposite side by the epicoracoids. This inner border is so broad,

that it almost touches the inner end of the clavicle (Fig. 24 _cl._). A

*coracoid foramen* is thus formed between the two bones, similar to the

_foramen ovale_ of the pelvis.

[Illustration: Fig. 30.

Left coracoid, twice nat. size.

_a_ From below.

_b_ From behind.

_l_ Outer extremity.

_m_ Inner extremity.

]

[Illustration: Fig. 31.

Clavicle of the left side, twice nat. size.

_l_ Outer extremity.

_m_ Inner extremity.

]

*b.* The small anterior bones of the ventral portion of the

shoulder-girdle (Figs. 24 _cl._, 31), I, like Cuvier and others,

regard as *clavicles*. Dugès (n. 33) names them ‘_acromial_.’ Each

bone is narrower than the coracoid, smaller internally (_m_), broader

externally (_l_), and articulates by this broader end with the

cartilage uniting the dorsal and ventral portions; the inner extremity

rests upon the epicoracoid. The clavicle is grooved on its superior

and posterior surfaces in almost its whole length; the groove widens

externally (_l_), and contains

*c.* The *precoracoid* (Fig. 34 _cl′_), a slender bar of cartilage,

connecting the anterior edge of the scapula with the sternum.

[Illustration: Fig. 32.

Right shoulder-girdle of _Rana esculenta_, from above.

_ca_ Cav. articularis.

_c c_ Cartilage.

_cl_ Clavicle.

_co_ Coracoid.

_sc_ Scapula.

_sc′_ Suprascapula.

]

[Illustration: Fig. 33

Hinder border of the scapula and coracoid, with the connecting

cartilage and shoulder-joint, twice nat. size.

_c_ Connecting cartilage.

_d_ Dorsal process.

_co_ Coracoid.

_sc_ Scapula.

_v_ Ventral process.

]

[Illustration: Fig. 34.

Clavicular cartilage of a young specimen of _Rana esculenta_, enlarged

to the adult size.

_c_ Connecting cartilage.

_e_ Epicoracoids.

_c′_ Ligament between the two cartilages.

_cl_ Clavicle.

_cl′_ Clavicle and precoracoid.

_co_ Coracoid.

_co′_ Cartilage of same.

_es_ Omosternum.

_hs_ Sternum proper.

]

*Cartilages* of the shoulder-girdle (Figs. 24, 32, 34). A cartilage

(_c_), described by Dugès as the ‘paraglénal,’ connects the scapula

with the coracoid, and converts the notch between the two processes

(_s_ and _v_) on the inner border of the scapula into an aperture

opening posteriorly into the glenoid cavity, and which is filled up by

the lining cartilage. This cartilage connects the two processes with

the coracoid; being thicker between this and the _proc. coracoideus_,

it takes an important share in the formation of the floor of the

glenoid cavity. At the outer extremity of the clavicle the cartilage

projects and is then continued in the groove on this bone towards

the middle line (Figs. 32, 34), where it widens out, and is united by

its external margin with the sternum, thus forming the precoracoid as

already described, see p. 40. These connections are best seen in young

animals, as is shown in Fig. 34.

The clavicle (_cl_) is of ossifying cartilage; the cartilage (_c_),

the epicoracoid, is very large, and hyaline, and at _cl_ runs directly

into the epicoracoids, which are here united mesially by ligament. In

the fully developed animal the whole of the epicoracoid changes to

calcified cartilage, as also does the precoracoid lying in the groove

of the clavicle.

B. BONES OF THE FORE LIMB.

*1.* The *arm* has but one bone.

[Illustration:

Fig. 35. Fig. 36. Fig. 37.

Humerus of _Rana Humerus of _Rana, Humerus of _Rana

esculenta_, female, esculenta_ male, esculenta_, female,

seen from below. seen from below. outer side.

Figures twice natural size.

_c_ Anterior extremity. _c_′ Posterior extremity. _cd_ Crista

deltoidea. _cm_ Crista medialis. _t_ trochlea.]

The *humerus*, _os humeri_ (Figs. 35, 36, 37). The shaft of the humerus

is, on the whole, cylindrical, the middle part being the narrowest;

its dorsal surface is slightly concave, the ventral surface convex. In

the upper half the cylindrical form is indistinct, from the presence

of a well-marked crest (_crista deltoidea_) upon the under or ventral

surface. The hinder half of the bone has very different shapes in

the two sexes: in the female it remains cylindrical as far as the

elbow-joint; in the male, on the posterior surface, where the _crista

deltoidea_ ceases, a marked ridge (_crista medialis_) arises, which

gradually winds inwards and terminates at the inner condyle. This

sexual difference is found in all the three species described in the

Introduction, and it would seem that this ridge increases in height

during the pairing season. The muscle which arises from this process

is the _flexor carpi radialis_ (Fig. 74 _Fc_), and it attains, in the

males, a very large size at the same period; indeed, it is at its

origin twice as broad as in the female. Dugès[35] has incorrectly

stated that this _crista_ is a peculiarity of _Rana temporaria_, and is

wanting in _Rana esculenta_. Pouchet[36] first observed that it was a

sexual distinction.

[Footnote 35: _l. c._, p. 68.]

[Footnote 36: Compt. rend. vol. XXV, 1847, p. 761.]

The anterior articular extremity presents a rounded head, which is

somewhat flattened externally. The inferior ventral border is sharp,

and developed into the above-mentioned bony crest (_crista deltoidea_);

the free border is arched and curved outwards. Above, on the outer

surface of the head, is found an eminence which may be a rudimentary

_tuberc. maius_. The inner half of the head is articular.

The posterior articular extremity consists of a rounded head (_c′_),

attached to the posterior extremity of the shaft; internal to this is

a small _trochlea_ (_t_), placed upon the inner condyle. The external

condyle is but small. The head is embraced by these condyles. Upon the

under surface, above the head, is the _fossa cubitalis anterior_. The

structure of the two articular extremities of the humerus differs; the

upper, as in long bones generally (see page 16), consists of hyaline

cartilage surrounded by calcified cartilage, which again is covered by

hyaline articular cartilage; the lower extremity is composed of spongy

bone, with roundish marrow spaces, and for the most part covered by a

layer of calcified cartilage.

The *shoulder-joint*. The socket is formed by the posterior border of

the scapula, by both the processes of that bone, the outer portion of

the coracoid, and besides these by cartilage, which forms the articular

surface already described (Fig. 33). In the macerated bone the floor

of the socket opens dorsally into a fossa between the processes of the

scapula (Fig. 32). This opening, in the natural condition, is closed by

synovial membrane only; the cartilage covering the rest of the space

being absent (the sac may easily be inflated, either from the joint

itself or through the opening). The margin of the socket consists of

a _labrum_, partly fibrous and partly cartilaginous, which is easily

removed entire. The capsule, which is attached beneath the _caput

humeri_, arises from this. From the articular cartilage of the ventral

margin of the socket, and from the adjacent part of the scapula,

proceeds a strong band, which is inserted into the flattened outer

surface of the head.

*2.* The *forearm*.

[Illustration: Fig. 38.

Radio-ulnar of the right side of _Rana esculenta_, twice nat. size.

_r_ Radius.

_r_′ Capitulum radii.

_u_ Ulna.

_u_′ Olecranon.

]

The *radio-ulnar*, _os antibrachii_ (Fig. 38). The two bony elements

of the forearm are fused into one, which, however, shows traces of its

compound character. Of these two bones, that which forms the inferior

(in man anterior) border is the radius, the other the ulna. The

relative positions of the bony elements are such that they appear to be

in a state of semi-pronation; by this the thumb-border of the arm is

directed downwards (in man forwards). The two bones so united form a

flattened, tube-like bone, the grooved surfaces of which are directed

inwards and outwards respectively; the borders being dorsal and

ventral. The shaft of the bone is narrower above, becoming gradually

wider below. In the upper half, the groove showing the line of junction

of the two bony elements is scarcely visible, and it appears as

though this part consisted of ulna alone, the radius not reaching so

far; below the groove is more marked. The upper articular extremity

appears to consist of an _olecranon_ (Fig. 38 _u′_) and a _processus

coronoideus_ (_r′_), between which is the greater sigmoid cavity

(_cavitas sigmoidea major_) for the _processus cubitalis humeri_. A

more careful examination shows this to be incorrect: the portion of the

concave articular surface, which lies in the immediate neighbourhood

of the ventral border, undoubtedly represents the articular surface of

the _capitulum radii_, and the upper portion of the articular surface

(at _u′_) belongs to the _ulna_ and articulates with the _trochlea_

(Figs. 35, 36 _t_) of the humerus. The inner border of the upper

extremity of the bone articulates with the groove _t_ of the humerus.

The inferior extremity forms one of the usual cartilaginous epiphyses,

enclosing the two bones. That part of the extremity corresponding with

the radius is triangular and convex, it is prolonged upon the inner or

under surface of the bone by means of a process; the ulnar portion of

the bone forms a _capitulum_. The structure of the inferior extremity

is that of epiphyses in general, except that there are two medullary

cavities in a common sheath of calcified cartilage; the anterior head

consists of cancellous tissue, with a covering of calcified cartilage.

*3.* Bones of the *hand* (Fig. 39).

*Dugès*, Pl. VIII, Figs. 37, 38, 39, 40, 41, 42, 43, 52.

*a.* Bones of the *wrist*, _ossa carpi_. These are arranged in two

rows, proximal and distal. The proximal or anterior row has three

bones, viz.:--

1. _Os lunatum_, the middle bone (Fig. 39 _l_), Dugès, n. 38,

articulates with the radial part of the articular surface of the

radio-ulnar by a concave surface; with the large bone of the second

row (_hc_) it articulates by means of a rounded head; and with the

two other bones of this row, _os naviculare_ (_n_) and _pyramidale_

(_p_), with the former by a saddle-shaped surface, with the latter

chiefly by ligament. The bone is irregularly cuboid; the dorsal and

palmar surfaces are non-articular. With Dugès I regard this bone as _os

lunatum_; Mertens[37] regards it as _os naviculare_.

[Footnote 37: Mertens, _l. c._]

[Illustration: Fig. 39.

Bones of the forearm and hand of _Rana esculenta_, right side, dorsum,

twice nat. size.

_a_ Os antibrachii.

_hc_ Os capitato-hamatum.

_l_ Os lunatum.

_m_ Ossa metacarpi.

_n_ Os naviculare.

_p_ Os pyramidale.

_r_ Radial portion of the radio-ulnar.

_t_ Os multangulum majus or trapezium.

_t′_ Os multangulum minus or trapezoides.

_u_ Ulnar portion of the radio-ulnar.

_I_-_V_ First to fifth fingers, the fifth only

of os metacarpi, the second and third of

two, fourth and fifth of three phalanges.

]

2. _Os pyramidale_ (Fig. 39 _p_), Dugès, n. 37, articulates with the

ulnar part of the posterior articular extremity of the radio-ulnar.

Anteriorly the bone has a facet which receives the _capitulum ulnae_,

and alongside this a small flat surface which articulates with the

radius. Posteriorly, opposite to the second row, it presents a long

raised articular surface, extending from the dorsum to the palmar

surface; and on the palmar aspect of the bone is a ridge running from

before backwards. Mertens names this bone _os lunatum_.

3. _Os naviculare_ (Fig. 39 _n_), Dugès, n. 39. The third bone of the

first row does not articulate with the bones of the forearm. It is

connected with the _os lunatum_ by a saddle-shaped surface, posteriorly

with the _os capitatum_ by means of a concave surface. On the inner

surface is a small concave articular facet for the _os multangulum

minus_ or _trapezoides_ (Fig. 39 _t′_). The convex palmar surface of

the bone articulates with the _os multangulum majus_ (_t_); the radial

side is non-articular, and with the _os lunatum_ forms a ridge directed

forwards. Mertens names this bone _os triquetrum_. It is evident,

however, that if the arrangement in man obtains as a standard, the

nomenclature of Dugès, which I have here used, is correct; but the _os

naviculare_ is no longer in contact with the articular surface of the

_os antibrachii_.

The second or distal row also consists of three bones:--

4. _Os capitato-hamatum_ (Fig. 39 _hc_), Dugès, n. 40, 41, 42,

corresponds with the _os capitatum_ and _hamatum_[38], which are here

fused together. It is the largest bone, and articulates with all three

bones of the first row; it is crescentic, with the concavity directed

posteriorly, and on the convex surface has three facets for the three

outer _ossa metacarpi_.

[Footnote 38: Dugès thinks that _os trapezoides_ is also included.]

5. _Os multangulum minus_, _s. trapezoides_ (Fig. 39 _t′_), Dugès, n.

43, is a small bone lying next to the foregoing; it articulates with

the _os metacarpi II_, and anteriorly with the _os naviculare_. Dugès

considers it to be the _os trapezium_.

6. _Os multangulum majus_, _s. trapezium_ (Fig. 39 _t_), is a small

bone, which is placed on the convex palmar auricular side of the _os

naviculare_. Dugès regards this bone as _os metacarpi pollicis_ (Dugès,

n. 44), Mertens as _multangulum minus_. It appears to me that the

interpretation here given is justified, because it articulates with the

rudiment of the thumb, and as it possesses no resemblance to a long

bone it cannot be regarded as an _os metacarpi_.

*Structure of the carpal bones.* All the carpal bones consist of

calcified cartilage, though variations due to age occur in the amount

of the deposit.

*Wrist joint.* As stated above, the _os lunatum_ and _os pyramidale_

are articulated with the forearm. In addition to the capsules uniting

the parts, there are strengthening ligaments, two of which require

special mention. One of these arises from the palmar surface of the

posterior extremity of the ulna, and passes to the sharp palmar ridge

of the _os pyramidale_, where it is attached, and is inserted into the

_os capitato-hamatum_; the second ligament passes in a similar manner

on the palmar surface from the radius to the _os naviculare_. Between

the two ligaments there is on the volar surface a deep excavation.

[Illustration: Fig. 40.

Bones of the forearm and hand of _Rana esculenta_, right side, dorsum,

twice nat. size.

_a_ Radio-ulnar.

_hc_ Os capitato-hamatum.

_l_ Os lunatum.

_m_ Ossa metacarpi.

_n_ Os naviculare.

_p_ Os pyramidale.

_r_ Radial portion of the radio-ulnar.

_t_ Os multangulum majus or trapezium.

_t′_ Os multangulum minus or trapezoides.

_u_ Ulnar portion of the radio-ulnar.

_I_-_V_ First to fifth fingers, the first only

of a metacarpal, the second and third of

two, fourth and fifth of three phalanges.

]

*Movements* of the hand. The two bones of the forearm being firmly

united, pronation and supination, as in man, are impossible. The

normal position, as already remarked (see radio-ulnar), is that of

semi-pronation (compare Figs. 2 and 74). If the animal be in its

natural posture, with the arm bent at the elbow (Fig. 1) it can touch

the ground with the ulnar border only; but when the whole surface of

the hand rests upon the ground, the action is brought about as follows:

the plane of the carpus is in extension parallel with that of the

forearm; in flexion, almost at right-angles with that plane (Fig. 74)

The _os lunatum_ glides on the prolongation of the articular surface

on the inner palmar surface of the radius, while the _os pyramidale_

is held firmly by the ligament just described. The hand follows the

movement of the _os lunatum_, and at the same time the three outer

_ossa metacarpi_ gliding with it inwards, the fingers of the hand turn

inwards.

*b.* The Bones of the *metacarpus*, _ossa metacarpi_ (Fig. 40 _m_).

Of these there are five: the *second* to the *fifth* are of about

equal length; the first, forming the rudiment of the thumb, hidden

beneath the skin, is much shorter. This metacarpal bone of the *thumb*

is regarded by Dugès and others as the first phalanx. My reason for

describing it as a metacarpal has already been given. This bone

also presents differences in the two sexes. Meckel’s[39] observation

that it exists only in the male is incorrect; it is a large, broad

and sickle-shaped mass of calcified cartilage or even true bone,

while in the female it is simply a small cartilaginous rod; as this

almost entirely disappears in dried specimens, Meckel’s statement is

explained. The *second* finger has in the frog assumed the functions

of the thumb, and, in the males, in the breeding season swells and

undergoes an essential alteration of structure. The metacarpal of

this finger also displays differences in the sexes, being in the male

stronger, broader, and on the inner side provided with a spine-like

prolongation, for the tendon of the _abductor digiti II (pollicis)

longus_, which attains an unusual development during the breeding

season. The remaining metacarpal bones, the *third*, *fourth*, and

*fifth*, are of similar form, the last being provided with a tubercle.

The anterior articular extremities are rather broad and concave, and

the posterior have rounded heads.

[Footnote 39: _l. c._, II, 464.]

*c.* The *phalanges*, _phalanges digitorum_. The hand has four fingers

and the rudiment of a fifth, which latter is completely hidden under

the skin, and as already explained, consists of a metacarpal only. Of

the four fingers, the fourth is the longest, the third the shortest,

the second and fifth of intermediate and about equal length; the fourth

and fifth fingers have each three phalanges, the second and third have

each two. Their shape is not unlike that of the human phalanges.

V. THE BONES OF THE HINDER EXTREMITY.

A. BONES OF THE PELVIC GIRDLE (Fig. 41).

The *pelvis* has a characteristic V-shape, the pubic and ischiatic

bones of the two sides together forming a vertical plate (_a p_), which

divides anteriorly into the two iliac bones (_il_). The pubic and

ischiatic bones being united by their median surfaces, one can only

speak of the pelvic cavity as the space between the two ilia.

*1.* The *ilia*, _ossa ilii_ (Fig. 42 _il_). In these a broader part

or body can be distinguished, which takes part in the formation of the

_acetabulum_ (_a_). The hinder and broadest part of the bone is joined

to the corresponding portion of its fellow by ligament; anteriorly the

two diverge, bounding a cone-shaped cavity, the pelvis. The suture, by

which the body of the iliac bone is united with the ischiatic and pubic

bones, runs from above downwards through the middle of the _acetabulum_

(Fig. 42, from _p_ through the _acetabulum_ to _is_). Inferiorly and

in front the united _ilia_ form a prominence (_s_), which represents

the _symphysis oss. pub._ of man. The expanded portions of the ilium

rest directly over the articular cavity by means of a somewhat

constricted part or root. The _ala_ is a sabre-like bone, with the

sharper, more convex border directed upwards; the more concave, thicker

border downwards. The two surfaces, of which the inner is grooved,

are directed respectively inwards and outwards. The broader part of

the blade springs directly from the neck or root (at _il′_), and at

this end of the sharp convex border is a process, _process. sup._,

from which the tendon of the _m. vastus externus_ arises. The anterior

end of the blade is a hollow cylinder, containing cartilage, movably

articulated to the transverse process of the last vertebra or sacrum.

[Illustration: Fig 41.

Pelvis of _Rana esculenta_, from below, nat. size.

_a_ Acetabulum.

_il_ Iliac bone.

_p_ Pubic bone.

]

[Illustration: Fig. 42.

Pelvis of _Rana esculenta_, left side, twice nat. size.

_a_ Acetabulum.

_d_ Upper sharp border.

_il_ Iliac bone.

_il′_ Superior process.

_is_ Ischium.

_is′_ Tuber ischii.

_p_ Os pubis.

_s_ Inferior process of the ilium.

_v_ Lower border of same.

]

*2.* The *ischia*, _ossa ischii_ (_is_), are irregular, four-cornered

plates of bone, united together by the whole extent of their inner

surfaces. The vertical plate formed by the union of the two bones takes

only a very small share in the formation of the articular cavity.

Above, it has a somewhat sharp border, continued with the ilium into

a process (_is′_) corresponding with the united _tubera ischii_; the

hinder border, which is united below with the pubic cartilage, is

curved.

[Illustration: Fig. 43.

Horizontal section through the iliac, pubic, and ischiatic bones.

_a_ Acetabulum.

_il_ Ilium.

_is_ Ischium.

_p_ Pubic cartilage.

_s_ Ligaments which unite the bones of opposite sides.

]

*3.* The *pubes*, _os pubis_ (Figs. 41, 42, 43 _p_). Ossified pubic

bones do not exist; even in old animals they are composed of calcified

cartilage. Each is a triangular cartilage, wedged between the ilium and

ischium, but also taking part in the formation of the _acetabulum_. The

pubes of opposite sides are, as shown in section in Fig. 43, united by

ligaments in the median plane.

B. BONES OF THE HIND-LIMB.

*1.* The *thigh*.

The *femur*, _os femoris_ (Fig. 44). The shaft is cylindrical, with a

slight double or S-shaped curve. In the anterior portion the convexity

is directed upwards, in the two hinder thirds downwards. Towards the

hinder extremity it is flattened from above downwards. The anterior

extremity is a rounded articular head placed directly on the shaft

like that of the _humerus_. This head is somewhat flattened from side

to side; and in structure resembles that of the humerus. The inferior

extremity is rounded below, but flattened above (towards the hollow of

the knee) and behind.

*Hip-joint.* The articular cavity is deepened by a fibrous ring,

_labrum cartilagineum_, which fits closely upon the _caput femoris_;

from this _labrum_ the joint-capsule arises, to be inserted behind the

head; close to the lower border of the cavity a _ligamentum teres_

arises, which is inserted into the _caput femoris_.

*2.* The *leg*.

[Illustration: Fig. 44.

Femur of _Rana esculenta_, right side, twice nat. size.

_a_ Anterior extremity.

_p_ Posterior extremity.

]

*Tibio-fibula*, _os cruris_ (Fig. 45). This consists of a single bone,

the _os cruris_; it is often called the _tibia_, but presents, however,

distinct indications of being formed of two bones, the _tibia_ and

_fibula_. The shaft is not quite straight; it is thinner in the middle

and flattened from above downwards. On the upper as also on the lower

surface of the bone is a groove, passing from the middle towards each

extremity; this deepens as it proceeds, and is an indication of the

compound nature of the bone. In the middle of each surface of the bone

is the opening of a canal, through which a bristle may be passed: the

canal has bony walls. The medullary cavity is, for the most part,

double, being single only in the middle. The anterior extremity is

articular, and presents a median groove, which is prolonged on to the

under surface. The posterior extremity forms a transverse condyle,

which has at its inner part a notch for the tendon of the _m. tibialis

posticus_.

[Illustration: Fig. 45.

Tibio-fibula of _Rana esculenta_, twice nat. size, seen from below.

_a a_ Grooves.

_f_ Foramen nutritium.

_t_ Anterior articular extremity.

_t′_ Posterior articular extremity.

_t″_ Groove on malleolus internus for tendon of _m. tibialis posticus_.

]

The *knee-joint* consists of a capsule strengthened by the insertions

of numerous tendons. Within this the articular surfaces of the _femur_

and tibio-fibula are connected by means of fibro-elastic bands, which

spring from the latter bone and cross each other, thus representing the

_lig. cruciata_. The articular surfaces are moreover deepened by loose

semilunar cartilages, corresponding with the inter-articular cartilages

of the knee-joint of man. In addition, on either side of the joint, are

found auxiliary ligaments.

[Illustration: Fig. 46.

Section of the tibio-fibula through the under extremity.

_a_ Cartilaginous epiphysis.

_m_ Marrow cavities.

_t″_ Connecting mass.

_t t′_ The two cylinders of bone.

]

*3.* The *foot* (Fig. 47).

*a.* The *ankle*, _ossa tarsi_. The *tarsus* consists of two rows of

bones.

α. The *proximal row* consists of two long tubular bones (Fig. 47

_ac_), which are parallel to each other, and undoubtedly correspond

with the *astragalus* (_talus_) and *calcaneum*. The bones are

separated in the middle, but approach each other at either end. The

outer of these bones (_c_) I have named calcaneum, following Cuvier

and Dugès; the inner, astragalus (_a_). The two bones are surrounded

and enclosed at their anterior and posterior extremities in a common

epiphysis of calcified cartilage, which has the usual structure found

elsewhere. The _ligamentum calcanei_, which will be described below

(see Muscles of Foot, Fig. 88 _lc_), may be regarded as an unattached

process of the heel[40].

[Footnote 40: Meckel, _l. c._, II, p. 488, regards the cartilage in the

_tendo Achillis_ as such.]

β. The *distal* row of the tarsals are all composed of calcified

cartilage. They are:--

1. _Os cuboideum_ (Fig. 47 _cb_), a flat plate of cartilage, concave

in front, convex behind, placed between the common epiphysis of the

astragalus and calcaneum (_ac_) and the _ossa metatarsi II_ and _III_.

This corresponds, as Dugès suggests, with the separated _os cuboideum_

and _cuneiforme III_ of other batrachians, _e. g._ Bufo.

2. On the inner side of the above, between the _astragalus_ and

_metatarsus I_ lies a small body, concave in front, convex behind,

which we may regard, with Dugès, as the analogue of the _os naviculare_

(Fig. 47 _n_).

[Illustration: Fig. 47.

Right foot of _Rana esculenta_, palmar surface, twice nat. size.

_a_ Astragalus.

_ac_ Common epiphysis.

_c_ Calcaneum.

_cb_ Os cuboideum.

_h_ First, _h′_ second cartilage of supplemental toe.

_m_ Ossa metacarpi.

_n_ Os naviculare.

_I_ to _V_ First to fifth toe.

]

3. This latter supports a cartilage which forms a projection on the

inner border, covered with skin. This consists of closely-united

pieces, a small round, basal piece (_h_), and a flattened beak-like

piece (_h′_). Dugès considers these pieces as the first and second

_ossa cuneiformia_; I rather incline to Meckel’s opinion that they

represent a rudimentary hallux, the first being the _os metatarsi_, the

second a _phalanx_. Both are composed of calcified cartilage; in _R.

temporaria_ the lime-deposit seems to be less than in _R. oxyrhinus_

and _R. esculenta_, and the projection consequently softer[41].

[Footnote 41: In the text these two cartilages are termed the

‘supplemental toe.’]

*b.* The *metatarsal bones*, _ossa metatarsi_ (Fig. 47, _m_ _I_-_V_).

The five _ossa metatarsi_ are long, thin tubular bones, the anterior

articular extremities of which are broader above and narrow below,

hence are wedge-shaped, and anteriorly present flat surfaces for

articulation with the _tarsus_. The shafts are long and thin; the

posterior extremities have rounded heads. The comparative length of

these is as follows: the fourth is the longest, then come the third,

fifth, second, and first in order. In structure they resemble other

tubular bones.

*c.* The *phalanges*, _phalanges digitorum_ (Fig. 47). The total

lengths of the separate toes correspond with that of the _ossa

metatarsi_, but the distinctive differences are increased, partly by

the difference in number and partly by the varying lengths of the

phalanges. The first and second toes have each only two phalanges, the

third and fifth have each three, the fourth, which is the longest, has

four. The terminal phalanges are somewhat hooked towards the plantar

surface.

SECTION II.

THE MUSCLES.

THE MUSCLES.

LITERATURE.

Anonymous, Ueber das Schultergerüst der Schildkröten mit den

daran sitzenden Muskeln. Isis, 1827. (Contains remarks on the

shoulder-muscles of the frog.)

*Carus*, *C. G.* (1) Lehrbuch der vergleichenden Zootomie. 2nd Edit.

Leipzig, 1834. (2) Erläuterungstafeln zur vergleichenden Anatomie.

1 Part. 1826. (Muscles of the Leg.)

*Chappuis*, Morphologische Stellung der kleinen hintern Kopfmuskeln.

Zeit. f. Anat. u. Entwickl. 1876. Vol. II, pages 287–297.

*Cuvier*, Leçons d’anatomie comparée. Paris, 1835. Vols. I, II.

*Dugès*, Recherches anatomiques et physiologiques sur la déglutition

dans les Reptiles. Ext. des Annales des Sciences naturelles.

Paris, 1827.

*Dugès*, Leçons d’anatomie comparée. Paris, 1835.

*Duméril and Bibron*, Erpétologie générale ou Histoire complète des

Reptiles. 1836.

*Führbringer*, Zur vergleichenden Anatomie d. Schultermuskeln.

Jenaische Zeitschr. 1873. Vol. VII.

*Klein*, Beiträge zur Anatomie der ungeschwänzten Batrachier (Rana

temporaria, L.): in Jahrshefte des Vereins für vaterländische

Naturkunde in Würtemburg. 6 Jahrgang. 1850. Page 1.

*Kuhl*, Beiträge zur Zoologie der Rana esculenta: in Beitr. z. Zool.

Frankfurt, a. M. 1820. Page 115.

*de Mann*, Vergelykende myologische en neurologische studien over

Amphibien en Vogels. Acad. Proefschrift. 1873.

*de Mann*, Myologie comparée de l’extrémité post. chez les

amphibiens. Niederl. arch. f. Zool. 1874. Vol. II.

*Marshall*, The Frog. London and Manchester. 2nd Edit. 1885. Pages

60–71.

*Martin St. Ange*, Annales des sciences naturelles. Vol. XXIV. 1831.

Page 393.

*Meckel*, System der vergleichenden Anatomie. Halle, 1828. Vols. III

and IV.

*Pfeiffer*, Zur Anatomie des Schultergürtels und der Schultermuskeln

bei Säugethieren, Vögeln und Amphibien. Giessen, 1854.

*Rymer Jones*, Article ‘Reptiles’ in Todd’s Cyclop. of Anatomy and

Physiology. 1847–1852. Vol. IV. Page 263.

*Stannius*, Handbuch der Anatomie der Wirbelthiere. Vol. II. Zootomie

der Amphibien. Berlin, 1856.

*Wagner*, Icones Zootomicae. Leipzig, 1841.

*Wagner*, Lehrbuch der vergleichenden Anatomie. Leipzig, 1834–35.

*Zenker*, Batrachomyologia. Diss. Jena, 1825.

THE MUSCLES.

I. MUSCLES OF THE HEAD.

I. MUSCLES OF THE EYE.

*1.* _M. rectus inferior_ (Fig. 48 _ri′_).

*Dugès*, post-orbito-sous-oculaire, n. 12.--*Zenker*, _depressor

oculi_.

This muscle arises by a very thin, almost thread-like tendon from the

parasphenoid, near the _foramen opticum_, and becomes rapidly broader;

it lies beneath the tendon of the _membrana nictitans_ and a part

of the _m. retractor bulbi_: then runs forwards and outwards to be

attached into the lower part of the circumference of the eyeball.

[Illustration: Fig. 48.

Eye muscles of _Rana esculenta_, from below.

On the right side (_i. e._ in the left eye) the lev. bulbi (_l_) is

taken away; on the left side it is still present.

_l_ M. levator bulbi.

_l.a_ M. lev. anguli scapulae

_oi_ M. obliquus inferior.

_os_ M. obliquus superior.

_p_ M. pterygoideus.

_r_ M. retractor bulbi.

_re_ M. rectus externus.

1 First cervical vertebra.

]

*2.* _M. rectus externus_ (Fig. 48 _re_).

*Dugès*, post-orbito-ex-oculaire, n. 14.

Arises by a thin, thread-like tendon from the parasphenoid near the

_foramen opticum_, internal to and a little behind the _m. rectus

inferior_: it then passes obliquely outwards over the _m. retractor

bulbi_ and the tendon of the _membrana nictitans_, to be inserted into

the outer and hinder part of the circumference of the eyeball.

*3.* _M. rectus internus_ (Fig. 48 _ri_).

*Dugès*, post-orbito-in-oculaire, n. 13.

A long, thin muscle, which arises by a thread-like tendon from the

parasphenoid at the inner and hinder angle of the orbit, passes along

its inner wall, and is inserted into the antero-internal part of the

circumference of the eyeball.

*4.* _M. rectus superior_ (Fig. 49 _rs._).

*Dugès*, post-orbito-sous-oculaire.

Arises by a slender tendon from the fronto-parietal bone, widens

rapidly, and runs forwards and outwards to be inserted into the upper

part of the circumference of the _bulbus_.

[Illustration: Fig. 49.

Eye-muscles of _Rana esculenta_, from above.

_n_ Membrana nictitans.

_n′_ Tendon of membrana nictitans.

_p_ M. pterygoideus.

_rs._ M. rectus superior.

_t_ M. temporalis.

]

*5.* _M. obliquus inferior_ (Fig. 48 _oi_).

*Dugès*, pré-sous-orbito-oculaire.

This muscle arises near the floor of the orbital cavity, at the inner

and anterior angle, from the palatine bone by a thin and long tendon,

runs under Harder’s gland outwards and backwards, and is inserted,

almost at a right angle with the axis of the eye, into the anterior

part of the circumference of the eyeball.

*6.* _M. obliquus superior_ (Fig. 48 _os_).

*Dugès*, pré-sus-orbito-oculaire, n. 7.

The tendon of this small muscle arises from the inner end of the

palatine bone, near the origin of the _m. obliquus inferior_, runs

upwards, backwards, and outwards, and passes into a flat muscle, which

rapidly becomes broader, and is inserted into the upper surface of the

eyeball.

[Illustration: Fig. 50.

Eye-muscles of _Rana esculenta_, from below, the m. recti and obliqui

have been removed.

_n′_ Tendon of membrana nictitans.

_r_ M. retractor bulbi, lower part.

_r′_ Second part of same.

]

*7.* _M. retractor bulbi_ (Figs. 48 _r_, 50 _r_, _r′_).

*Dugès*, orbito-post-oculaire or choanide, n. 10.--*Zenker*, _m.

opticus_.

This muscle surrounds the optic nerve, and is situated within the

cone formed by the four recti muscles; it consists of three portions,

which take their broad and fleshy (the _recti_ and _obliqui_ arise

by thin tendons) origins from the under surface of the parasphenoid.

The first portion (Fig. 50 _r_) arises from the under surface of the

parasphenoid, nearly as far as the middle line, is broad and fleshy,

and covers, from below, the tendon of the _rectus externus_, then runs

forwards and outwards. The second portion (_r_′), which lies over

this and the tendon of the _m. rectus externus_, has also a broad and

fleshy origin, but runs more directly forwards, slightly crossing the

preceding. The third portion arises above the two others by a strong

thin tendon from the parasphenoid: it runs almost in the direction of

the _m. rectus internus_, but more directly forwards. The first two

portions pass more to the upper surface of the eyeball, the third more

to the under surface; consequently the whole muscle forms a sheath

surrounding the eyeball. The fibres to the upper surface of the eyeball

are inserted somewhat in front of the equator of the eyeball, those to

the under surface somewhat behind it.

*8.* _M. levator bulbi_, Dugès (Figs. 48 _l_, 51 _l_, 52 _l_).

*Dugès*, fronto-pterygoidien, n. 9.--*Klein*, _sustentator bulbi_.

[Illustration: Fig. 51.

Transverse section through the skull and orbital cavities of _Rana

esculenta_.

_b_ Bulb of eye.

_c_ Skull cavity.

_g_ Mucous membrane.

_l_ M. levator bulbi.

_m_ Upper jaw.

_n_ Membrana nictitans.

_n_′ Continuation of same to the lower eyelid.

_p_ Upper eyelid.

]

[Illustration: Fig. 52.

_M. levator bulbi_ of _Rana esculenta_, seen from above; the eyeball

which rests on it has been removed.

_d_._p_ M. depressor palpebrae inferioris.

_l_ M. levator bulbi.

_n_ Membrana nictitans.

_p_ M. pterygoideus.

_t_ M. temporalis.

]

There is no bony floor to the orbital cavity, and the oral and orbital

cavities are only separated by soft tissues; these are:--the mucous

membrane of the mouth, a thin fascia, and above this the muscle under

consideration. The fibres of this muscle run from before and from the

inner side obliquely backwards and outwards, below all the remaining

muscles of the eye, with the exception of the origin of the _m.

retractor bulbi_, and the tendons of the _m. obliquus inferior_, and

the _m. pterygoideus_; they arise on the inner side from the upper

and outer border of the fronto-parietal bone (Fig. 51), also from the

transverse portion of the parasphenoid and pterygoid bones, anteriorly

from the sphenethmoid and palatine bones; they run first downwards,

then, in the direction indicated, underneath the eye; and are inserted

externally by means of tendinous fibres into the superior border of the

upper jaw.

*9.* From the foregoing muscle certain fibres pass to the lower

eyelid, which may be regarded as a *depressor palpebrae inferioris*

(Fig. 52 _d.p_).

Manz[42] has described a muscle as _depressor palpebrae inferioris_

which arises in the neighbourhood of the external angle of the eye,

somewhat behind and below the lower border of the orbit, thence

passes obliquely upwards to be inserted into the lower eyelid. This

description is right as regards the insertion, but needs correction as

regards the origin. The muscle is attached to the orbital border in so

far that the tendinous expansion, into which the _levator bulbi_ passes

at its external border, is attached into the upper jaw; this bundle

splits off from the upper surface of the _levator bulbi_ at its middle,

and runs outwards on the under surface of the eyeball to the hinder

part of the lower eyelid.

[Footnote 42: Manz, Berichte der naturf. Gesellschaft zu Freiburg, 2

Band, 4 Heft, 1862, p. 391.]

[Illustration: Fig. 53.

Eye-muscles of _Rana esculenta_, from below, the m. recti and obliqui

have been removed.

_n_′ Tendon of membrana nictitans.

_r_ M. retractor bulbi, lower part.

_r_′ Second part of same.

]

*10.* Tendon of the _membrana nictitans_[43] (Fig. 53 _n_′).

The pigmented free border of the nictitating membrane passes, at the

inner and outer angles of the eye, into a tendon which passes for some

distance through fibrous tissue, then descends to the under surface

of the eyeball and joins that of the opposite side, so that the free

border of the eyelid, together with this tendon, form a complete ring.

The tendinous part of the ring, which is thin and thread-like, is found

on the under surface of the eyeball, and lies under the _m. retractor

bulbi_, and is bound to this muscle by connective tissue. Manz is

right in saying that the two muscles described by Dugès as _levatores

palpebrae inferiores_ (orbito-palpebral anterieur and posterieur, n. 5

and 6) do not exist; as is also the suggestion that he has confounded

the tendon of the nictitating membrane with these; this is clearly

shown in the figure in Plate VI of Dugès. With regard to the *actions*

of these eye-muscles, it is plain that the four straight and two

oblique muscles rotate the eyeball about three axes, one the axis of

the eye, one vertical, and the third passing from behind forwards and

inwards. The _m. retractor_ pulls the eyeball into the orbital cavity

backwards and downwards, while the _levator_ raises it.

[Footnote 43: Compare Manz, _l. c._]

There is less agreement about the movements of the eyelids, or rather

of the lower eyelid, for the upper has no independent movements.

Dugès (_l. c._) says that the two muscles, which he considers to be

_levatores palpebrae inferiores_, and which, according to him, are

connected with the _m. retractor bulbi_, draw up the lid as it is being

drawn backwards and downwards by the latter muscle. The depression

of the lid on the relaxation of the retractor and projection of the

eyeball is due simply to elasticity. Manz[44], on the contrary,

has shown that the sinking of the eyeball by the contraction of

the _retractor_ must necessarily cause a rising of the nictitating

membrane, as its tendons are attached to that muscle and so must follow

its movements.

[Footnote 44: _l. c._, p. 4.]

The depression of the lower lid occurs simultaneously with the raising

of the eyeball by means of the _m. levator bulbi_, through the

contraction of the _m. depressor palpebrae inferioris_, which proceeds

from that muscle; this is easily understood, as they are but parts of

the same muscle.

II. MUSCLES OF THE FACE.

*11.* _M. intermaxillaris s. dilatator narium_ (Fig. 54 _im_).

*Dugès*, intermaxillaire, n. 1.--*Zenker*, intermaxillaris medius.

This small muscle lies in the space between the vertical processes of

the two premaxillary bones, and consists of obliquely-crossing fibres

arising from the one bone and inserted into the opposite one. It brings

together the processes of the intermaxillary bones, so that their upper

extremities approach each other. At the same time, the cartilaginous

nasal coverings or wings (_an_) diverge and the nares expand; hence we

may regard this muscle as corresponding to a _dilatator narium_.

[Illustration: Fig. 54.

Facial muscles of _Rana esculenta_.

_an_ Cartilaginous nasal alae.

_i_′ Vertical process of premaxillary bone.

_im_ M. intermaxillaris.

_ln_ M. lateralis narium.

_n_ Cartilaginous nose capsule.

]

*12.* _M. lateralis narium_ (Ecker), (Fig. 54 _ln_).

*Zenker*, _m. intermaxillaris lateralis_.--*Klein*, _m. nasalis

inferior_.--*Dugès*, sus-maxillo-pré-nasal, n. 2.

This small muscle occupies the space between the anterior portion

of the maxillary bone and the ascending process of the premaxillary

bone. It arises from the maxillary, and ascends obliquely forwards

to be inserted into the outer border of the ascending process of the

premaxillary bone. It is antagonistic to the foregoing.

A third muscle is described by Dugès as _sus-maxillo-post-nasal_, n.

3 (_nasalis externus_, Klein); according to him it passes from the

_proc. nasalis_ of the maxillary bone to the outer border of the nasal

opening, which it widens. According to Klein (_l. c._, p. 9), this

muscle in _R. temporaria_ runs to the upper border of the maxillary

bone beneath the lower eyelid, and is long and narrow. I have, however,

never been able to find muscular fibre in this situation.

III. MUSCLES OF THE LOWER JAW.

[Illustration: Fig. 55.

Muscles of the back and shoulder.

_c_ M. cucullaris.

_dm_,_dm_ M. depressor maxillae, arising from the fascia dorsalis.

On the right side it is cut through and reflected.

_fd_ Fascia dorsalis.

_fd_′ Same cut near the spinous processes.

_i_ M. infraspinatus.

_l.a_ M. levator anguli scapulae.

_ld_ M. latissimus dorsi.

_r_ M. retrahens scapulae.

_sc_ M. sternocleidomastoideus.

]

*13.* _M. depressor maxillae inferioris_, Carus, Zenker (Figs. 55,

56, 63 _d.m_.).

*Cuvier*, _l. c._, II, 141, _digastricus_.--*Dugès*,

sus-occipitodorso-angulaire, 32.

This is a strong, triangular muscle, wide above, pointed below. It

is placed between the head and the shoulder-blade. It arises in two

portions; the greater, from the _fascia dorsalis_ (_fd_), covers the

scapula, and passes inwards and backwards over the muscles of the back;

anteriorly it passes over the _m. temporalis_ and is attached to the

fronto-parietal and squamosal, being continued to the fascia of the

upper eyelid. The second part is smaller, and arises by fleshy fibres

from the postero-superior arm of the squamosal, and from the posterior

and inferior border of the cartilaginous tympanic ring. The two parts

unite and are inserted into the hinder angle of the lower jaw. Cuvier

correctly considers that only the anterior portion corresponds to the

_digastricus_, the posterior answering to the _m. cervico-maxillaris_

of snakes. It draws the lower jaw down and opens the mouth.

*14.* _M. temporalis_ (Figs. 49, 52, 56, 57 _t_).

*Cuvier*, _m. temporalis_, I, _l. c._, p. 138.--*Dugès*,

sous-rupéo-temporo-coronoidien.

This muscle, together with the _M. pterygoideus_, occupies the space

between the prootic and the eyeball; it arises by the greater part of

its fibres from the upper surface of the prootic; it passes, narrowing,

beneath the anterior arm of the squamosal, then over the pterygoid

bone, between this and the maxillary and quadrato-jugal, receiving

on the way other fibres which arise from the anterior border of the

inferior arm of the squamosal and from the anterior circumference of

the tympanic ring. The collected fibres end in a broad, flat tendon,

which is inserted into the inner surface of the so-called _proc.

coronoideus_.

[Illustration: Fig. 56.

Muscles of the lower jaw of _Rana esculenta_.

_d_ M. deltoideus.

_dm_ M. depressor maxillae, its origin where covering

the m. temporalis reflected at *.

_ld_ M. latissimus dorsi.

_m_ M. masseter.

_t_ M. temporalis, origin.

_t_′ Lower portion of same.

]

*15.* _M. pterygoideus_ (Figs. 49 _p_, 52, 57 _pt_).

*Cuvier*, _m. temporalis_, II, _l. c._--*Zenker*,

_m. massetericus_, _l. c._, p. 25.--*Dugès*,

pré-rupéo-pterygo-maxillaire, 31.

This is partly covered by the foregoing, and lies between it and

the eyeball. It arises at the inner wall of the orbit from the

fronto-parietal, and from that portion of the prootic which may be

regarded as the _ala magna_, above the _m. retractor bulbi_. The muscle

is broad, and compressed from before backwards: it runs downwards

covered by the _m. temporalis_, and soon passes into a long, thin

tendon (Fig. 57 _pt_), which is inserted, behind the _m. temporalis_

and immediately in front of the joint, into the inner surface of the

lower jaw.

*16.* _M. masseter_ (Figs. 56, 57 _m_).

*Dugès*, zygomato-maxillaire, n. 29.

Arises as a somewhat broad muscle from the horizontal process of the

quadrato-jugal and is inserted into the outer side of the lower jaw,

opposite the insertion of the _m. temporalis_.

[Illustration: Fig. 57.

Muscles of the lower jaw of _Rana esculenta_.

_dm_ M. depressor maxillae cut through and reflected.

_la_ M. levator anguli scapulae.

_l.d._ M. latiss. dorsi, cut through and reflected.

_m_ M. masseter.

_pt_ M. pterygoideus.

_s_ Scapula.

_s_′ Suprascapula.

_s.c._ M. sternocleidomastoideus.

_t t_ M. temporalis cut through and reflected.

]

*17.* _M. submaxillaris_ (Figs. 58, 59, 61 _sm_, _sm_′).

*Cuvier*, _m. mylo-hyoideus_, _l.c._, II, 536.--*Zenker*,

_mylo-sternoideus_, p. 25.--*Dugès*, sous-maxillaire, n. 15.

This muscle forms the floor of the mouth as does the _mylo-hyoideus_ in

man; it is, however, not inserted into the hyoid bone.

It arises--1. From the whole of the inner surface of the upper border

of the lower jaw[45], except near the angle (Fig. 58 _sm_);

[Footnote 45: A second origin from the lower border of the mandible,

described by Klein, is not to be found. The appearance is due to the

exit of vessels and nerves from the groove between the mandible and the

muscle.]

2. By a small portion (Figs. 58, 59, 61 _sm_′) near the skull, from

the cartilaginous anterior cornu of the hyoid-bone which projects from

the cartilaginous part of the prootic bone. This portion descends

along the cornu to unite with the other portion. The two together form

a membranous layer, the fibres of which run transversely and meet in

a band of connective tissue along the middle line, forming a kind of

_linea alba_. The sheath runs forwards to the most projecting part of

the lower jaw superficially to the _m. submentalis_. Slightly in front

of its posterior border the muscle is attached in its whole breadth to

the skin of the throat by a fine lamella of connective tissue, which

is not always complete. The posterior border of the muscle is attached

to a fascia, which passes between the deeper muscles and is attached

to these. It is known that this muscle plays an important part in the

movements of respiration and swallowing; but its precise action has not

yet been worked out.

*18.* _M. submentalis_ (Fig. 59 _smt_).

*Cuvier*, _transversus_, _l. c._, p. 588.--*Dugès*, 4,

sous-mentonnier.--*Zenker*, _m. lingualis_ (incorrectly brought

into connection with _m. hyoglossus_).

This small muscle lies in the anterior angle of the lower jaw between

the dentary bones. It consists of transverse fibres which pass from one

bone to the other. Below it is covered by the most anterior portion of

the _m. submaxillaris_.

[Illustration: Fig. 58.

Muscles of the throat, chest, and abdomen of _Rana esculenta_.

_sm_ M. submaxillaris.

_sm_′ Hinder portion of same arising from

the anterior cornu of the hyoid bone.

(For other references see page 68.) ]

Its action is to approximate the sides of the lower jaw: according to

Dugès (_l. c._, p. 123), it assists in closing the nostrils, by drawing

together the sides of the lower jaw and thus raising the premaxillae.

IV. MUSCLES OF THE HYOID BONE AND THE TONGUE.

*19.* _M. geniohyoideus_ (Figs. 59 _gh_, 60 _gh_).

*Cuvier*, IV, 1, 536.--*Dugès*, n. 16.--*Zenker*, p. 30.--*M. St.

Ange*, _l. c._, Pl. XXVI, f. 1, 13, p. 423.

Arises from the lower jaw in two portions, the one from near the middle

line above the _m. submentalis_, the other more externally from the

upper border of the lower jaw. The two parts unite to form a long,

flat muscle, which lies on the ventral surface of the body of the

hyoid bone, and divides posteriorly into two portions. One of these,

the median (Figs. 59, 60 _gh_″), is inserted into the inner border of

the posterior horn of the hyoid bone, and is here attached to a fascia

which covers the _m. hyoglossus_ from beneath. By the same fascia the

muscles of opposite sides are connected in the space between the two

posterior cornua. The lateral division (Figs. 59, 60 _gh_′) is attached

to the posterior cartilaginous process of the hyoid bone.

[Illustration: Fig. 59.

Muscles of the hyoid bone and the tongue of _Rana esculenta_.

The m. submaxillaris has been removed close to its insertion.

_cm_′ Origin of the hinder portion of same from the hyoid bone.

_gh_ M. geniohyoideus.

_gh_′ External portion of same.

_gh_″ Internal portion of same.

_H_ Body of hyoid bone.

_H_′ Anterior horn.

_hg_ M. hyoglossus.

_oh_ M. omohyoideus.

_ph_ Mm. petrohyoidei.

_sh_ M. sternohyoideus.

_sh_′ External origin of same.

_sh_″ Internal origin of same.

_sm_ M. submaxillaris.

_sm_′ Origin of hinder portion of same.

_smt_ M. submentalis.

]

*20.* _M. sternohyoideus_ (Figs. 59 60 _sh_).

*Cuvier*, IV, 1, 538 (_pubio-hyoideus_).--*Dugès*, n. 17,

sterno-xipho-hyoidien.--*Zenker*, p. 30.

This muscle forms the cervical part of the _m. rectus abdominis_

and is, in fact, a direct continuation of it. It arises by an inner

portion (Fig. 59 _sh_″) from the upper surface of the inner extremity

of the coracoid and from the xiphisternum, while the outer portion

(_sh_′) is an immediate continuation of the _m. rectus abdominis_,

which passes into this muscle at about the fifth _inscriptio tendinea_.

The muscle passes forwards on the upper surface of the coracoid

and of the clavicle, under the pericardium towards the hyoid bone;

here the muscle, hitherto horizontal, suddenly becomes (compare

Fig. 60) vertical, and passes between the two insertions of the _m.

geniohyoideus_, and is inserted for a considerable length into the

lower surface of the hyoid bone and the anterior extremity of the

posterior cornu.

[Illustration: Fig. 60.

Muscles of the tongue and of the hyoid bone of _Rana esculenta_, from

below.

_gh_ M. geniohyoideus.

_gh_′ External origin }

_gh_″ Internal origin } of same.

_H_ Body of hyoid.

_H_′ Anterior cornu.

_H_″ Posterior cornu.

_hg_ M. hyoglossus.

_hg_′ Hinder origin of same.

_oh_ M. omohyoideus.

_ph_. 1, 2, 3, 4, first, second, third, and fourth

m. petrohyoideus.

_sh_ M. sternohyoideus.

]

*21.* _M. omohyoideus_ (Figs. 59, 60 _oh_).

*Cuvier*, _l.c._, p. 539.--*Dugès*, n. 18, interscapulo-hyoidien ou

omohyoidien.-- *Zenker*, p. 31.--*M. St. Ange*, _l.c._

Arises from the anterior border of the bony scapula, and is inserted

into the outer part of the ventral surface of the body of the hyoid: it

is partially covered by the _m. sternohyoideus_.

*22.* _M. petrohyoideus anterior_ (Ecker), (Fig. 61 _ph_1).

*Dugès*, 10, rupéo-cérato-hyoidien.--*Zenker*, _m. petro-ceraus_?

p. 30.-- *Klein*, _basio-hyoideus_, _l.c._, p. 71.

A thin, flat muscle, narrow above, but broadening rapidly below,

which arising from the outer extremity of the prootic bone, partially

surrounds the pharynx below, into which some of its fibres are

inserted; it is attached to the ventral surface of the body of the

hyoid bone, near the lateral notch. The chief action of this muscle

appears to be that of a _m. constrictor pharyngis_.

*23, 24, 25.* _Mm. petrohyoidei posteriores_ (Ecker), (Figs. 61

_ph_ 2, 3, 4).

*Cuvier*, _mm. stylohyoidei_, _l. c._, p. 537.--*Dugès*, 20, 21,

22, masto-hyoideus (Fig. 4).--*Zenker*, _petro-hyoideus superior_

and _inferior_.--*Klein*, _stylo-hyoideus_, _l. c._, p. 18.

These are three long, thin muscles, which lie nearly parallel

to one another, and run from the prootic bone to the posterior

cornu of the hyoid bone. All three arise, covered by the _m.

sternocleidomastoideus_, from the lateral extremity of the prootic,

pass round the pharynx and are attached, the _m. petrohyoideus

posterior primus_ (Fig. 61 _ph_ 2) to the anterior extremity, the

_secundus_ (Fig. 61 _ph_ 3) to the middle; and the _tertius_ (Fig. 61

_ph_ 4) to the hinder extremity of the posterior horn of the hyoid bone.

*26.* _M. hyoglossus_ (Figs. 59, 60, 61 _hg_).

*Cuvier*, _l. c._, p. 588.--*Dugès*, 24, hyo-glosse.

Arises from the bony posterior extremity of the hinder process of the

hyoid bone (_hg_′). The muscles of the two sides pass forwards and

converge to the middle line, where they unite. The azygos muscle thus

formed runs forwards on the ventral surface of the hyoid bone, between

the _mm. geniohyoidei_, over the anterior border of the hyoid, and

passes into the tongue, where it turns backwards and runs to the tip.

(For action, see tongue.)

[Illustration: Fig. 61.

Muscles of the hyoid bone of _Rana esculenta_, seen from the right side

and below.

_H_′ Anterior cornu.

_H_″ Posterior cornu.

_hg_ M. hyoglossus.

_H H_ Body of hyoid bone.

_ph_ 1 M. petrohyoideus anterior.

_ph_ 2, 3, 4 Mm. petrohyoidei posteriores.

_sm_ M. submaxillaris.

_sm_′ Hinder portion of same.

]

*27.* _M. genioglossus._

*Cuvier*, _l. c._, p. 587.--*Meckel*, _l. c._, IV, 339.--*Dugès*,

23, genio-glosse.

Arises above the insertion of the _m. geniohyoideus_ on either side

near the middle line from the two anterior bones of the lower jaw. The

two origins quickly unite to form a thick, very firm muscular belly,

which in its course backwards rapidly thins, and is distributed by

numerous bundles; decussating with the fibres of the _m. hyoglossus_ at

an acute angle, it runs on to the anterior extremity of the tongue.

II. MUSCLES OF THE TRUNK.

I. MUSCLES OF THE ABDOMEN.

In the formation of the abdominal wall the following muscles take

part:--(1) _m. rectus abdominis_, (2) _m. obliquus externus_, (3)

_m. obliquus internus_, (4) the _portia abdominalis_ of the _m.

pectoralis_. Above, the _m. obliquus internus_ forms a kind of

*diaphragm*, partially separating the abdominal from the thoracic

cavity (p. 70).

*28.* _M. rectus abdominis_ (Fig. 62 _r_).

*Dugès*, pubio-thoracique, n. 52.

These muscles, situated on either side of the middle line of the

abdomen, are as a rule traversed by five notched _inscriptiones

tendineae_. Each muscle arises, by a narrow, strong tendon, from the

inferior border of the pubes, passes forwards, and quickly widening,

divides at the second _inscriptio tendinea_ (counting from behind) into

two portions.

(_a_) The Outer portion (Fig. 62 _r_) runs into the _portio

abdominalis_ of the _m. pectoralis_, and forms the greater lateral

division of that muscle.

(_b_) The Inner (_r_′) continues as the _m. rectus abdominis_ and

gradually narrows anteriorly. At the sternum some of the fibres (the

median) are inserted into the dorsal surface of the cartilaginous plate

of the xiphisternum, while the greater part of the muscle continues

forwards over the coracoid to become the _m. sternohyoideus_ at about

the fifth _inscriptio tendinea_ (p. 64).

*29.* _M. obliquus externus_ (Fig. 63 _oe_, _oe_′).

*Zenker*, _l. c._, p. 31, _m. obliquus_.--*Dugès*,

dorso-sous-abdominal, n. 54.

The _m. obliquus externus_ arises from--

(_a_) The aponeurosis covering the long muscles of the back, and by

this from the spinous processes. The anterior edge of the _m. obliquus

externus_ covers the posterior edge of the _m. latissimus dorsi_,

which arises from the inferior, ventral surface of the aponeurosis.

The aponeurosis also divides laterally into two parts, one of which,

the posterior, passes into the _obliquus externus_, while the anterior

forms the tendon of origin of the _depressor maxillae inf._ (_dm_)

(compare Fig. 66).

(_b_) A second, narrow portion, _portio omo-abdominalis_ (Fig. 63

_oe_′) (_m. xipho-adscapulaire_, Dugès, n. 62), arises from the

posterior border of the scapula by a thin tendon, and becomes broader

as it runs downwards and backwards, to join the anterior border of the

other and larger portion.

[Illustration: Fig. 62.

Muscles of the chest, throat, and belly of _Rana esculenta_.

_ch_ M. coraco-humeralis.

_d_ M. deltoideus.

_oi_ M. obliquus abdom. internus.

_oe_ M. obliq. abdom. externus.

_oe_′ Scapular portion of same.

_oh_ M. omohyoideus.

_p_′ Port. sternalis anterior of m. pectoralis.

_p_″ Port. sternalis posterior of same.

_p_‴ Port. abdominalis of same.

_r_ M. rectus abdominis.

_r_′ Inner portion of same.

_sh_ M. sternohyoideus.

_sm_ M. submaxillaris.

_sm_′ Hyoid origin of same.

_sr_ M. sternoradialis.

]

The whole muscle is attached by its most anterior fibres to the

cartilage of the xiphisternum, the rest passing into an aponeurosis

which, inseparably connected with the _inscriptiones tendineae_,

traverses the lower surface of the _m. rectus abdominalis_ to the

_linea alba_.

[Illustration: Fig. 63.

Muscles of trunk of _Rana esculenta_, from the right side.

_cd_ M. cutaneus femoris.

_d_ M. deltoideus.

_d.m._ M. depressor maxillae.

_i_ M. infraspinatus.

_ld_ M. latiss. dorsi.

_oe_ M. obliquus abdom. externus.

_oe_′ Scapular origin of same.

_ss_ M. subscapularis.

_t_ M. triceps brachii.

]

*30.* _M. obliquus internus_ (and _transversus_), (Figs. 64, 65

_oi_).

*Dugès*, ileo-transverso-sous-sternal, p. 53.--*Zenker*,

_transversus_, _l. c._, p. 31.--*Kuhl*, _transversus_, _l. c._,

p. 116.

This muscle corresponds with the combined _obliquus internus_ and

_transversus_, and has therefore been described either as the one or

the other, by various authors.

The fibres arise tendinously--

*a.* From the transverse processes of the vertebrae from the fourth

backwards, and from the fascia covering the _mm. intertransversarii_.

*b.* From the iliac bone, by a strong tendon from its upper border, and

by a few weaker fibres from its outer surface. The latter fibres are

covered by the former, and these end posteriorly in a sharp concave

border.

The muscular bundles diverge from these points, some running forwards,

some backwards; the former are only partially covered by the _portio

omo-abdominalis_ of the _m. obliquus externus_ (compare Figs. 62, 63),

in front of the anterior edge of that muscle. The posterior border of

the muscle is uncovered.

[Illustration: Fig. 64.

Second layer of abdominal muscles of _Rana esculenta_, from the right

side and below. The m. obliquus externus and the right anterior

extremity has been removed.

_dm_ M. depressor maxillae.

_hg_ M. hyoglossus.

_ld_ M. latissimus dorsi and infraspinatus.

_oi_ M. obliquus internus.

_oi_′ Insertion of third portion of same.

_oi_″ Second portion inserted into the pharynx.

_pc_ Pericardium.

_ph^4._ M. petrohyoideus quartus.

_r_ M. rectus abdominis.

_ss_ M. subscapularis.

]

The insertions of the muscle are very various, and as a consequence

very various functions are accomplished by it.

*a.* The most anterior part of the muscle (Fig. 65 _oi_‴), the fibres

of which run forwards, is attached--

1. Partly to the xiphisternum and the coracoid.

2. A second portion, _viz._ that arising from the transverse processes

of the fourth vertebra (Fig. 64 _oi_″), surrounds the pharynx like a

diaphragm (Fig. 65 _oe_), and is attached to its side as far as the

dorsal aspect (Fig. 65 _oi_′). This portion is lightly separated from

the following.

3. A third portion, placed behind the preceding, runs from the pharynx

over the pericardium (Fig. 64 _pc_), and is attached to this nearly

as far as the middle line (Figs. 64 _oi_′ and 65 _oi_″), resting on

the sternum, the _m. rectus_ and _m. sterno-hyoideus_. The lines of

insertion of the muscles of opposite sides form an angle, open in

front. In the thin borders of this portion the muscular bundles lie

almost entirely in simple layers, and they are therefore well adapted

for microscopical purposes.

*b.* The middle and posterior portions of the muscle pass downwards

and slightly backwards, towards the outer border of the _m. rectus_;

there they pass into a tendon which, for the most part, runs on the

upper surface of the _rectus_ to the _linea alba_.

II. MUSCLES OF THE BACK.

The muscles of the back are covered by a fascia, the *fascia dorsalis*

(Fig. 66 _fd_), attached to the spinous processes of the vertebrae.

Anteriorly, where it covers the _m. temporalis_, it is attached to the

fronto-parietal and squamosal bones, posteriorly to the superior border

of the iliac bones and the extremity of the urostyle. The posterior

part, which covers the origins of the _m. longissimus dorsi_, of the

_m. coccygeo-iliacus_, and of the _coccygeo-lumbaris_, is especially

strong. From the anterior part several muscles arise laterally, viz.

a portion of the _m. obliquus abdominalis externus_, of the _m.

latissimus dorsi_, and of the _m. depressor maxillae inferioris_. In

the uppermost layer, immediately covered by the _fascia dorsalis_, lie

the _m. cucullaris_, the _m. retrahens scapulae_, the _m. latissimus

dorsi_ arising from the fascia itself, and the scapula with its

muscles; then come the long muscles of the back; and in the third and

deepest layer the short muscles of the back.

[Illustration: Fig. 65.

M. obliquus internus, showing its attachments superiorly.

_h_ Xiphisternum.

_oe_ Oesophagus.

_oi_ M. obliquus internus.

_oi_′ Fibres of m. obliq. int., which are inserted

into the oesophagus.

_oi_″ _oi_‴ Fibres of same, which are attached to

coracoid and xiphisternum.

_p_ Pericardium.

_r_ M. rectus abdominis.

]

The individual muscles are as follows:--

A. _LIMB MUSCLES OF THE BACK_ (for moving the shoulder-girdle and arm).

*31.* _M. cucullaris_ (Fig. 66 _c_).

*Cuvier*, _angularis_.--*Dugès*, n. 38, sous-occipito

adscapulaire.--*Zenker*, _levator scapulae sublimis_.

In order to see this muscle, it is necessary to separate the _fascia

dorsalis_ from the spines of the vertebrae and to turn it back with

the _m. depressor maxillae inf._ arising from it. It is then seen to

rise from the posterior surface of the exoccipital as far as the middle

line; the muscles of the two sides run backwards, diverging each from

the other, so as to form an angle, and are then attached on each side

to the anterior superior angle of the suprascapula at its under surface

and median border.

Cuvier considers this muscle as the analogue of the _m. levator anguli

scapulae_. Dugès and Meckel consider it to be the anterior part of the

_m. cucullaris_, the former thinking the posterior part of the muscle

to be the _m. rhomboideus_. I cannot agree with these opinions, and I

regard the muscle as simply representing the _m. cucullaris_ of man

(Ecker).

*32.* _M. latissimus dorsi_ (Figs. 56, 66 _ld_).

*Dugès*, n. 66, lombo-huméral.--*Zenker*, _depressor brachii_.

This thin, triangular muscle arises from the ventral surface of the

_fascia dorsalis_, and is posteriorly covered in part by the anterior

border of the _m. obliquus abdominis externus_. The muscle becomes

narrower towards the sides, and passes into a flat tendon, which

blends with that of the _m. infraspinatus_, and is attached to the

outer surface of the _crista deltoidea_ of the humerus by a triangular

expansion.

*33.* _M. retrahens scapulae_ (Ecker), (Fig. 66 _r_).

*Dugès*, n. 59, lombo-adscapulaire.--*Klein*, _m. rhomboideus_, _l.

c._, p. 26.--*Zenker*, _omoplateus rectus_? p. 37, _l. c._--*Kuhl*,

_l. c._, 124, _retrahens rhomboideus_.

[Illustration: Fig. 66.

Muscles of the back and shoulder-blade.

_c_ M. cucullaris.

_dm_ M. depressor maxillae arising from the fascia dorsalis,

cut through and reflected on the right side.

_fd_ Fascia dorsalis.

_fd_′ Same cut through at the spines.

_i_ M. infraspinatus.

_l.a_ M. levator anguli scapulae.

_ld_ M. Latissimus dorsi.

_r_ M. retrahens scapulae.

_sc_ M. sternocleidomastoideus.

]

This flat, oblong muscle arises from the transverse process of

the fourth vertebra, and is connected with the third tendinous

inscription (counting from before) of the _m. longiss. dorsi_.

Posteriorly its origin forms a convex border, whence it runs forwards

and to the side; it is inserted into the ventral surface of the

suprascapula, near its inner border. This muscle lies near the _m.

serratus_ or _transv. scap. tertius_ (Fig. 68 _ts_″), and has an

analogous position. It would, therefore, possibly be more correct

to regard it also as a _m. serratus_. It arises from parts which

represent ribs, and is inserted into the scapula. It has, at any

rate, no analogy to the _m. cucullaris_, with which Dugès connects

it, nor with the _m. rhomboideus_, which Klein holds it to be, as it

does not arise from spinous processes.

B. _LONG MUSCLES OF THE BACK._

*34.* _M. extensor dorsi communis_ (Fig. 67 _lg.d._).

*Dugès*, n. 33–40, masse des muscles surspinaux devisée en

huit faisceaux principaux: 1. vertebro-sus-occipital, 2–5.

transverso-spinaux, 6. transverso-coccyg., 7. sacro-coccygien,

8. ileo-coccygien. The muscles 1–7 form the lombo-costal, 8 the

ischicoccyg., Cuv.--*Zenker*, _m. sacrolumbaris_.--*Klein*,

_longiss. dorsi_, _coccygeo-lumb._, _coccyg.-iliac_.

This muscular mass is covered by the _fascia dorsalis_, the _m.

cucullaris_ and _retrahens scapula_, and the suprascapula: it

corresponds with (1) the _m. ileo-costalis_ and _longissimus dorsi_,

_dorsi_ and _cervicis_ (Henle); (2) the _semispinalis_ and _spinalis_

(Henle).

The following separate portions may be made out:--

*35.* (1) _M. longiss. dorsi_ (Fig. 67 _lg.d._).

This muscle arises from the most anterior part of the urostyle, runs

forwards near the middle line, separated from the muscle of the

opposite side by the spinous processes of the vertebrae. The inner

portion passes over the posterior spinous processes without being

connected with them, as far as the fifth, it is then inserted into

the succeeding spines as far as the exoccipital bone. The lateral

portion runs outwards and forwards to be attached to the transverse

processes and oblique processes of all the vertebrae, from the sixth

forwards, while accessory muscular slips arise from the transverse

processes and from the anterior spinous processes which, merging in

the general muscle-mass, are inserted with this into the occiput.

The muscle is subdivided by a series of wavy, tendinous partitions

(Fig. 67), which arise from the transverse processes, and traverse

the muscle in planes, the upper surfaces of which are directed

upwards and forwards. This arrangement of the muscle indicates

its relation with the lateral trunk muscles of fish and fish-like

amphibians.

*36.* (2) _M. coccygeo-sacralis_ (Fig. 67 _c.l._).

This muscle is partially covered by the preceding, it arises

from the lateral surfaces of the anterior half of the urostyle;

from this origin the fibres pass forwards and outwards to be

inserted in two portions, the anterior set into the arch and the

posterior set into the transverse process of the last vertebra. The

former may be regarded as _m. intercruralis_, the latter as _m.

intertransversarius_.

*37.* (3) _M. coccygeo-iliacus_ (Fig. 67 _c.i._).

This muscle arises from nearly the whole length of the lateral

surface of the urostyle, the anterior portion being concealed by

the preceding muscle. The fibres pass outwards and forwards to be

inserted into the anterior two-thirds of the inner surface of the

ilium.

[Illustration: Fig. 67.

Muscles of the back and pelvic girdle of _Rana esculenta_.

_c.i._ M. coccygeo-iliacus.

_c.l._ M. coccygeo-sacralis.

_g.l._ M. glutaeus.

_i_ MM. intertransversarii.

_i′_ MM. intercrurales.

_i.c.i._ M. intertransv. capitis inferior.

_i.c.s._ M. intertr. cap. sup.

_il._ M. ilio-lumbaris.

_lg.d._ M. longissimus dorsi.

_o.i._ M. obliq. internus.

_t_ M. temporalis.

]

*38.* _M. ilio-lumbaris_ (Klein), (Fig. 67 _il._).

*Dugès, n.* 51, transverso-iliaque.--*Cuvier*, *Zenker*, etc.,

_quadr. lumborum_.

Evidently represents the _quadratus lumborum_ of human anatomy;

it takes its origin from the anterior extremity of the ilium, and

runs forwards to be inserted into the transverse processes of the

vertebrae from the seventh to the fourth.

C. _SHORT MUSCLES OF THE BACK._

*39.* _M. intertransversarius capitis superior_ (Ecker), (Fig. 67

_i.c.s._).

*Dugès*, n. 41, ex-occipito-transversaire sup.

A small muscle arising from the prootic: it lies above the _levator

anguli scapulae_, and is inserted into the transverse process of the

second vertebra.

*40.* _M. intertransversarius capitis inferior_ (Fig. 67 _i.c.i._).

*Dugès*, n. 42, ex-occipito-transv. inf.

This muscle lies on the ventral surface of the preceding one, but is

easily separable from it; it arises from the angle of the prootic and

is inserted, together with the preceding muscle, into the transverse

process of the third vertebra. The _n. vagus_ passes outwards between

these two muscles.

*41.* _MM. intertransversarii dorsi_ (Fig. 67 _i_).

*Dugès*, n. 43 to 50, intertransversaires.

Seven small muscles, placed between adjacent transverse processes of

the vertebrae; from the second to the last, in continuity with the

muscles last mentioned.

*42.* _MM. intercrurales_ (Ecker), (Fig. 67 _i_′).

*Klein*, p. 29, interspinales, interobliqui.

Those muscular fibres found between the arches of adjacent vertebrae

are so named. On account of the shortness of the spinous processes

they can scarcely be named _m. interspinales_.

III. MUSCLES OF THE ANTERIOR EXTREMITY.

I. MUSCLES OF THE SHOULDER-GIRDLE.

A. _DORSAL MUSCLES OF THE SHOULDER-BLADE_.

(I.) Muscles arising from the head or trunk, and inserted into

the scapulae, which they act upon primarily, and upon the

shoulder-girdles secondarily.

Some of these muscles, such as _m. cucullaris_ and _retrahens_,

are situated on the back, and have already been described with

the muscles of that region. Others are attached partly to the

under-surfaces of the scapulae and are only visible from below; they

are best seen in such a preparation as is shown in Fig. 68, made by

cutting through the sternum in the middle line, and reflecting each

half outwards.

(*a*) _Muscles which arise from the skull_.

*43.* _M. levator anguli scapulae_ (Fig. 68 _la_).

*Dugès*, sous-occipito-adscapulaire, n. 60.--*Zenker*, _protractor

scapulae_, Pl. I, Fig. 5.--*Cuvier*, Leçons I, p. 379, describes

this muscle as part of the _m. serratus anticus magnus_.

A fairly strong muscle, which has a broad origin from the prootic

and exoccipital bones as far as the _foramen magnum_, a few fibres

are also attached to the lateral portions of the parasphenoid. The

fibres run backwards and outwards, first in a frontal plane and then

in a sagittal plane, to be inserted into the under surface of the

suprascapula near the posterior border, where it encloses the hinder

border of the _m. interscapularis_.

Its action is to draw the shoulders forwards and inwards, or the head

downwards.

*44.* _M. sternocleidomastoideus_ (Fig. 68 _sc_).

*Dugès*, scapulo-mastoidien, n. 65.--*Cuvier*,

sterno-mastoidien.--*Zenker*, _protractor scapulae_, Pl. I, Fig. 5,

1.

A narrow muscle placed in front of and externally to the preceding

muscle; its general direction resembles that of the _m. levator ang.

scap_.

The fibres arise from the most external part of the prootic and

from the hindermost part of the squamosal, under cover of the

_m. digastricus maxillae_, and from the hinder portion of the

cartilaginous tympanic ring. Covered by these muscles, it passes

backwards and downwards to be inserted into the concavity of the

anterior border of the scapula; externally to the origin of the

deltoid.

The action of this muscle is to draw the shoulder forwards and

towards the middle line, or to bend the head downwards.

*45.* _M. protrahens scapulae_ (Fig. 68 _ps_).

*Dugès*, *Zenker*, _protractor acromii_, Pl. I, Fig. 5, 3.

This muscle lies to the outer side of the _rectus capitis inferior_:

it arises from the prootic, and runs backwards and outwards upon the

_levator anguli scapulae_, to be inserted into the scapula.

It pulls the scapula forwards.

(*b*) _Muscles which arise from the vertebrae._

*46.* _M. transverso-scapularis major_ (Ecker), (Fig. 68 _ts_).

*Dugès*, transverso-interscapulaire, n. 63.--*Cuvier*,

_l. c._, p. 380, probably regarded as a portion of the

_serratus_.--*Zenker*, Pl. I, Fig. 5, 3; _depressor acromii_,

Pl. II, Fig. 3, 15.--*Klein*, _depressor scapulae_.

This muscle arises by tendon from the cartilage of the transverse

process of the fourth vertebra and by a slip from the cartilage of

the transverse process of the third vertebra. Diminishing in size,

the muscle passes outwards, forwards, and upwards to be inserted

into the posterior border of the scapula, opposite the insertion

of the _m. sternocleidomastoideus_. It draws the shoulder inwards,

backwards, and downwards.

*47.* _M. transverso-scapularis minor_ (Ecker), (Fig. 68 _ts_′).

*Dugès*, transverso-adscapulaire, n. 61.--*Cuvier*, regarded as

part of the _serratus_.--*Zenker*, Pl. I, Fig. 5, 7, _detractor

scapulae_.

This is much shorter than the muscle just described; it arises under

cover of the _m. levator scapulae_, from the transverse process of

the third vertebra in front of the _transverso-scapularis major_; and

runs outwards and forwards to be inserted into the ventral surface of

the cartilage of the scapula. The action of the muscle is to draw the

shoulder inwards, backwards, and outwards. Dugès regards this muscle

as a portion of the _serratus anticus magnus_, the remainder of which

he considers is to be found in the _portio scapularis_ of the _m.

obliquus externus_ (xipho-adscapulaire, Dugès, n. 62).

[Illustration: Fig. 68.

Muscles of the shoulder, from below; the sternum has been cut through

and the sides drawn apart.

_d_ Deltoideus.

_is_ Interscapularis.

_it_ Intertransversarii.

_la_ Lev. anguli scapulae.

_ps_ Protrahens scapulae.

_sc_ Sternocleidomastoideus.

_ss_ Subscapularis.

_ts_ Transverso-scap. major.

_ts_′ Transverso-scap. minor.

_ts_″ Transverso-scap. tertius.

]

*48.* _M. transverso-scapularis tertius s. serratus_ (Ecker),

(Fig. 68 _ts_″).

This is larger than the muscle just described; arising by a broad

and flat origin from the transverse processes of third and fourth

vertebrae, it runs upwards and inwards to be inserted into the

ventral surface of the cartilaginous scapula near its inner border.

In position, course, and action the muscle most nearly represents the

_m. serratus anticus_ of human anatomy.

(II.) Muscles on the scapula, that is, arising from the scapula and

inserted into the scapula or humerus.

(*a*) _Muscles on the deeper surface._

*49.* _M. interscapularis_ (Fig. 69 _i.s._).

*Dugès*, interscapulaire, n. 64.

[Illustration: Fig. 69.

Muscles of right shoulder and upper arm. The coracoids (_co_) and

the clavicles (_cl_) have been drawn apart after cutting through the

sternum.

_c.h._ M. coraco-humeralis.

_cl_ Clavicle.

_c.o._ Coracoid.

_d_ M. deltoideus.

_d_′ Clavicular portion of m. deltoideus.

_i_ M. infraspinatus.

_i.s._ M. interscapularis.

_l.a._ M. levator anguli scapulae.

_l.d._ M. latissimus dorsi.

_p_′ Sternal portion of m. deltoideus.

_p_″ Anterior sternal portion of the m. pectoralis.

_p_‴ Posterior sternal portion of the m. pectoralis.

_s_ Scapula.

_s.r._ M. sternoradialis.

_t.r._ M. triceps brachii.

]

One end of this muscle is tendinous, and is attached to the outer

extremity of the suprascapula; from this the muscle, becoming fleshy

and narrower, runs towards the middle line to become attached to

the ridge on the ventral surface of the scapula, from which the _m.

deltoideus_ and the _m. subscapularis_ arise, and between which it

passes. It approximates the scapula and suprascapula and lessens the

angle between them.

*50.* _M. subscapularis_ (Fig. 70 _ss_).

*Dugès*, sous-scapulo huméral, n. 72.

This muscle is situated on the upper or visceral surface of the

scapula. It arises from a bony ridge found on the upper surface of

the coracoid and of the _proc. coracoideus_ of the scapula. The

muscle runs outwards and is inserted, after widening, into the inner

surface of the _crista deltoidea_ of the humerus; it is antagonistic

to the deltoid, and draws the raised arm backwards and towards the

trunk.

[Illustration: Fig. 70.

Right shoulder, from below, arm strongly abducted.

_cl_ Clavicle.

_co_ Coracoid.

_d_ Deltoideus.

_d_′ Clavicular portion of deltoid.

_h_ Humerus.

_ss_ Subscapular muscle.

]

(*b*) _Muscles on the superficial surface._

*51.* _M. infraspinatus_ (Fig. 57, between _ld_ and _dm_, Fig. 71

_i_).

*Dugès*, adscapulo-huméral, n. 67.

The _m. infraspinatus_ represents the _m. infraspinatus_, _teres

major_ and _minor_ of human anatomy. It is of triangular form, with

the broad base directed inwards; it arises from the whole of the

upper surface of the suprascapula except along the inner border which

is formed of hyaline cartilage alone: from this origin the fibres

converge outwards to a flat tendon which unites with that of the _m.

latissimus dorsi_ to be inserted into the _crista deltoidea humeri_,

by means of a thin, triangular, tendinous expansion.

[Illustration: Fig. 71.

Muscles of the back and shoulder.

_c_ M. cucullaris.

_dm_ M. depressor maxillae.

_fd_ Fascia dorsalis.

_fd_′ Fascia dorsalis reflected.

_i_ M. infraspinatus.

_l.a_ M. levator anguli scapulae.

_ld_ M. latissimus dorsi.

_r_ M. retrahens scapulae.

_sc_ M. sternocleidomastoideus.

]

*General arrangement of the muscles of the shoulder-blade.*

The muscles attached to the scapula and suprascapula are:

1. Muscles of the back (_m. cucullaris_ and _m. retrahens scap._).

2. Muscles of the neck (_m. omohyoideus_).

3. Muscles of the abdomen (_portio scapularis_ of the _m. obliquus

externus_).

4. True shoulder muscles (_m. levator anguli scapulae_; _m.

sternocleidomastoideus_; _m. protrahens scapulae_; _mm.

transverso-scapularis major_, _minor_, and _tertius s. serratus_; _m.

interscapularis_; _m. subscapularis_; and the long head of the _m.

triceps_).

*From above* the following are visible: _m. cucullaris_, a portion of

the _m. subscapularis_, and the _transverso-scapularis maior_.

*From below* (after cutting through and drawing aside the

two halves of the sternum) may be seen: the _omohyoideus_,

_sternocleidomastoideus_, _levator anguli scapulae_, _protrahens

scapulae_, _transverso-scapularis minor_ and _tertius_,

_interscapularis_ and _subscapularis_.

B. _VENTRAL MUSCLES OF THE SHOULDER (PECTORAL MUSCLES)_

These arise from the sternum or shoulder-girdle and are inserted

either into the humerus or into the radio-ulnar.

*52.* _M. pectoralis_ (Fig. 72).

This is placed on the ventral surface of the shoulder-girdle and

consists of the following parts:

(α) _Portio sternalis anterior_ (Ecker), (Figs. 72, 73 _p_′).

*Dugès*, clavi-huméral, n. 70 (port. clavic. du grand pectoral).

This is the anterior portion of the _m. pectoralis_, it arises from

the sternum proper and the epicoracoids. Broad at the origin, it

becomes narrower as it runs outwards into a tendon attached to the

_crista deltoidea humeri_.

(β) _Portio sternalis posterior_ (Ecker), (Figs. 72, 73 _p_″).

*Dugès*, sterno-huméral, n. 71 (port. sternal du grand

pectoral).--*Klein*, _humero-sternalis_.

This muscle is placed immediately behind the _portio sternalis

anterior_, it has a somewhat broad origin from the sternum and

xiphisternum; the muscle runs outwards and slightly forwards to be

inserted into the groove beside the _crista deltoidea humeri_. The

tendon of the _m. sterno-radialis_ passes between these two portions

of the _m. pectoralis_.

(γ) _Portio abdominalis_ (Figs. 72, 73 _p_‴).

*Dugès*, abdomino-huméral, n. 69 (port. costal du grand

pectoral).--*Zenker*, _brachio-abdominalis_.--*Klein*,

_humero-abdominalis_.

[Illustration: Fig. 72.

Muscles of the chest, throat, and belly of _Rana esculenta_.

_ch_ M. coraco-humeralis.

_d_ M. deltoideus.

_oi_ M. obliquus abdom. internus.

_oe_ M. obliq. abdom. externus.

_oe_′ Scapular portion of same.

_oh_ M. omohyoideus.

_p_′ Port. sternalis anterior to m. pectoralis.

_p_″ Port. sternalis posterior to same.

_p_‴ Port. abdominalis of same.

_r_ M. rectus abdominis.

_r_′ Inner portion of same.

_sh_ M. sternohyoideus.

_sm_ M. submaxillaris.

_sm_′ Hyoid origin of same.

_sr_ M. sternoradialis.

]

This portion represents that part of _m. pectoralis major_ of

human anatomy which arises from the costal cartilages, and more

particularly that part which is connected with the aponeurosis of

the _m. obliquus abdominis externus_. The muscle arises (1) from

the _m. rectus abdominis_ (Fig. 72 _r_), that is, it forms a direct

continuation of the outer portion of this muscle; (2) the inner,

smaller part arises from the superficial surface of the aponeurosis

of the _m. obliquus abdominis_. The muscle becomes narrower as it

courses outwards and forwards, the outer fibres being longer and more

oblique than the inner. The action of this muscle will necessarily

vary according to the part or parts which are brought into action;

speaking generally, the anterior extremities will be drawn downwards

and towards each other.

*53.* _M. coraco-humeralis_ (Dugès), (Figs. 72, 73 _c.h._).

*Dugès*, n. 72.--*Klein*, _adductor humeri_.

A long, narrow muscle, lying upon the posterior border of the

coracoid and the lowest part of the _m. subscapularis_. By its

position it corresponds most nearly with the _m. pectoralis minor_,

although its insertion does not. It has a narrow origin from the

coracoid near the sternum, whence it courses outwards, under cover

of the _port. sternalis anterior_ and _posterior_ of the _m.

pectoralis_, to be inserted into the middle of the humerus between

the deltoid and internal head of the triceps. It draws the limb

towards the trunk.

*54.* _M. sternoradialis_ (Cuvier), (Figs. 72, 73 _s.r._).

*Dugès*, pré-sterno-clavi-radial, n. 74.

This muscle evidently represents the biceps of man, it lies in front

of the _portio sternalis anterior_ of the _m. pectoralis_, its hinder

border being under cover of the latter muscle. It takes origin from

the episternum, omosternum, and the epicoracoid; from this broad

origin the fibres converge while coursing backwards and outwards

to be attached to a strong tendon. This tendon plays in a groove

along the _crista deltoidea_, and is held in position by tendinous

bands arising from the insertion of the _m. pectoralis_; it then

pierces the lower portion of the muscular belly of the deltoid, and

is inserted into the anterior extremity of the radial side of the

radio-ulnar. The muscle is a powerful flexor of the forearm.

*55.* _M. deltoideus_ (Figs. 70, 72, 73 _d_).

*Dugès*, pré-sterno-scapulo-huméral, n. 68.

It is placed in front of and external to the _m. sternoradialis_, and

represents both the _m. deltoideus_ and _supraspinatus_ of man. The

muscle has two points of origin:--

(*a*) _Portio scapularis_, the larger, external portion is attached

to the outer end of the clavicle, and to the anterior process of the

precoracoid, but it arises chiefly from the dorsal surface of the

scapula (from the same ridge as the _m. subscapularis_), and lastly

from the anterior border and ventral surface of the scapula. The

fibres pass outwards over the shoulder-joint.

[Illustration: Fig. 73.

Muscle of right shoulder and upper arm. The coracoids (_c.o._) and

the clavicles (_cl_) have been drawn asunder after cutting through

the sternum.

_c.h._ M. coraco-humeralis.

_cl_ Clavicle.

_c.o._ Coracoid.

_d_ M. deltoideus.

_d_′ Clavicular portion of m. deltoideus.

_i_ M. infraspinatus.

_i.s._ M. interscapularis.

_l.a._ M. levator anguli scapulae.

_l.d._ M. latissimus dorsi.

_p_′ Sternal portion of m. deltoideus.

_p_″ Anterior sternal portion of the m. pectoralis.

_p_‴ Posterior sternal portion of the m. pectoralis.

_s_ Scapula.

_s.r._ M. sternoradialis.

_t.r._ M. triceps brachii.

]

(*b*) _Pars clavicularis_ (_musc. cleido-humeralis_, Klein) arises

from the inner extremity of the clavicle and from the omosternum and

joins the _portio scapularis_.

The muscle is inserted into the humerus, the under part of the

_portio scapularis_ being attached to the upper portion of the

_crista deltoidea_; the outer fibres of this part, together with

those of the _pars clavicularis_, are attached to the extremity of

the _crista deltoidea_ and to the inner surface of the humerus as

far as the distal extremity; just before its insertion the muscle is

pierced by the tendon of the _m. sternoradialis_. The deltoid draws

the limb forwards.

II. MUSCLES OF THE FORE-LIMB.

A. _MUSCLES OF THE ARM._

_M. sternoradialis (biceps)._ See page 82.

*56.* _M. triceps brachii_ (Fig. 73 _t.r._).

*Dugès*, scapulo-huméro-olecranien, n. 75.--*Capes*, *Zenker*, _m.

anconaeus_.

This muscle lies on the upper or dorsal surface of the arm, and

has relations similar to those of the corresponding muscle in man.

The long head arises from the posterior border of the scapula at

the upper border of the glenoid cavity, and is here attached to

the capsule of the joint; the inner head arises from the upper and

inner surfaces of the anterior half of the humerus as far as the

extremity; the outer head arises from the outer surface of the

humerus. Additional fibres, which may be regarded as a fourth origin

(_m. subanconaeus_), arise from the upper surface of the hinder

half of the humerus. The fibres from these several origins unite

to form a strong muscle, which covers the upper, inner, and outer

surfaces of the bone, and then passes into a tendon. This tendon is

attached to the capsule of the elbow-joint and inserted into the

proximal extremity of the radio-ulnar. A cartilage, representing the

olecranon, is found in the capsule at the point of attachment of the

tendon.

B. _MUSCLES OF THE FOREARM._

(I.) Muscles of the flexor surface.

The muscles of this surface are arranged in two groups, so as to

form, at the elbow-joint, a triangular depression (_plica cubiti_),

into which the tendon of the _m. sternoradialis_ sinks.

A. Muscles of the Inner Group.

*57.* _M. flexor carpi I s. radialis_ (Fig. 74 _Fc_).

*Dugès*, sous-huméro-carpien, n. 83.

This represents either the _flexor carpi radialis longior_ or

_radialis brevior_ of man; it has a broad origin from the inner

border of the humerus above the epicondylus medialis. It becomes

narrower as it passes downwards to be inserted by a tendon into the

_os semilunare_ and _naviculare_. The muscle is much stronger in

males than in females, and the crest from which it takes origin is

correspondingly larger in the former sex (p. 42). Tendinous fibres

connect it with the following muscles:--

*58.* _M. flexor carpi II s. ulnaris_ (Fig. 74 _Fc_′).

*Dugès*, epitrochlo-carpien, n. 84.

This muscle represents the _flexor carpi ulnaris_ or _ulnaris

anterior_, and is close to the inner side of the foregoing. It arises

from the _epicondylus medialis_, being connected by fibres with the

_flexor digitorum communis_, and is inserted into the _os naviculare_.

[Illustration: Fig. 74.

Muscles of the right arm of _Rana esculenta_.

_ed_ M. extens. digitorum communis.

_ei_ M. abductor digiti II longus.

_Fa_′ M. flexor antibrachii lateralis superficialis.

_Fc_ M. flexor carpi radial.

_Fc_′ M. flexor carpi ulnar.

_Fd_ M. flexor digitorum communis.

_sr_ Tendon of the M. sternoradialis.

]

*59.* _M. flexor digitorum communis_ (Figs. 75, 76, 77 _Fd_).

*Dugès*, epitrochlo-sous-phalangettien, n. 118.

This muscle lies most internally, and corresponds to the _m. flexor

digitorum sublimis_ of man. It arises in common with the foregoing

muscle from the _epicondylus medialis_, and runs downwards to the palm

of the hand. At its entrance into this it diminishes and passes into

the flat _aponeurosis palmaris_. (See hand, B.)

*60.* _M. flexor antibrachii medialis_ (Ecker), (Fig. 75 _Fa_).

*Dugès*, epitrochlo-sous-radial, n. 79. (Frequently described as the

_pronator rotundus_, a description which cannot be retained, as the

implied movement does not occur in the frog.--Ecker.)

This muscle lies deeply on the inner side of the forearm, covered by

the _flexores carpi_; it arises tendinously from the _epicondylus

medialis_ and is inserted into the inner surface of the radial side of

the radio-ulnar as far as the inferior articular extremity.

*61.* _M. flexor antibrachii lateralis superficialis_ (Ecker),

(Figs. 74, 75 _Fa_′).

*Dugès*, premier ex-huméro-radial, n. 76. (Usually described as

_supinator longus_, this, for reasons similar to those in the case

of _m. flexor antibrachii medialis_, cannot be retained.)

The situation of this muscle corresponds with that of the _supinator

longus_ of man; it lies to the outer side of the tendon of the _m.

sternoradialis_, and arises by two heads, the one, above from the

outer edge of the humerus, the other deeper from the _epicondylus

lateralis_. They unite and pass into a tendon which, running over the

lower articular extremity of the radio-ulnar, is partly attached to

the carpus and partly connected with the tendon of the _m. extensor

digiti II proprius longus_. From its position and insertion, this

muscle evidently flexes the forearm and extends the hand.

[Illustration: Fig. 75.

Muscles of the right arm of _Rana esculenta_, deep layer.

_ed_ Extensor digit. communis.

_Fa_ Flexor antibrachii medialis.

_Fa_′ Flexor antibrachii lat. superficialis.

_Fa_″ Flexor antibr. lat. profundus.

_Fc_ Flexor carpi radialis.

_Fc_′ Flexor carpi ulnaris.

_Fd_ Flexor digitorum communis.

]

*62.* _M. flexor antibrachii lateralis profundus_ (Ecker), (Fig. 75

_Fa_″).

*Dugès*, epicondylo-sus-radial, n. 78.--*Klein*, _flexor

antibrachii_. (_Supinat. brevis_ autt.)

This muscle arises under cover of the foregoing, from the

_epicondylus lateralis_ of the outer surface of the humerus, it

becomes broader and is inserted into the whole length of the lower

(volar) ridge of the radio-ulnar. It is a powerful flexor of the

forearm.

(II.) Muscles of the extensor surface.

*63.* _M. extensor digitorum communis longus_ (Fig. 75 _ed_).

*Dugès*, huméro-sus-digital, n. 95.

This muscle lies on the outer border of the forearm; it arises

in common with the short head of the _flexor antibr. lateralis

superficialis_ from the _epicondylus lateralis_, and from the surface

above it; it runs downwards along the outer side of the forearm,

in order to pass into an aponeurosis on the back of the hand,

which expands upon the third, fourth, and fifth fingers, and ends

between them in a free concave border. This aponeurotic expansion is

connected with the tendons of the _extensor brevis digitorum_.

*63*.* _M. abductor digiti II_ (_i. e. pollicis_) _longus_ (Fig. 74

_ei_).

**Dugès*, cubito-métacarpien, n. 87.

This muscle lies between the foregoing and the _flexor antibrachii

lateralis superficialis_, and appears superficially in the interval

between them. It arises from the lateral surface of the radio-ulnar,

and runs obliquely over the _m. flexor antibrachii lateralis

profundus_ and the hinder extremity of the radio-ulnar towards the

second finger into the metacarpal bones of which it is inserted.

*64.* _M. extensor carpi ulnaris s. ulnaris post._ (Fig. 76 _ec_).

*Dugès*, epicondylo-sous-carpien, n. 85.--*Klein*, _abductor carpi

internus_.

This muscle has a somewhat narrow origin from the _epicondylus

lateralis_; during its course through the forearm, where it lies

between the _mm. anconaei_ on the one side and the _extensor

digitorum communi_ on the other, it becomes broader and is inserted

into the outer surface of the carpus in the following manner. By

one tendinous slip it is attached to the palmar border of the

_os pyramidale_, by a second slip to the outer border of the _os

capitato-hamatum_. This latter slip is usually connected in its

dorsal portion with the _extensor digitorum communis brevis_.

[Illustration: Fig. 76.

Muscles of forearm of _Rana esculenta_, dorsal view.

_aa_ M. anconaei.

_ec_ M. extens. digit. communis.

_Fd_ M. flex. digit.

]

*65, 66.* _M. anconaei s. extensores antibrachii_ (Fig. 76 _aa_).

*Dugès*, epicondylo-cubital and epitrochlo-cubital, n. 80, 81.

Both these muscles lie on the extensor side of the forearm; they

spring, the one from the _epicondylus lateralis_, the other from the

_epicondylus medialis_; converging, they run downwards, enclose the

olecranon, and are inserted into the upper (dorsal) border of the

ulna, each forming a penniform muscle. They are extensors of the

forearm and assist the _m. triceps_.

C. _MUSCLES OF THE HAND._

(I.) Muscles on the palmar surface.

(1) Muscles common to all the digits.

*67.* _M. flexor digitorum communis_ and _m. palmaris brevis_

(Fig. 77 _fd_, _pb_).

The origin and course of this muscle has been described above. At

its entrance into the palm it suddenly diminishes in size and passes

into a triangular aponeurosis, _aponeurosis palmaris_. Into the outer

border of this aponeurosis is inserted the _m. palmaris brevis_

(Fig. 77 _pb_); this latter muscle arises from the posterior or ulnar

border of the lower extremity of the radio-ulnar below the insertion

of the _mm. anconaei_, and runs obliquely downwards to this insertion.

From the lower free border of the aponeurosis a number of muscles

arise which will be described with the rest of the digital muscles.

(2) Muscles of the rudimentary thumb.

*68.* _M. abductor pollicis_ (Figs. 77, 78 _ap_).

*Dugès*, cubito-pollicien, n. 100.

This short muscle arises from the upper border of the lower extremity

of the radio-ulnar, passes transversely outwards, and is then

inserted into the anterior border of the rudimentary thumb.

*69.* _M. adductor pollicis_ (Figs. 77, 78 _ad.p_).

*Dugès*, sous-carpo-pollicien, n. 101.

This lies more superficially than the last, it arises from the palmar

surface of the _os capitato-hamatum_, and is inserted together

with the _abductor pollicis_ into the thumb. Several fibres of the

_extens. dig. II brevis_ are also inserted into the thumb. As this

member is completely covered with skin, the motions of which it is

capable are very limited.

(3) Muscles of the second finger, which serves the purposes of a

thumb.

*70.* _M. flexor digiti II longus s. sublimis_ (Fig. 77 _f2_).

*Dugès*, sous-carpo-phalangettien de l’index, n. 133.

[Illustration: Fig. 77.

Muscles of hand of _Rana esculenta_, volar surface, larger than

nature.

_A_ Aponeurosis palmaris.

_ab5_ M. abductor dig. V primus.

_ab5_′ M. abductor dig. secundus.

_ac_ MM. anconaei.

_adp_ M. adductor pollicis.

_ap_ M. abductor pollicis.

_ec_ M. extensor digitorum communis.

_e5_ M. extens. dig. commun. brevis.

_f2_ M. flexor dig. II longus.

_f3_ M. flexor dig. III longus.

_f4_ M. flexor dig. IV longus.

_f5_ M. flexor dig. V longus.

_fb2_ M. flexor dig. II brevis.

_fb3_ and _fb3_′ MM. flexores dig. III breves.

_fb4_ and _fb4_′ MM. flexores dig. IV breves.

_fb5_ M. flexor dig. V primus.

_fd_ M. flexor digitorum communis.

_o_ M. opponens.

_pb_ M. palmaris brevis.

I First finger, rudimentary thumb.

II Second finger, functions as thumb.

III-V Third to fifth fingers.

]

This is a flat muscle, which arises for the most part by fleshy

fibres from the hinder free border of the _aponeurosis palmaris_,

also by a smaller part from the palmar surface of the _os

capitato-hamatum_. It quickly passes into a thin long tendon which is

inserted into the terminal phalanx of the thumb (second finger).

[Illustration: Fig. 78.

Second layer of muscles on volar surface of hand of _Rana esculenta_,

larger than nature.

_ab5_ M. abductor dig. V primus.

_ab5_′ M. abductor dig. V secundus.

_ad2_ M. adductor dig. II.

_ad.p_ M. adductor pollicis.

_ap_ M. abductor pollicis.

_ec_ M. extensor carpi.

_fb2_ M. flexor dig. II brevis.

_fb2_′ M. flexor dig. II tertius.

_fp_ MM. flexores proprii phalangum.

_i3_ }

_i4_ } MM. interossei volares.

_i4_′}

_i5_ }

_o_ M. opponens dig. II.

_o_′ M. opponens dig. V.

_tt_ ′ MM. transversi metacarpi.

I-V as in Fig. 77.

]

*71.* _M. flexor digiti II brevis s. profundus_ (Fig. 77 _fb2_).

*Dugès*, sous-carpo-phalangien de l’index, n. 102.

This flat muscle, covered by the preceding, has a tendinous origin

from the palmar surface of the _os capitato-hamatum_, together with

one head of the _flexor sublimis_, and is inserted into the basal

phalanx.

*72.* _M. flexor digiti II tertius_ (Fig. 78 _fb2_′).

*Dugès*, tendini-phalangien de l’index, n. 103.

A small, thin muscle which arises close to the foregoing from the _os

capitato-hamatum_, and is inserted into the basal phalanx (see below,

_m. interosseus volaris I_).

*73.* _M. adductor digiti II_ (Fig. 78 _ad2_).

*Dugès*, métacarpo-métacarpien de l’index, n. 90.

This muscle represents the _adductor pollicis_ of man; it arises

from the base of the third os metacarpal and is inserted into the

metacarpal of the second finger, which latter it draws towards the

third finger.

*74.* _M. opponens digiti II_ (Figs. 77 and 78 _o_).

*Dugès*, sous-carpo-métacarpien de l’index, n. 90.

This muscle represents the muscle of like name in man. It arises from

the palmar process of the _os capitato-hamatum_ by a narrow tendon

and passes into a broad, flat muscle, which is inserted into the

under surface and outer border of the metacarpal of the second finger

throughout its length. It has, undoubtedly, the same action as the

corresponding muscle in man.

(4) Muscles of the third finger.

*75.* _M. flexor digiti III longus s. sublimis_ (Fig. 77 _f3_).

The smaller portion of this muscle arises from the free border of the

_aponeurosis palmaris_, the chief portion, together with the _flexor

brevis_, from the palmar surface of the _os capitato-hamatum_. The

two portions pass into one long tendon, which is inserted into the

last phalanx.

*76, 77.* _MM. flexores digiti III breves s. profundi I, II_

(Fig. 77 _fb3_ and _fb3_′).

*Dugès*, tendini-phalangiens du medius, n. 104, 105.

These two muscles arise in part from the free border of the

_aponeurosis palmaris_, in part from the volar surface of the _os

capitato-hamatum_, and pass backwards on either side of the muscle

last described, to be inserted into the basal phalanx. Dugès regards

these as analogues of the _mm. lumbricales_.

*78.* _M. flexor metacarpi dig. III._

This is a somewhat strong muscle, which arises by fleshy fibres from

the palmar surface of the _os capitato-hamatum_, and is inserted into

the palmar surface of the metacarpal bone.

(5) Muscles of the fourth finger.

*79.* _M. flexor digiti IV longus s. sublimis_ (Fig. 77 _f4_).

It arises at the free border of the _aponeurosis palmaris_, and is

inserted into the terminal phalanx by a thin tendon.

*80, 81.* _MM. flexores digiti IV breves s. profundi I, II_

(Fig. 77 _fb4_ and _fb4_′).

*Dugès*, tendini-phalangiens de l’annulaire, n. 108, 109.

Dugès regards these also as analogues of the _m. lumbricales_; they

arise on either side of the foregoing muscle from the free border

of the _aponeurosis palmaris_, and are inserted by tendons into the

basal phalanx.

*82.* _M. flexor proprius phalangum dig. IV_ (Fig. 78 _fp_).

*Dugès*, phalango-phalangien, n. 115.

A small muscle which arises by two slips from the palmar surface

of the basal phalanx, and is inserted into the base of the middle

phalanx.

*83.* _M. flexor metacarpi dig. IV_ (Fig. 78 _i4_).

This somewhat strong muscle arises from the _os capitato-hamatum_,

and is inserted into the palmar surface of the metacarpal bone.

(6) Muscles of the fifth finger.

*84.* _M. flexor dig. V longus s. sublimis_ (Fig. 77 _f5_).

This corresponds exactly with the corresponding muscle of the fourth

finger.

*85.* _M. flexor dig. V brevis_ (Fig. 77 _fb5_).

This muscle arises from the free border of the aponeurosis and is

inserted into the basal phalanx.

*86.* _M. flexor proprius phalangum dig. V_ (Fig. 78 _fp_).

*Dugès*, phalango-phalangien, n. 17.

It corresponds exactly with the corresponding muscle of the fourth

finger.

*87.* _M. opponens dig. V_ (Fig. 78 _o_′).

*Dugès*, deuxième sous-carpo-métacarpien du digitule, n. 92.

This muscle has a narrow origin from the palmar surface of the _os

capitato-hamatum_, it widens and is inserted into the outer border of

the metacarpal of the small finger. It draws this finger towards the

thumb.

*88.* _M. abductor dig. V primus_ (Figs. 77 and 78 _ab5_).

*Dugès*, sous-pyro-pré-métacarpien du digitule, n. 93.

This muscle arises from the palmar border and the outer surface of

the _os pyramidale_, and is inserted into the basal phalanx by a thin

tendon.

*89.* _M. abductor dig. V secundus_ (Figs. 77 and 78 _ab5_′).

*Dugès*, sous-pyro-post-métacarpien du digitule, n. 94.

It arises from the posterior part of the _os pyramidale_, is short,

and is inserted into the outer side of the metacarpal.

(II.) Muscles on the dorsal surface.

_M. extensor digitorum communis longus_ (Figs. 75 _ed_ and 77 _ec_).

*90.* _M. extensor digitorum communis brevis_ (Fig. 79 _e_3, _e_4,

_e_5).

*Dugès*, sous-pyro-phalangettien du III, IV, V, n. 124, 128, 131.

Covered by the foregoing, this muscle arises from the _os pyramidale_

in conjunction with the _extensor pollicis_ and from the _os

capitato-hamatum_. The part destined for the third finger, together

with that for the fourth, arises from the _os pyramidale_, passes

obliquely backwards into a tendon which runs along the inner side

of the corresponding finger to be inserted into the last phalanx.

That for the fifth finger arises from the inner projection of _os

capitato-hamatum_ being here attached to the insertion of the _m.

extensor carpi ulnaris_, and is inserted in a similar manner into the

terminal phalanx of the fifth finger.

*91.* _M. abductor digiti II_ (_i.e._ _pollicis_) _longus_

(Figs. 74 _ei_ and 79 _ab.2_).

For the origin and course of this muscle in the forearm see No. 63.

The muscle passes obliquely over the wrist-joint from the little

finger towards the inner side and from before backwards, it is

inserted into the metacarpal of the second finger (thumb). The place

of insertion of this muscle into the metacarpal (compare p. 47) is,

in the males, developed into a ridge and process, the muscle itself

is enlarged during the breeding season. This muscle abducts the thumb

and by this means presses upon the chest of the female when clinging

to her.

*92.* _M. extensor digiti II proprius longus_ (Fig. 79 _e2_, _e2_′).

*Dugès*, cubito-radio-sous-phalangien de l’index, n. 96.

[Illustration: Fig. 79.

Muscles of hand of _Rana esculenta_. Dorsal view, twice natural size.

_ab.2._ M. abductor dig. II longus.

_ab.2_′ M. abductor dig. II brevis.

_ad.2_ M. adductor dig. II.

_e2_ }

_e2_′ } M. extensor dig. II longus.

_e3_ }

_e4_ } M. extensor digitorum communis brevis.

_e5_ }

_eb2_ M. extensor dig. II proprius brevis.

_ec_ M. extensor carpi ulnaris.

_fa_ M. flexor antibr. lat. superficial.

_i_′3 }

_i_′4 } MM. interossei.

_i_′5 }

II-V as in Fig. 77.

]

This muscle lies obliquely on the back of the hand, parallel with

the _m. abductor digiti II_. It arises by two heads, one from the

tendon of the _flexor antibr. lat. superf._ (n. 60) upon the hinder

extremity of the radio-ulnar. This tendon conceals the hinder end

of this bone by means of a patella-like thickening, and sends forth

three tendinous prolongations; of these, the one is attached to the

ulnar border of the radio-ulnar, the second to the _os naviculare_,

while the third constitutes that portion of the _m. extens. dig. II

proprius long._ which is under consideration. The second head arises

from the _os pyramidale_ in connection with the _extensor digitorum

communis brevis_ (n. 90). The heads unite to form a long muscle,

which runs obliquely towards the second finger, passes into a tendon,

which, after forming connections by some muscular fibres with the

metacarpal, is inserted into the terminal phalanx.

*93.* _M. extensor dig. II proprius brevis_ (Fig. 79 _eb2_).

*Dugès*, sus-luno-phalangettien de l’index, n. 120.

This muscle lies between the preceding muscle and the _abductor dig.

II longus_. It arises from the _os naviculare_, and also includes

some fibres which arise from the second metacarpal. The tendon runs

along the radial side and is inserted along with the last muscle into

the terminal phalanx.

*94.* _M. abductor dig. II brevis_ (Fig. 79 _ab._2′).

*Dugès*, sus-luno-métacarpien de l’index, n. 88.

This represents the _abduct. pollic. brevis_ of man, lies on the

outer side of the hand, arises from the _os naviculare_, and is

inserted into the second metacarpal.

*95.* _M. extensor dig. III proprius._

*Dugès*, sus-luno-phalangettien du médius, n. 122.

This muscle has two heads of origin, from the _os naviculare_ and

_lunatum_. These form one muscular belly, which rapidly diminishes

and passes into a tendon; this, after receiving a few muscular fibres

from the second metacarpal, runs to the outer part of the dorsal

surface of this finger as far as the terminal phalanx, where it is

inserted.

*96.* _M. extensor dig. IV proprius._

*Dugès*, sus-pyro-phalangettien de l’annulaire, n. 128.

By a somewhat broad base, this muscle arises from the _os naviculare_

and _lunatum_. The tendon receives fibres from the fourth metacarpal

and runs on the outer half of the dorsal surface of this finger as

far as the middle phalanx, where it is inserted.

(III.) _MM. interossei._

*97, 98.* [*a.*] _MM. transversi metacarpi_ (Ecker), (Fig. 78 _t_,

_t_′, _ad2_).

These lie transversely between the metacarpals. There are three; the

first has been already described as _m. adductor digiti II_ (n. 73);

the second is stretched between the third and fourth metacarpals, and

the third between those of the fourth and fifth fingers.

*99, 100, 101.* [*b.*] _MM. interossei volares._

One may regard as such, four long, thin muscles, which take origin

near one another on the palmar surface of the carpal bones, and

diverge from this towards the second and fifth fingers.

1. The first of these muscles (Fig. 78 _fb2_′) has already been

described above as the _m. flexor digiti II tertius_ (n. 72).

2. The second (Fig. 78 _i3_) passes from the origin already described

to the palmar surface of the basal phalanx of the third finger.

3. The third (Fig. 78 _i4_) runs, in like manner, to the fourth

finger.

4. The fifth (Fig. 78 _i5_) to the fifth finger[46].

[Footnote 46: Whether these muscles may be regarded as _mm.

interossei_ is open to discussion, as they appear to act more as _mm.

opponentes_.]

*102* to *108*. _MM. interossei dorsales._

These arise in common from the metacarpals, and are inserted into the

basal phalanx of the corresponding surface on the finger.

1. The _third_ finger has two, one to the radial side (Fig. 79

_i_′3), one to the ulnar side.

2. The _fourth_ finger has three, the third (Fig. 79 _i_′4) arises

from the carpus, and perhaps ought rather to be regarded as an

extensor.

3. The _fifth_ finger has two.

IV. MUSCLES OF THE HINDER LIMB.

I. MUSCLES OF THE THIGH.

General arrangement (Figs. 80, 81).

*a.* The _m. triceps femoris_ forms the _outer border_ of the thigh,

and extends on to both the dorsal and ventral surfaces. The outer

border therefore corresponds with the anterior surface of the thigh

of man, in consequence of the thigh in the frog being normally

rotated outwards.

*b.* The _m. vastus internus_ (_v.i._) and a portion of the _adductor

longus_ (_ad_′), the _sartorius_ (_s_), a portion of the _adductor

magnus_ (_ad_‴), and the _rectus internus major_ (_r.i_′) are found

on the _inner surface_ of the thigh.

*c.* The _rectus internus minor_ (_r.i_″) forms the _inner border_.

*d.* On the _dorsal surface_ are to be seen, nearest the trunk,

the _glutaeus_ (_gl_), on the outer border the _vastus externus_

(_v.e._), the _pyriformis_ and _biceps_ (_p_ and _b_), then the

_semimembranosus_ (_sm_), and lastly the _rectus internus minor_

(_ri_″).

*e.* By reflecting the _rectus internus major_ (compare Fig. 82)

there are exposed to view: the _semitendinosus_ (_st_), the whole

length of the _adductor longus_, _magnus_ (_ad_′ and _ad_‴), and

_brevis_ (_ad_″). By reflecting the triceps, one sees (Figs. 84,

85) the _ilio psoas_ (_ip_), and beneath the _semimembranosus_, the

_quadratus_ (_qf_), and _capsularis femoris_ (_og_).

_Description of the separate Muscles._

(A.) Muscles of the dorsal surface.

*109.* _M. glutaeus_ (Fig. 80 _gl_).

*Dugès*, ex-ilio-trochantérien.--*Zenker*, _glutaeus maior_.

The tendinous origin of this muscle is found on the outer surface

and upper border of the two hinder thirds of the iliac bone. Thence

it runs outwards and backwards between the head of the _m. vastus

externus_ (_ve_) on the one side, and the _ilio-psoas_ and _rectus

anticus_ (_ra_) on the other, to be inserted into a tubercle

(trochanter) of the femur.

*110.* _M. Pyriformis_ (Fig. 80 _p_).

*Cuvier*, pyramidal.--*Dugès*, coccy-fémoral, n. 136.

A narrow, slender muscle, arising from the apex of the coccyx; it

passes obliquely outwards and backwards to the thigh, to be inserted

deeply between the _m. vastus externus_ and the _biceps_, into the

inner surface of the femur, but towards the ventral surface.

_M. triceps femoris s. extensor cruris communis_ (Fig. 80).

*Dugès*, pelvi-fémoro-rotulien, n. 145, 146, 147.

This is a large, strong muscle, having three heads of origin, and

inserted by a tendon into the tibio fibula. The three heads are the

_caput longum_ or the _rectus femoris anticus_ (_r.a._), the _caput

externum_ or the _vastus externus_ (_v.e._), and the _caput internum_

or the _vastus internus_ (Fig. 82 _vi_); of these, the first is

situated on the outer border of the thigh, the second on the dorsal

surface, and the third on the ventral surface.

[Illustration: Fig. 80.

Muscles of left thigh of _Rana esculenta_. Dorsal view.

_b_ M. biceps.

_ci_ M. coccygeo-iliacus.

_g_ M. gastrocnemius.

_gl_ M. glutaeus.

_p_ M. pyriformis.

_pe_ M. peroneus.

_r.a._ M. rectus anterior.

_ri_″ M. rectus internus minor.

_sm_ M. semimembranosus.

_t.a._ M. tibialis anticus.

_tr._ M. triceps fem.

_v.e._ M. vastus externus.

]

*111.* [*a.*] _M. rectus femoris anticus_ (Figs. 80, 82 _ra_).

This arises from the under (ventral) surface of the iliac bone about

its middle and beneath the _m. glutaeus_. The muscular belly quickly

passes into an aponeurosis, which covers the other two muscular

masses and combines with their tendons.

*112.* [*b.*] _M. vastus externus_ (Fig. 80 _v.e._).

It arises from the upper and hinder extremity of the iliac bone

(Fig. 42 _il_′) behind the _glutaeus_, which here sinks in between

this muscle and the _rectus anterior_.

*113.* [*c.*] _M. vastus internus_ (Figs. 81, 82 _vi_).

It arises from the whole of the under and outer surfaces of the

hip-joint capsule, and from tendinous slips which pass from this

capsule downwards on the under surface of the muscle, (from the

pubis and the femur I have not seen any fibres of origin, as stated

by others). Anteriorly the _vastus_ may easily be separated into

two heads. The two _vasti_ unite and form one muscular belly, in

the formation of which, as above stated, the _rectus anticus_ takes

no part. This united muscle passes into a tendon which, without the

mediation of a patella, passes over the knee-joint to be inserted

into the anterior extremity of the tibio fibula, and is prolonged

into the fascia of the leg.

*114.* _M. ilio-fibularis s. biceps_ (Fig. 80 _b_).

*Dugès*, ilio-péronien, n, 149.--*Zenker*, flexor externus tibiae,

p. 42.

This is a long, somewhat narrow muscle, which lies to the inner side

of, and partly covered by, the _vastus externus_, between which and

the semimembranous it is placed. It arises by a tendon between the

_ilio-psoas_ and _pyriformis_ from the ilium above the acetabulum

and behind the origin of the _vastus externus_. From here it passes

backwards, separated from the _vastus externus_ by a strong _ligam.

intermusculare_. On the inner side of this muscle, between it and

the _m. semimembranosus_, run the vessels of the thigh and the _N.

ischiadicus_. The muscle then divides into two portions, of which

one is inserted high up on the inner and under surface of the femur.

This is probably the muscle which Klein (_l. c._, p. 61) describes as

the _extensor brevis_. The other long muscular belly passes into a

thin tendon, opposite the hinder extremity of the thigh, it divides

into two portions; the anterior is inserted into the dorsal surface

of the hinder extremity of the femur, the other into the dorsal

surface of the tibio-fibula. Between the two is stretched an arch

with the concavity outwards. This insertion of the biceps is covered

by the stronger tendon of origin of the _m. gastrocnemius_ (compare

Fig. 87).

*115.* _M. semimembranosus_ (Figs. 80, 86 _sm_).

*Dugès*, sus-ischio-poplité, n. 148.--*Klein*, p. 60, _extensor

femoris sublimis_.

This muscle lies on the dorsal surface of the thigh, to the inner

side of the _biceps_ and _pyriformis_: it is a broad muscle, which

takes a tendinous origin from the hinder and upper angle of the

_symphysis ossis ilei_, and is inserted by a flat tendon into the

dorsal surface of the ligaments of the knee-joint, and through these

into the tibio-fibula. The tendon is formed rather abruptly from

the muscle, and is surrounded (Fig. 86) by the concave arch, which

forms the tendon of origin of the _gastrocnemius_ (_q. v._). A little

behind its middle (compare Fig. 81) the muscle is divided obliquely

into an anterior and a posterior portion by an aponeurotic septum. As

all the fibres are interrupted by this structure, the muscle may very

easily be torn through.

(B.) Muscles on the ventral surface.

*116.* _M. sartorius_ (Figs. 81, 82 _s_).

*Dugès*, sous-iléo-tibial, n. 150.--*Klein*, _gracilis_.

This long, flat muscle lies along the middle of the ventral surface

of the thigh; it arises by a tendon from the anterior inferior angle

formed by the symphysis of the innominate bones, and terminates in a

tendon behind the knee-joint, forming a kind of _pes anserinus_ in

the fascia of the leg, similar to that of the _semitendinosus_[47].

[Footnote 47: The sartorius has lately been used by Kühne in his

researches on the nerve-endings in muscle and on muscle contraction

without nerve supply (du Bois-Reymond’s and Reichert’s Archiv, 1859.

Page 314).]

*117.* _M. rectus internus major_ (Ecker), (Figs. 81 _r.i_′, 82

_ri_′).

*Dugès*, post-ischio-tibial profond, n. 152.--*Klein*,

_semimembranosus_.

This is a broad and flat muscle, thinned out at either extremity;

it occupies the inner half of the ventral surface of the thigh,

and is in relation with the _adductor magnus_ externally and

anteriorly, with the _sartorius_ posteriorly, and the _rectus minor_

muscle internally. The muscle arises by a tendon from the pubic

symphysis; it covers in its course the _semitendinosus_, then passes

into a strong tendon, which blends with the tendon of the _rectus

minor_, to be inserted by a slip into a process of the tibio-fibula

representing the _tuberositas tibiae_; first passing between the

tendons of the _sartorius_ and _semimembranosus_, which here form

an aponeurotic arch; a second slip passes under the tendon of the

_m. semitendinosus_ to be inserted into the upper surface of the

tibio-fibula, under it passes the _N. tibialis_. A third thread-like

tendon passes to the _m. semimembranosus_. Somewhat behind its middle

(Fig. 82) the muscle is traversed by an _inscriptio tendinea_, by

which the whole of the fibres are interrupted.

[Illustration: Fig. 81.

Muscles of left thigh of _Rana esculenta_. Ventral surface.

_ad_′ M. adductor longus.

_ad_″ M. adductor brevis.

_ad_‴ M. adductor magnus.

_e.c._ M. extensor cruris.

_f.t._ M. flexor tarsi.

_g.c._ M. gastrocnemius.

_r.i_′ M. rectus internus major.

_r.i_″ M. rectus internus minor.

_s_ M. sartorius.

_t.a._ M. tibialis anticus.

_t.p._ M. tibialis posticus.

_v.i._ M. vastus internus.

]

*118.* _M. rectus internus minor_ (Ecker), (Figs. 80, 81 _r.i_″).

*Dugès*, post-ischio-tibial superficiel, n. 151.--*Zenker*, _flexor

tibiae magnus_.--*Klein*, _ischio-tibialis_.

Situated entirely on the inner surface of the thigh, this narrow

muscle arises by a tendinous band which runs the whole length of the

pubic symphysis, and is connected posteriorly with the _sphincter

ani_, anteriorly with the _rectus abdominis_; posteriorly it passes

into a thin tendon which is also connected with the muscle just

described (117). In its course this muscle is attached to the skin

by strands of vessels and connective tissue.

The vastus internus major and minor together undoubtedly correspond

with rectus internus or gracilis of human anatomy. The rectus

internus major is, however, regarded as corresponding with the

adductor magnus by Dugès, and with the semimembranosus by Klein; by

considering the position and attachments of these muscles one must

arrive at a different conclusion.

_MM. adductores femoris._

*119.* [*a*] _M. adductor longus_ (Fig. 82 _ad_′).

*Dugès*, sous-ilio-fémoral, n. 141.

Without further dissection this muscle is only partially visible

between the _m. sartorius_ and _vastus internus_ (Fig. 81 _ad_′). The

muscle is flat and long, and arises by a tendon from the anterior

inferior angle of the pubic symphysis (Fig. 42 _s_), partially

covered by the origin of the _sartorius_. Thence the muscle passes

between the _vastus internus_ and the _adductor magnus_, and finally

is inserted below the middle of the femur, together with the latter

muscle.

[Illustration: Fig. 82.

Muscles of the left thigh of _Rana esculenta_. Ventral surface.

_ad_′ M. adductor longus.

_ad_‴ M. adductor magnus.

_gl_ M. glutaeus.

_ip_ M. ilio-psoas.

_ra_ M. rectus fem. anticus.

_ri_′ M rectus int. major.

_s_ M. sartorius.

_st_ M. semitendinosus.

_vi_ M. vastus internus.

]

*120.* [*b*] _M. adductor magnus_ (Figs. 81, 82 _ad_‴).

*Dugès*, sous-ischio-pubi-fémoral (adductor 1 and 2), n. 142,

143.--*Klein*, _adductor magnus_ and _extensor femoris profundus_.

This muscle is visible between the _sartorius_ and _rectus internus

major_ in the anterior half of the thigh, posteriorly it is covered

by these muscles. It arises by a tendon from (_a_) the pubic

symphysis and _ischii_, and is in this position divided into two

portions, between which passes the tendon of one head of the _m.

semitendinosus_ (Fig. 83 _st_″); (_b_) a second part arises from

this same tendon (Fig. 83 _st_‴) and joins with the first part. Thus

formed the muscle passes to be inserted into the distal half of the

inner surface of the femur as far as the condyle. Near its hinder

extremity, however, it passes on to the dorsal and outer surfaces,

and thus forms a kind of muscular sheath surrounding the distal

portion of the bone.

*121.* [*c*] _M. adductor brevis_ (Figs. 81, 83 _ad_″),

and

*122.* [*d*] _M. pectineus_ (Fig. 83 _pe_).

*Dugès*, sous-pubio-fémoral (analogue of the _pectineus_, possibly of

the _add. brevis_ also).

These two small muscles which lie under cover of the _sartorius_,

_adductor longus_ and _magnus_, appear externally as one muscle, but

may easily be separated (Fig. 83). They arise close together from

the pubic symphysis, and are inserted into the inner surface of the

anterior half of the femur. They are not attached to the _adductor

magnus_.

[Illustration: Fig. 83.

Deep muscles of left thigh of _Rana esculenta_.

_ad_′ M. adductor longus.

_ad_″ M. adductor brevis.

_ad_‴ M. adductor magnus.

_ip_ M. ilio-psoas.

_pe_ M. pectineus.

_ra_ M. rectus anticus,

_ri_′ M. rectus internus major }

_ri_″ M. rectus internus minor } cut through.

_s_ M. sartorius.

_st_ Belly of m. semitendinosus.

_st_′ M. semitendinosus (cap. posticum).

_st_″ M. semitendinosus (cap. anticum).

_st_‴ Fibres passing into the adductor magnus.

_vi_ M. vastus internus.

]

*123.* _M. semitendinosus_ (Fig. 83 _st_, _st_′, _st_″).

*Dugès*, bis-ischio-tibial, n. 153.--*Zenker*, _biceps_.

This is a long thin muscle, only visible after removal of the _m.

rectus internus_, by which it is entirely covered. It arises by two

long thin tendons, one (_caput posticum_, _st_′) from the pubic

symphysis, the other (_caput anticum_, _st_″) from the hollow between

the symphysis and the acetabulum. The latter passes through a slit

in the _adductor magnus_, and so affords this muscle another point of

origin; then backwards between the _adductor magnus_ and the _rectus

internus major_ and under the lower third of the thigh, and unites

with the posterior head of origin. The muscle so formed passes into

a thin tendon, and, together with that of the _sartorius_, forms an

aponeurotic arch, connected with the fascia of the leg, under which

the tendons of the _rectus internus major_ and _minor_ pass.

(C.) Deep muscles of the thigh.

To these, in addition to the _adductor brevis_, _pectineus_, and the

portion of the _biceps_ already described, belong also the following:

[Illustration: Fig. 84.

Deep muscles of the left thigh of _Rana esculenta_. Dorsal view.

_ad_′ M. adductor brevis.

_ci_ M. coccygeo-iliacus.

_cx_ Coccyx.

_gl_ M. glutaeus.

_il_ Ileum.

_ip_ M. ilio-psoas.

_o.g._ M. obturatorius.

_qf_ M. quadratus femoris.

]

[Illustration: Fig. 85.

Left half of pelvis of _Rana esculenta_. Thigh turned forwards under

the abdomen.

_gl_ M. glutaeus.

_og_ M. obturatorius.

]

*124.* _M. ilio-psoas_ (Figs. 82, 83, 84 _ip_).

*Dugès*, intra-ilio-fémoral (*_iliacus_ and _psoas_).--*Cuvier*,

*Zenker*, *Klein*, _iliacus internus_.

The fibres of this muscle take a broad origin from the pelvic surface

of the ilium, pass outwards over the lower margin of the bone and

form a flat muscle with its apex pointed posteriorly. It crosses the

hip-joint between the _m. rectus_ and _vastus internus_, covering

the hinder part of the _m. glutaeus_, and is inserted into the outer

surface of the middle third of the femur.

*125.* _M. quadratus femoris_ (Fig. 84 _qf_).

*Dugès*, post-ilio-fémoral (Homologue of the _glutaeus maximus_), n.

197.--*Zenker*, _glutaeus minor_, p. 42.

This muscle is of a somewhat elongated, triangular shape, and

arises, under cover of the _pyriformis_, from the ilium behind the

acetabulum; it passes backwards over the joint to be inserted into

the inner and under surfaces of the femur, between the _pyriformis_

and _ilio-psoas_; below, it is in direct relation with the _adductor

brevis_.

*126.* _M. obturatorius_ (Figs. 84, 85 _og_).

*Dugès*, ischio-pubi-fémoral, n. 139 (?).--*Zenker*, _m. capsularis

femoris_.

This small muscle is deeply situated upon the hip-joint, and covered

by all the other muscles that arise from the symphysis. Its origin

from the innominate bone extends from the superior posterior angle of

the _symph. ossis ilii_ to about the middle of the pubic symphysis;

the muscle fills the space between the crest of the symphysis and the

acetabulum. The fibres converge to pass into a strong tendon, which

is inserted into the posterior and anterior surfaces of the head of

the femur. The muscle appears to represent collectively the small

rotators of human anatomy, _m. obturator externus_ and _internus_ and

the _gemelli_.

II. MUSCLES OF THE LEG.

_General description._

On the outer surface (the anterior of man), covered by the _fascia

cruris_, into which the tendon of the _extensor communis cruris_ or

_triceps_ is prolonged, are found the following muscles: nearest to

the bone, the _m. extensor cruris brevis_ (_ec_); externally, the

_m. tibialis anticus_ (_ta_) and the _m. peroneus_ (_pe_). On the

inner surface (the hinder of man) lie the _gastrocnemius_ (_gc_) and

_tibialis posticus_ (_tp_), and deeply the _flexor tarsi sup._ (_ft_).

*127.* _M. gastrocnemius_ (Figs. 80, 81, 86).

*Dugès*, bi-fémoro-plantaire, n. 159.

This important muscle arises by two tendinous heads. The one (Fig. 86

_g_′) rather nearer the middle line of the popliteal space, is

by far the stronger: it arises by a broad, tendinous expansion,

which covers the knee-joint, and is attached to the femur and the

tibio-fibula by anterior and posterior bands respectively; it ends

internally in a concave aponeurotic arch: the second head is a thin,

tendinous slip, arising from the common tendon of the _triceps_ as

this lies on the knee-joint. The muscle is thickest near its origin,

and then gradually diminishes in size as it passes backwards; the

superficial surface is convex, the deep surface is flat. By tracing

the tendons of origin, they are seen soon to unite and form a common

flat tendinous surface, broad in front, contracted behind; this

aponeurosis lies in a plane running obliquely from the deep surface

towards the superficial surface, without, however, reaching this.

The muscular fibres arise from both surfaces of the aponeurosis, the

fibres of the two sides diverging slightly as they pass backwards

so as to be inclined at a small angle to each other. The fibres

are inserted into a fibrous expansion (_Ap_) which lies on the

superficial surface of the muscle; thin and slight in front, this

aponeurosis rapidly thickens posteriorly until it passes into the

_tendo Achillis_.

[Illustration: Fig. 86.

Muscles of the right leg and foot of _Rana esculenta_. Dorsal view.

_a.h._ M. abductor hallucis.

_a.l1_ M. abductor longus digiti I.

_A.p._ Aponeurosis plantaris.

_b_ M. biceps.

_e.t._ M. extensor tarsi.

_F.l._ M. flexor digitorum III, IV, V longus.

_F.l_′. M. flexor digitorum I and II longus.

_g_ M. gastrocnemius, drawn towards the right.

_g_′ Tendon of origin of gastrocnemius.

_l.c._ Ligamentum calcanei.

_p_ M. plantaris.

_r.i._ MM. internus maior and minor.

_s.m._ M. semimembranosus.

_s.t._ M. semitendinosus.

_t.A._ Tendo Achillis.

_t.A_′. Thickening of tendo Achillis.

_t.p._ M. tibialis posticus.

_v.e._ M. vastus externus.

_v.i._ M. vastus internus.

]

*128.* _M. tibialis posticus_ (Fig. 86 _t.p._).

*Dugès*, cruro-astragalien, n. 160.

Covered by the _gastrocnemius_ this muscle occupies the whole of the

inner and upper surfaces of the tibio-fibula, from which it arises

by numerous fibres. Opposite the hinder extremity of the bone the

muscle becomes free and passes somewhat abruptly into a tendon, which

courses behind and over the _malleolus internus_, on to the dorsum

of the foot, and is here inserted into the anterior extremity of the

_astragalus_. I cannot agree with Dugès that this muscle corresponds

with the _soleus_ (Ecker).

*129.* _M. extensor cruris brevis_ (Fig. 87 _ec_).

*Dugès*, pré-fémoro-tibial, n. 154.

Covered by the strong _fascia cruris_, and lying between the _m.

tibialis anticus_ and the tibio-fibula, this muscle arises by a long

tendon from the inner condyle of the femur; it passes backwards in

the groove on the anterior extremity of the tibio-fibula, covered by

the tendon of the triceps femoris, and is then inserted by muscular

fibres into the outer surface of the tibio-fibula almost as far as

the posterior third of the bone. (Dugès regards this muscle as the

upper, i. e. anterior, portion of the _tibialis anticus_.)

*130.* _M. tibialis anticus_ (Figs. 80, 81, 87 _ta_).

*Dugès*, pré-fémoro-astragalien and pré-fémoro-calcanien, n. 156 and

157; the former he regards as one part of the _tib. anticus_, the

latter as the _peroneus longus I_.--*Zenker*, _tib. ant. biceps_.

The tendon of origin of this muscle is attached to the femur

near the tendon of the _m. extensor cruris brevis_, and to the

joint-ligaments. The muscle is placed under the _fascia cruris_, its

long tendon of origin passing beneath the tendon of the _triceps_

in a special sheath through the joint; the tendon then gains a

muscular belly which bifurcates at about the middle of the leg,

forming two muscular masses, coursing along the outer and the inner

side of the leg respectively. The latter is inserted into the dorsal

surface of the _astragalus_ by a tendon, and the former into the

dorsal surface of the _calcaneus_ in a similar manner. The muscle

extends the foot and flexes the tarsus; the latter movement is very

distinct on account of the insertion of the muscle being so near the

_hypomochlion_.

*131.* _M. flexor tarsi anterior_ (Ecker), (Fig. 87 _Ft_).

*Dugès*, ex-tibio-astragalien, n. 155.--*Zenker*, _tibialis anticus

simplex_.

This muscle arises where the insertion of the _extensor cruris

brevis_ ceases, from the middle of the under surface of the

tibio-fibula, and passes backwards between the two tendons of the _m.

tibialis anticus_ along the outer side of this muscle, to be inserted

by a tendon into the inner side of the anterior articular extremity

of the _astragalus_ and the _fascia dorsalis pedis_.

[Illustration: Fig. 87.

Muscles of leg and foot of _Rana esculenta_, seen from below.

_a.b.5_ M. abductor dig. V longus.

_ec_ M. extensor cruris brevis.

_e.d.1_ M. extensor digit I longus.

_e.d.4_ M. extensor digit IV longus.

_e.d.5_ M. extensor digit V longus.

_Ft_ M. flexor tarsi anterior.

_Ft_′ M. flexor tarsi posterior.

_g_ M. gastrocnemius.

_p_ M. peroneus.

_ta_ M. tibialis anticus.

_tF_ M. triceps femoris.

]

*132.* _M. peroneus_ (Figs. 80 _pe_, 87 _p_).

*Dugès*, génio-péronéo-calcanien, n. 158.

This is a strong muscle which is attached by a long tendon of

origin to the hinder extremity of the femur and the ligaments of

the knee-joint. The tendon runs in a sheath through the joint and

posteriorly becomes muscular; the muscle passes downwards on the

outer side of the tibio-fibula to be inserted into the _malleolus

externus_ of this bone and into the anterior extremity of the

_calcaneus_. It has an action similar to that of the _m. tibialis

anticus_.

III. MUSCLES OF THE FOOT.

A. _MUSCLES ON THE PLANTAR SURFACE._

*133.* _Aponeurosis plantaris_ (Fig. 88 _A.p._).

As already mentioned, the tendon of the _m. gastrocnemius_ is, at

the heel, continued into a strong aponeurosis, and possesses at this

place a thickening[48]. The aponeurosis is of triangular form, the

base directed towards the toes and attached to these. The lateral

borders of this triangle are continued into weaker _fasciae_, which

are attached to the two long tarsal bones. The inner border in

particular gives a strong bundle of fibres to the astragalus, and is

thus connected with the fascia of the dorsum of the foot; externally

this aponeurosis is attached to a cartilage (Dugès, os sesamoïde, _l.

c._, n. 66), situated on the plantar surface of the joint between the

_calcaneus_ and the _os metatarsi IV_.

[Footnote 48: Lehmann, Ueber den Knorpel in der Achillessehene des

Frosches, Zeitschs. f. U. Zool. XIV, p. 109. (See also literature on

cartilage, Sect. VII.)]

*134.* _Ligamentum calcanei_ (Ecker), (Figs. 88, 89 _l.c._).

If the _tendo Achillis_ be divided and the _aponeurosis plantaris_

reflected towards the toes, there is seen, arising from the posterior

head of the tibio-fibula, in its whole breadth, a ligament (between

the _tib. posticus_ and _peroneus_). This ligament forms a firm

surface on which the thickened portion of the _tendo Achillis_ may

move, and which evidently represents the tuberosity of the heel bone.

I have therefore named the entire structure the _ligamentum calcanei_.

[Illustration: Fig. 88.

Muscles of the right leg and foot of _Rana esculenta_. Dorsal view.

_a.h._ M. abductor hallucis.

_a.l_I M. abductor longus digiti I.

_A.p._ Aponeurosis plantaris.

_b_ M. biceps.

_e.t._ M. extensor tarsi.

_F.l._ M. flexor digitorum III, IV, V longus.

_F.l._′ M. flexor digitorum I and II longus.

_g_ M. gastrocnemius, drawn towards the right.

_g_′ Tendon of origin of gastrocnemius.

_l.c._ Ligamentum calcanei.

_p_ M. plantaris.

_r.i._ MM. internus maior and minor.

_s.m._ M. semimembranosus.

_s.t._ M. semitendinosus.

_t.A._ Tendo Achillis.

_t.A_′. Thickening of tendo Achillis.

_t.p._ M. tibialis posticus.

_v.e._ M. vastus externus.

_v.i._ M. vastus internus.

]

Several muscles arise from this ligament, viz.:--

*135.* _M. extensor tarsi_ (Klein), (Figs. 88, 89 _e.t._).

*Dugès*, tibio-sous-astragalien (regarded by him as the homologue of

the _tibialis posticus_).

Covered by the two succeeding muscles, this muscle arises from the

_ligamentum calcanei_, runs backwards, and is inserted into the whole

length of the plantar surface of the _astragalus_. It extends the

foot.

*136.* _M. plantaris_ (Figs. 88, 89 _p_).

This takes its origin from the _ligamentum calcanei_ to the inner

side and below the last muscle. It passes backwards and is inserted

into the dorsal surface of the _aponeurosis plantaris_.

[Illustration: Fig. 89.

Muscles of the plantar surface of foot of _Rana esculenta_, twice

natural size.

_A.p._ Aponeurosis plantaris, divided.

_ab.5._ M. abductor dig. V.

_ad.5._ M. adductor dig. V.

_ad.l.1._ M. adductor longus dig. I.

_e.t._ M. extensor tarsi.

_F.5._ M. flexor brevis dig. V.

_F.l._ M. flexor digitorum III, IV, V longus.

_F.l_′. M. flexor digitorum I, II longus.

_l.1._}

_l.2._}

_l3_ }

_l3_′ } Musculi lumbricales.

_l4_′ }

_14_″ }

_l.5._}

_l.c._ Ligamentum calcanei.

_p_ M. plantaris.

_t.A._ Tendo Achillis.

_t.A_′. Thickening in tendo Achillis.

_t.p._ M. transversus plantae posterior.

_t.p._′ M. transversus plantae anterior.

]

*137.* _M. flexor digitorum III_, _IV_, _V longus_ (Ecker),

(Figs. 88, 89 _F.l._).

*Dugès*, péronéo-sous-phalangettien (_flexor dig. longus_).

This muscle also arises from the _ligamentum calcanei_ to the outer

side of the _m. plantaris_. It runs backwards over the _aponeurosis

plantaris_, and at the free margin of this passes somewhat suddenly

into a strong tendon, which traverses an aponeurotic canal, formed by

fibres of the aponeurosis passing above and below the tendon, from

the cartilaginous enlargement in the tendon to the median border of

the aponeurosis. The tendon forms three slips for the fifth, fourth,

and third toes: these are slightly attached to the aponeurosis,

but reach beyond its free border to be inserted into the terminal

phalanges of the toes mentioned, and are held down by fibrous bands.

(The small _m. lumbricales_ arising from these tendons are described

below.)

*138.* _M. flexor digitorum I_, _II longus_ (Ecker), (Figs. 88, 89

_F.l_′.).

The tendons, by means of which the terminal phalanges of the first

and second toes are flexed, arise directly from the posterior border

of the aponeurosis.

*139, 140.* _MM. transversi plantae_ (Ecker), (Fig. 89 _t.p._,

_t.p_′.).

The movements of the aponeurosis and through it the movements of

the tendons attached to it are influenced not only by the _m.

gastrocnemius_ and _plantaris_, but also by two other muscles, which

together have been described as the _flexor dig. longus internus_

(Klein), or as the tarso-sous-phalangettien (Dugès, 221).

*a.* _M. transversus plantae posterior_ (_t.p._).

This takes its origin from the plantar cartilage, widens as it

courses backwards and inwards to be inserted into the dorsal surface

of the _aponeurosis plantaris_; from this the tendons for the first

and second toes (_flexor dig. longus I_ and _II_) arise, thus there

is no direct connection between the muscular fibres of the two

muscles.

*b.* _M. transversus plantae anterior_ (_t.p_′.).

This muscle is covered by the last; broader anteriorly, it arises

from the calcaneum, and is inserted to the inner side of the last

muscle into the dorsal surface of the aponeurosis. This muscle, like

that just described, is not continued directly into the tendon of the

flexor of the toes, they cannot therefore be regarded as flexors of

the toes.

*141* to *149*. _Musculi lumbricales_ (Fig. 89 _l.1–5_).

(A.) From the posterior border of the _aponeurosis plantaris_ arise:--

(1) _M. lumbricalis digiti I_ (_l.1_).

*Dugès*, tendini-sous-phalangien de pouce, n. 185.

This is inserted into the anterior extremity of the proximal phalanx.

(*2*) _M. lumbricalis digiti II_ (_l.2_).

*Dugès*, tendini-sous-phal. du I doigt, n. 186 (and 187), double

according to Dugès.

Similarly inserted.

(*3* and *4*) _MM. lumbricales digiti III_ (_l.3_).

*Dugès*, tendini-sous-phal. du III doigt, n. 188, single according to

Dugès.

One of these (_l3_) is inserted into the anterior extremity of the

basal phalanx, the other (_l3_′) into a corresponding position on the

middle phalanx.

(*5*) _M. lumbricalis digiti IV._

*Dugès*, tendini-sous-phal. du IV doigt.

Inserted into the anterior extremity of the basal phalanx.

(B.) Arising from the tendons of the _flexor longus III_, _IV_, _V_:

(*6* and *7*) _MM. lumbricales digiti IV_ (_l.4_′. and _l.4_″.).

*Dugès*, tendini-sous-phalanginien du IV doigt, n. 201, 202.

The first (_l.4_′) is inserted into the posterior end of the basal

phalanx, the second (_l.4_″) in a similar position into the second

phalanx.

(*8*) _M. lumbricalis digiti V_ (_l.5._).

*Dugès*, tendini-sous-phalangien du V doigt, n. 203.

This arises from the tendon for the fifth toe, runs backwards as a

thin tendon alongside that of the flexor, and is inserted into the

middle phalanx.

_Special Muscles of the Separate Toes._

(I.) Muscles of the rudimentary great toe.

*150.* _M. abductor hallucis_ (Figs. 88 _a.h._, 90).

*Dugès*, tibio-sous-tarsien, n. 163.

This muscle arises from the inner border of the _aponeurosis

plantaris_, at the spot where the _m. plantaris_ is inserted, it may

therefore almost be regarded as a continuation of this muscle; it is

inserted into the anterior border of the rudimentary toe.

(II.) Muscles of the great and first toes.

*151.* _M. adductor longus digiti I_ (Figs. 89, 90 _ad.l1_).

*Dugès*, calcanéo-scaphoidien, n. 164.

This powerful muscle arises from the plantar surface of the calcaneum

and astragalus; its fibres converge towards a tendon lying in its

middle, which becomes free towards its posterior end; the tendon

passing inwards under the _ligamentum tarsi transversum_ through

a groove on the hinder extremity of the astragalus in an arched

direction, to be inserted into the _os naviculare_, which bears the

rudimentary great toe and the _os metatarsi I_.

(III.) Muscles of the first toe.

*152.* _M. abductor longus digiti I_ (Figs. 88 _al.1_, 90 _a.b.1_).

It takes its origin from the inner border of the _aponeurosis

plantaris_, and lies on the plantar surface of the _abductor

hallucis_. Its tendon runs in the hollow which the great toe forms,

and is inserted into the inner side of the _os metatarsi I_.

*153.* _M. flexor brevis digiti I_ (Fig. 90 _F.b.1_).

A small, thin muscle, which arises from the anterior extremity of

the _os metatarsi II_, and is inserted into the basal phalanx of the

first toe. In this small muscle, and also in the _m. lumbrical. dig.

V_ (Fig. 89 _l.5._), the nerve distribution may be very well seen.

*154.* _M. opponens dig. I_ (Fig. 90 _op_).

*Dugès*, sous-metatarso-phalangien du pouce, n. 192.

This is situated to the inner side of the preceding muscle, arises

from the _os metatarsi II_ somewhat narrow, widens in a fan-like

manner towards the _os metatarsi I_, into the anterior half of which

it is inserted.

(IV.) Muscles of the second toe.

*155.* _M. flexor metatarsi dig. II_ (Fig. 90 _F.m.2_).

*Dugès*, sous-tarso-metatarsien du II doigt.--*Klein*, extensor

metatarsi.

By a narrow origin from the hinder end of the calcaneum, posteriorly

this muscle becomes broader and is inserted, somewhat thinned out,

into the plantar surface of the second _os metatarsi_.

*156.* _M. flexor dig. II proprius_ (Fig. 90 _F.p.2_).

*Dugès*, sous-metatarso-phalangien, n. 194.

It arises from the plantar surface of the _os metatarsi II_, and

is inserted by a thin tendon into the plantar surface of the first

phalanx.

Dugès describes this and the analogous muscles of the third, fourth,

and fifth toes (_F.p.3_, _4_, _5_) as the _interossei plantares_, a

nomenclature which, to me, does not appear correct, as these muscles

lie entirely on the plantar surface, and do not adduct or abduct the

toes, but flex them.

(V.) Muscles of the third toe.

*157.* _M. flex. metatarsi dig. III_ (Ecker), (Fig. 90 _F.m.3_).

*Dugès*, sous-calcaneo-métatarsien du III doigt, n. 174.--

*Klein*, _extensor metatarsi_.

Like the corresponding muscle of the second toe (n. 155), this

arises by a small tendon from the hinder extremity of the calcaneum,

runs backwards and is inserted into the plantar surface of the _os

metatarsi III_. The greatest portion of the muscle lies beneath

the _m. transversus metatarsi_ (_tm_), the most anterior portion,

however, lies above this. Dugès has described this latter portion as

a special muscle (n. 176, metatarso-métatarsien du III), the same

description holds good for the second toe (n. 177, Dugès).

*158.* _M. flexor dig. III proprius_ (Ecker), (Fig. 91 _Fp3_).

*Dugès*, sous-métatarso-phalangien du III doigt, n. 195.

Like the corresponding muscle of the second toe, this arises from

the plantar surface of the _os metatarsi III_, and is inserted by a

thin tendon into the plantar surface of the anterior extremity of the

first phalanx.

[Illustration: Fig. 90.

Muscles of plantar surface of foot of _Rana esculenta_.

_a.b.1_ M. abductor longus digiti I.

_ab5_ M. abductor digiti V.

_ad.l.1_ M. adductor longus digiti I.

_a.h._ M. abductor hallucis.

_F.b.1_ M. flexor brevis digiti I.

_F.m.2_, _3_, _4_ M. flexores metatarsi II, III, IV.

_F.p.2_, _3_, _4_, _5_ MM. flexor. proprii digit. 2, 3, 4, 5.

_F.ph.3_, _4_, _4_′, _5_ MM. flexor. phalang. digit. 3, 4, 5.

_op_ M. opponens digiti I.

_t.m_′ }

_t.m_″ } MM. transversi metatarsi 1, 2, 3.

_t.m_‴ }

]

*159.* _M. flexor phalangum proprius digiti III_ (Ecker), (Fig. 91

_F.ph.3_).

*Dugès*, phalango-phalangien, n. 204.

This arises by tendon from the plantar surface of the basal phalanx,

and is inserted into the anterior extremity of the terminal phalanx.

(VI.) Muscles of the fourth toe.

*160.* _M. flexor metatarsi dig. IV_ (Fig. 91 _F.m.4_).

*Dugès*, métatarso-métatarsien, n. 175.

Similarly to the corresponding muscle of the third toe, this one

arises from the hinder extremity of the calcaneum, and is inserted

into the anterior portion of the plantar surface of the _os metatarsi

IV_, but not into the greater portion of this surface, as in the

second and third _os metatarsi_.

*161.* _M. flexor dig. IV proprius_ (Fig. 91 _Fp4_).

*Dugès*, sous-métatarso-phalangien du IV doigt (_interosseus

plantaris_), n. 196.

The origin and insertion are similar to those of the muscles of the

second and third toes.

[Illustration: Fig. 91.

Muscles of plantar surface of foot of _Rana esculenta_.

_a.b.1_ M. abductor longus digiti I.

_ab5_ M. abductor digiti V.

_ad.l.1_ M. adductor longus digiti I.

_a.h._ M. abductor hallucis.

_F.b.1_ M. flexor brevis digiti I.

_F.m.2_, _3_, _4_ M. flexores metatarsi II, III, IV.

_F.p.2_, _3_, _4_, _5_ MM. flexor. proprii digit. 2, 3, 4, 5.

_F.ph.3_, _4_, _4_′, _5_ MM. flexor. phalang. digit. 3, 4, 5.

_op_ M. opponens digiti I.

_t.m_′ }

_t.m_″ } MM. transversi metatarsi 1, 2, 3.

_t.m_‴ }

]

*162.* _M. flexor brevis dig. IV._

A small muscle, which arises with the _m. flexor brevis_ of the fifth

toe, from enlargement of _aponeurosis plantaris_, and is inserted

into the basal phalanx of the fourth toe.

*163.* _M. flexor phalangum proprius dig. IV anterior_ (Fig. 91

_F.ph.4_).

*Dugès*, phalango-phalanginien, n. 205.

It resembles the corresponding muscle of the third toe.

*164.* _M. flexor phalangum proprius posterior_ (Fig. 91 _F.ph.4_′).

*Dugès*, phalangino-phalangettien, n. 209.

This extends from the plantar surface of the second phalanx to the

anterior extremity of the third.

(VII.) Muscles of the small toe.

*165.* _M. abductor dig. V_ (Fig. 91 _a.b.5_).

*Dugès*, calcanéo-ex-métatarsien du digitule, n. 169.

This muscle arises from the hinder extremity of the calcaneum, and is

inserted into the outer surface of the fifth metatarsal.

*166.* _M. adductor dig. V_ (Fig. 89 _a.d.5_).

*Dugès*, sous-tarso-in-phalangien, n. 191.

This arises from the cartilage of the plantar aponeurosis, and is

inserted into the hinder end of the fifth metatarsal.

*167.* _M. flexor brevis dig. V_ (Fig. 89 _F.5_).

*Dugès*, sous-tarso-ex-phalangien, n. 190.

Arising from the same place as the preceding, it extends to the basal

phalanx.

*168.* _M. flexor dig. V. proprius_ (Fig. 91 _F.p._5).

(Analogues of Muscles, n. 156, 158, 161; see note at n.

156).--*Dugès*, sous-métatarso-phalangien, n. 197 (*_inteross.

plant._).

This muscle arises from the plantar and inner surfaces of the fifth

metatarsal, and is inserted into the basal phalanx.

*169.* _M. flexor phalangum proprius_ (Fig. 91 _F.ph.5_).

*Dugès*, phalango-phalanginien, n. 206.

Its origin and insertion are as in the corresponding muscles of the

second, third, and fourth toes, n. 159, 163, 164, 169.

*170* to *172*. _MM. interossei_ (Fig. 91 _t.m._).

*Dugès*, 1, 2, and 3, intermétatarsien, n. 170, 171, 172.

These are three muscles with their fibres arranged transversely,

which occupy the anterior portion of the spaces between the

metatarsals, and approximate these. The first arises (_tm_′) from the

edge of the plantar surface of the first metatarsal, and is inserted

into the corresponding part of the second: the second (_tm_″) passes

from this point to the third, and the third (_tm_‴) to the fifth.

B. _MUSCLES OF THE DORSAL SURFACE._

*173.* _M. flexor tarsi posterior_ (Ecker), (Fig. 92 _F.t._).

*Dugès*, péronéo-sus-astragal, n. 161.

This is a strong muscle, which has a narrow origin from the outer

side of the tibio-fibula, directly over the distal extremity, and is

inserted into the dorsal surface of the astragalus from the middle to

the distal extremity.

Muscles of the first toe.

*174.* _M. extensor longus dig. I_ (Fig. 92 _e.l.1_).

*Dugès*, calcanéo-sus-métatarsien du pouce, n. 166.

This lies to the outer side of and near the foregoing muscle; it

arises by a long, strong head from the middle of the calcaneum, runs

obliquely backwards and inwards to unite with a small head which

arises from the common epiphysis of the astragalus and calcaneum, in

common with the _m. extensor dig. II_. The muscle so formed passes

into a flat tendon to be inserted into the basal phalanx of the first

toe.

*175.* _M. extensor brevis dig. I_ (Fig. 92 _e.b.1_).

A small, short muscle which arises from the _os naviculare_ and is

inserted into the dorsal surface of the first metatarsal.

*176.* _M. abductor brevis dig. I_ (Fig. 92 _a.b.1_).

*Dugès*, ex-tarso-métatarsien du pouce, n. 168.

This small muscle is situated near the preceding; it arises from the

first bone of the supplemental toe, and is inserted into the inner

side of the first metatarsal.

Muscles of the second toe.

*177.* _M. extensor dig. II longus_ (Fig. 92 _e.l.2_).

*Dugès*, calcanéo-sus-phalangien du II doigt, n. 182.

This arises from the calcaneum, in common with the _m. extensor

longus_ of the third toe, and is inserted into the basal phalanx of

the second toe by a tendon.

*178.* _M. extensor dig. II brevis_ (Fig. 92 _e.b.2_).

*Dugès*, astragalo-sus-phalangien du II, n. 183.

This is partly covered by the preceding muscle: it arises from the

astragalus, and unites with the tendon of the preceding muscle.

Muscles of the third toe.

*179.* _M. extensor dig. III longus_ (Fig. 92 _e.l.3_).

*Dugès*, sus-astragalo-phalangien du médius, n. 199.

This arises with the _extensor longus_ of the second toe from the

calcaneum: its tendon unites with that of the following muscle.

*180.* _M. extensor dig. III brevis_ (Fig. 92 _e.b.3_).

*Dugès*, astragalo-sus-phalangien du médius, n. 181.

This muscle arises from the astragalus, and is inserted with the

tendon of the preceding muscle into the dorsal surface of the basal

phalanx.

[Illustration: Fig. 92.

Dorsal view of muscles of foot of _Rana esculenta_.

_a.b.1_ M. abductor brevis dig. I.

_e.b.1_ M. extensor brevis dig. I.

_e.b.2_ M. extensor brevis dig. II.

_e.b.3_ M. extensor brevis dig. III.

_e.b.4_ M. extensor brevis 1 dig. IV.

_e.b.4_′ M. extensor brevis 2 dig. IV.

_e.b.5_ M. extensor brevis dig. V.

_e.l.1_ M. extensor longus dig. I.

_e.l.2_ M. extensor longus dig. II.

_e.l.3_ M. extensor longus dig. III.

_e.l.4_ M. extensor longus dig. IV.

_e.l.5_ M. extensor longus dig. V.

_F.t_ M. flexor tarsi posterior.

_i.2_ to _i.10_ MM. interossei.

]

Muscles of the fourth toe.

*181.* _M. extensor longus dig. IV_ (Fig. 93 _e.l.4_).

*Dugès*, péroneo-sus-phalangien du IV, n. 178.

A narrow muscle arising in common with the _flexor tarsi inf._, but

external to it. It is inserted into the tendinous expansion on the

dorsum of the toe.

*182.* _MM. extensores dig. IV breves_ (Fig. 93 _e.b.4_ and

_e.b.4_′).

*Dugès*, calcanéo-sus-phalangien and sus-calcanéo-phalanginien, n.

179 and 198.

These two small muscles arise from the calcaneum: one is inserted

into the basal phalanx, the other, by means of a long thin tendon,

passes along with the tendon of the corresponding _m. interosseus_ to

the second phalanx, to which it is attached.

Muscles of the fifth toe.

*183.* _M. extensor dig. V longus_ (Fig. 93 _e.l.5_).

*Dugès*, calcanéo-sus-métatarsien, n. 165.

A strong muscle which covers almost the whole length of the

calcaneum; it arises from the anterior extremity of this bone,

runs backwards to the outer side of the dorsum of the foot, and is

attached to the outer surface of the fifth metatarsal.

*184.* _M. extensor dig. V brevis_ (Fig. 93 _e.b.5_).

*Dugès*, calcanéo-sus-phalangien du V, n. 180.

This is a long thin muscle, which, together with the muscle last

described, arises from the calcaneum and is inserted into the basal

phalanx of the fifth toe.

*185.* _M. abductor dig. V brevis_ (Fig. 93 _i.10_).

This, the last _m. interosseus_, arises from the outer side of the

fifth metatarsal, and is inserted into the hinder extremity of the

second phalanx of the same toe.

*186* to *195*. _MM. interossei dorsales_ (Fig. 93 _i.1_ to _i.10_).

First toe.

(1) The analogue of the first _m. interosseus_ is the _m. abductor

brevis dig. I_ (_a.b.1_).

(2) The second _m. interosseus_ (_i.2_) arises from the outer side

of the first metatarsal, and is inserted into the outer surface of

the second phalanx of the same toe, dorsally to the _m. opponens_ and

_flexor brevis dig. I_.

Second toe.

(3) The third _m. interosseus_ (_i.3_) takes origin from the anterior

part of the outer surface of the first metatarsal by a thin tendon;

it is inserted into the inner surface of the second phalanx by a

similar tendon.

(4) The fourth _m. interosseus_ (_i.4_) arises from the outer side of

the second metatarsal, and is inserted into the outer surface of the

second phalanx.

Third toe.

(5) The fifth _m. interosseus_ arises from the outer surface of the

second metatarsal, and is inserted by a thin tendon into the inner

surface of the second phalanx of the same toe.

(6) The sixth arises from the outer surface of the third metatarsal,

and is inserted laterally by a thin tendon into the hinder extremity

of the second phalanx of the same toe.

[Illustration: Fig. 93.

Dorsal view of muscles of foot of _Rana esculenta_.

_a.b.1_ M. abductor brevis dig. I.

_e.b.1_ M. extensor brevis dig. I.

_e.b.2_ M. extensor brevis dig. II.

_e.b.3_ M. extensor brevis dig. III.

_e.b.4_ M. extensor brevis 1 dig. IV.

_e.b.4_′ M. extensor brevis 2 dig. IV.

_e.b.5_ M. extensor brevis dig. V.

_e.l.1_ M. extensor longus dig. I.

_e.l.2_ M. extensor longus dig. II.

_e.l.3_ M. extensor longus dig. III.

_e.l.4_ M. extensor longus dig. IV.

_e.l.5_ M. extensor longus dig. V.

_F.t_ M. flexor tarsi posterior.

_i.2_ to _i.10_ MM. interossei.

]

Fourth toe.

(7) The seventh _m. interosseus_ (_i.7_) arises by a thin tendon from

the hinder extremity of the astragalus, and by fleshy fibres from

the hinder extremity of the fourth metatarsal; it is inserted by a

thin tendon into the inner surface of the second phalanx of the same

toe.

[Illustration: Fig. 94.

Dorsal view of muscles of foot of _Rana esculenta_.

_a.b.1_ M. abductor brevis dig. I.

_e.b.1_ M. extensor brevis dig. I.

_e.b.2_ M. extensor brevis dig. II.

_e.b.3_ M. extensor brevis dig. III.

_e.b.4_ M. extensor brevis 1 dig. IV.

_e.b.4_′ M. extensor brevis 2 dig. IV.

_e.b.5_ M. extensor brevis dig. V.

_e.l.1_ M. extensor longus dig. I.

_e.l.2_ M. extensor longus dig. II.

_e.l.3_ M. extensor longus dig. III.

_e.l.4_ M. extensor longus dig. IV.

_e.l.5_ M. extensor longus dig. V.

_F.t_ M. flexor tarsi posterior.

_i.2_ to _i.10_ MM. interossei.

]

(8) The eighth _m. interosseus_ (_i.8_) takes its origin from the

outer surface of the fourth metatarsal and from the inner surface of

the fifth: it is inserted into the hinder extremity of the second

phalanx.

Fifth toe.

(9) The ninth _m. interosseus_ (_i.9_) arises from the inner surface

of the fifth metatarsal, and is inserted into the hinder extremity of

the second phalanx.

(10) The tenth _m. interosseus_ has already been described as the

_abductor digiti V brevis_ (Fig. 94 _i.10_).

If we take the fourth toe as the axis of movement, the first, third,

fifth, seventh, eighth, and tenth _mm. interossei_ will abduct from

an imaginary line which runs through this toe. The second, fourth,

sixth, and ninth will adduct towards this line. If we compare them

with those of man (in whose case the second toe affords the central

line), those of the first group must be regarded as _mm. interossei

dorsales_, of the latter as _plantares_. Dugès holds all these

_interossei_ (with the exception of the first, _abductor brevis dig.

I_) to be _mm. interossei dorsales_ (métatarso-sus-phalangettiens, n.

210, 211, 212, 214, 215, 216, 218, 219). Those which I have described

as _flexores proprii digitorum_, he regards as _interossei volares_.

V. MUSCLES OF THE SKIN.

*196.* _M. cutaneus pectoris_ (Fig. 95 _cp_).

*Dugès*, abdomino-guttural, n. 53.--*Klein*,

abdomino-cutaneus.--*Zenker*, subcutaneus pectoris.

This is an elongated quadrangular muscle, attached by its hinder

margin to the side of the xiphisternum, and to the superficial

surface of the aponeurosis of the _m. obliquus externus_. The fibres

course forwards and slightly outwards to be inserted, at a very acute

angle, into the skin: lying between the two muscles and attached

to them on either side is a thin fascia, which, together with the

muscles and the _septum thoracicum_, assists in closing a triangular

space between these structures and the deeper muscles.

The thinness and transparency of this muscle render it especially

suitable for the investigation of nerve-terminations[49].

[Footnote 49: Kölliker, Mikrosk. Anatomie, 1866, vol. II. 1,

p. 247; Reichert, Müller’s Archiv, 1851, p. 29, Pl. I; Kölliker,

Untersuchungen über die letzten Endigungen der Nerven in den Muskeln

des Frosches; Leipzig, Engelmann, 1862.]

[Illustration: Fig. 95.

Pectoral region of _Rana esculenta_.

_cp_ M. cutaneus pectoris.

_p_″ Port. abdominalis of the m. pectoralis.

]

*197.* _M. cutaneus dorsi_ (Fig. 96 _cd_).

*Dugès*, pubio-dorso-cutané, n. 56.--*Zenker*, _cutaneus iliacus_.

This muscle has a narrow origin in the space between the muscles of

the belly and of the thigh, and is here inserted into the fasciae,

which are attached to the pubic symphysis; it passes inwards and

upwards to the outer side of the anterior insertion of the _m. rectus

femoris anticus_, then widens and is inserted by diverging fibres

into the inner surface of the skin of the hinder portion of the back.

*198.* _M. coccygeo-cutaneus._

*Dugès*, coccy-dorso-cutanés, n. 57.

The muscular fibres which Dugès has described under this name, lie

covered by the _m. pyriformis_, behind the _m. coccygeo-iliacus_,

etc., and above the muscles of the rectum. They arise from the hinder

extremity of the urostyle: they appear to be connected with the

rectal muscles, and pass outwards to be attached to the skin.

Among the cutaneous muscles may perhaps be included the _m.

submaxillaris_ (p. 62).

The fibrils passing from the flexor tendons to the warty bodies on

the plantar surface of the toes are described[50] with the skin

(p. 373).

[Footnote 50: Compare Klein, Beiträge zur Anatomie der ungeschwänzten

Batrachier (R. temporaria, L.): Jahreshefte des Vereins für

vaterländische Naturkunde in Württemberg, 1850, p. 72.]

[Illustration: Fig. 96.

Hind portion of back and thigh of _Rana esculenta_.

_cd_ M. cutaneus dorsi.

_cx._ Coccyx.

_gl._ M. glutaeus.

_H_ Skin reflected to left side.

_oc_ M. obliquus externus.

_rf_ M. rectus femoris anterior.

_sm_ M. semimembranosus.

_v.e_ M. vastus externus.

]

SECTION III.

THE NERVOUS SYSTEM.

THE NERVOUS SYSTEM.

LITERATURE.

*Ahlborn, F.*, Ueber die Bedeutung der Zirbeldrüse. Zeit. f. wiss.

Zool. 1884. Vol. XL, pp. 331–337.

*Allen, H.*, The Spinal Cord in Batrachia and Reptilia. Proc. Acad.

Nat. Sci. Philadelphia. 1883, pp. 56–57.

*Arndt, A. W.*, Untersuchungen über die Ganglienkörper des Nervus

sympathicus. Arch. f. mikrosk. Anat. 1874. Vol. X, pp. 208–241.

*Arndt, R.*, Untersuchungen über die Endigung der Nerven in den

quergestreiften Muskelfasern. Arch. f. mikrosk. Anat. 1873.

Vol. IX, p. 481.

*Arndt, R.*, Untersuchungen über die Ganglienkörper der

Spinalganglien. Arch. f. mikrosk. Anat. 1875. Vol. XI, p. 140.

*Arnold, J.*, Zur Histologie der Lungen des Frosches. Virchow’s Arch.

1863. Vol. XXVIII, p. 433.

*Arnold, J.*, Histologische Verhältnisse des Frosch-Sympathicus.

Centralbl. f. d. med. Wiss. 1864, p. 657.

*Arnold, J.*, Ueber die feineren histologischen Verhältnisse der

Ganglienzellen in dem Sympathicus des Frosches. Virchow’s Arch.

1865. Vol. XXXII, p. 1.

*Arnold, J.*, Die Spinalfasern im Sympathicus des Frosches. Arch. f.

Anat. und Physiol. 1866, p. 398.

*Arnold, J.*, Das Gewebe der organischen Muskeln. Leipzig, 1869; and

Chap. IV in Stricker’s Handbook.

*Arnstein, C.*, and *Gonjaew*, *K.*, Ueber die Nerven des

Verdauungskanals. Bericht. f. Physiol. u. Histologie. Mittheilung.

aus. d. 4 Vers. wissensch. Naturforsch. zu Kasan. Pflüger’s Arch.

d. ges. Physiol. 1874. Vol. VIII, pp. 614–615.

*Aubert, H.*, Die Innervation der Kreislaufsorgane, Hermann’s

Handbuch der Physiologie. Leipzig, 1880. Vol. IV, Pt. I, p. 377.

*Axmann*, De Gangliorum systematicis structura persitiori eiusque

functionibus. Berolini, 1847.

*Axmann*, Beiträge zur mikroskopischen Anatomie und Physiologie des

Ganglien-nervensystems. 1853, p. 20.

*Baculo, B.*, Nuove ricerche intorno l’apparechio ganglionare

intrinseco dei cuori linfatici. Naples, 1885.

*Balfour, F. M.*, Treatise on Comparative Embryology. London, 1880.

*Balfour, F. M.*, Handbuch der vergleichenden Embryologie. Aus dem

Englischen von B. Vetter. Jena, 1880.

*Beale, L. S.*, On the Structure and Formation of the so-called

Apolar, Unipolar, and Bipolar Nerve-cells of the Frog. Phil.

Trans. 1863, p. 543. (Hyla arborea.)

*Beale, L. S.*, Further observations in favour of the view that

nerve-fibres never end in voluntary muscle. Proc. Roy. Soc. 1863.

Abstract in Quart. Journ. Micros. Sci. 1863. Vol. XII, p. 668.

*Beale, L. S.*, New observations upon the structure, etc. of certain

nervous centres. Proc. Roy. Soc. 1860. Vol. III.

*Beale, L. S.*, On the Distribution of Nerves to the elementary

fibres of Striped Muscle. Phil. Trans. 1864, p. 611.

*Beale, L. S.*, Of very fine nerve-fibres ramifying in certain

fibrous tissues and trunks, and plexuses consisting entirely

of very fine nerve-fibres in the bladder of the frog. Beale’s

Archives of Medicine, 1864. Vol. IV, pp. 19–251.

*Beard, J.*, The Ciliary or Motor-oculi ganglion and the Ganglion

of the ophthalmicus profundus in Sharks. Anatom. Anzeiger. 1887.

Vol. II, p. 565.

*Beck, K.*, Zur Kenntniss der Herznerven. Arch. f. mik. Anat. 1885.

Vol. XXIV, pp. 11–19.

*Bellonci, G.*, Intorno alla struttura e alle connessioni dei

lobi olfattorii negli arthropodi superiori e nei vertebrati.

Atti Accad. Lincei Ann. 279 Mem. Accad. Bologna, Vol. XIII,

pp. 555–564: and in Arch. Ital. Biol. 1883; Vol. III, pp. 191–196.

*Bellonci, G.*, Sulla terminazione centrale del nervo ottico nei

mammiferi. Mem. Accad. Bologna, Vol. VI, pp. 199–205: and in Arch.

Ital. Biol. 1884, Vol. VI, pp. 405–411.

*Bidder, F. H.*, Die Endigungsweise der Herzzweige des N. vagus beim

Frosch. Arch. f. Anat. u. Physiol. 1839, p. 1.

*Bidder, F. H.*, Zur Lehre von dem Verhältniss der Ganglienkörper

zu den Nervenfasern. Nebst einem Anhange von Dr. A. W. Volkmann.

Leipzig, 1847.

*Bidder, F. H.*, Zur Lehre vom Verhalten der Ganglienkörper, u. s. w.

Leipzig, 1847.

*Bidder, F. H.*, Ueber functionelle verschiedene und räumlich

getrennte Nerven-centra im Froschherzen. Arch. f. Anat. u.

Physiol. 1852, p. 163.

*Bidder, F. H.*, Endigungsweise der Herzzweige des N. vagus beim

Frosch. Arch. f. Anat. und Physiol. 1868, p. 1.

*Bidder, F. H.*, Zur näheren Kenntniss des Froschherzens und seiner

Nerven. Arch. f. Anat. u. Physiol. 1866, p. 1.

*Bidder, F. H.*, and *Kupffer*, Untersuchungen über die Textur des

Rückenmarks. Leipzig, 1857.

*Bidder, F. H.*, and *Volkmann*, *A. W.*, Die Selbständigkeit des

sympathischen Nervensystems, durch anatomische Untersuchungen

nachgewiesen. Leipzig, 1842.

*Bidder* and *Gregory*, Beiträge zur Physiologie der Herzbewegung

beim Frosche. Dorpat, 1865.

*Bikfalvi, K.*, Ueber die Hornscheide der markhaltigen Nervenfasern.

Orvoster-meszettudomanyi Ertesito, 1884, p. 133. Abstract in

Centralbl. f. d. med. Wiss. 1886, p. 34.

*Biroege, E. A.*, Die Zahl der Nervenfasern u. der motor.

Ganglienzellen im Rückenmark des Frosches. Arch. f. Anat. u.

Physiol. 1882. Physiol. Abth., p. 435.

*Bischoff*, Nervi accessorii Willisii anatomia et physiologia.

Heidelberg, 1832.

*Blasius, G.*, Gerardi Blasii Anatomia Animalium. Amstelodami, 1681.

*Blattmann, A.*, Mikrosk. anatom. Darstellung der Centralorgane des

Nervensystems bei den Batrachiern. Dissert. Zürich, 1850.

*Boll, F.*, Studj sulle immagini microscopiche della fibra nervosa

midollare. Atti della R. accademia dei Lincei, 1876–1877. Vol. II.

*Boll, F.*, Ueber Zersetzungsbilder der markhaltigen Nervenfasern.

Arch. f. Anat. u. Physiol. 1877, p. 288.

*Bojanus*, Testudinis anatomia. 1819.

*Bosse*, De ganglior. spinal. vi in nutr. radic. poster. nervor.

spinal. Dissert. 1859.

*Boveri, T.*, Beiträge zur Kenntniss der Nervenfasern. Abhandl. d.

bayr. Akad. d. Wiss. 1885. Vol. XV, p. 421.

*Bremer, D. L.*, Die Nerven der Capillaren, der kleinen Arterien und

Venen. Arch. f. mik. Anat. Vol. XXI. 1882, p. 663.

*Bremer, L.*, Ueber die Muskelspindeln sowie über Structur,

Neubildung u. Innervation der quergestreiften Muskelfaser.

Waldeyer’s Arch. 1883. Vol. XXII, p. 2.

*Bremer, L.*, Ueber die Endigungen der markhaltigen und marklosen

Nerven im quergestreiften Muskel. Arch. f. mikrosk. Anat. 1882.

Vol. XXI, p. 165.

*Brenner, A.*, Ueber das Verhältniss des nervus laryngeus inferior

vagi zu einigen Aortenvarietäten des Menschen und zu dem

Aortensystem der durch Lungen athmenden Wirbelthiere überhaupt.

His u. Braune’s Arch. 1883, pp. 373–397.

*Budge, J.*, Ueber den Verlauf der Nervenfasern im Rückenmark des

Frosches. Arch. f. Anat. u. Physiol. 1844, p. 160.

*Budge, J.*, Ueber die Bewegung der Iris. (Concerning ganglia of

motor-oculi, p. 36.)

*Budge, J.*, Wagner’s Handwörterb. d. Physiologie. 1846. Vol. III,

p. 451.

*Burdon-Sanderson, J.*, Circulation of the Blood. Handbook for the

Physiological Laboratory. London, 1873.

*Calberla, E.*, Ueber die Endigungsweise der Nerven in den

quergestreiften Muskeln bei Amphibien. Zeit. f. wiss. Zool. 1874.

Vol. XXIV, p. 164.

*Calberla, E.*, Studien über die Entwicklung der quergestreiften

Muskeln und Nerven der Amphibien und Reptilien. Arch. f. mik.

Anat. 1875. Vol. XI, p. 442.

*Carus*, Versuch einer Darstellung des Nervensystems und Gehirns.

Leipzig, 1814.

*Champness, F.*, The septum atrium of the Frog and Rabbit. Journ. of

Anat. and Physiol. 1874.

*Christmas-Dirckinck-Holmfeld, J.*, Experimentaluntersuchungen über

den Bau der Regio olfactoria. Nord. med. ark. 1883. Vol. XV, No. 3.

*Ciaccio, G. V.*, On the nerves of the Cornea, and their distribution

in the corneal tissue of man and animals. Proc. Roy. Soc. 1863.

Abstract in Trans. Micros. Soc. London, 1863. Vol. XI, pp. 77–93.

*Ciaccio, G. V.*, On the distribution of nerves to the skin of the

frog, with physiological remarks on the ganglia connected with the

cerebro-spinal nerves. Quart. Journ. Micros. Sci. 1864. Vol. IV,

pp. 15–31.

*Ciaccio, G. V.*, Intorno alla minuta fabbrica delle pelle Rana

esculenta. Giornale di Scienze naturali ed economiche. Palermo,

1866. Vol. II, p. 103.

*Clarke, J. Lockhart*, Ueber den Bau des Bulbus olfactorius und der

Geruchsschleimhaut. Zeit. f. wiss. Zool. 1862. Vol. XI, p. 31.

*Cornil* and *Ranvier*, A Manual of Pathological Histology. London,

1880, p. 359, Fig. 199 (Ganglia of heart).

*Courvoisier, G.*, Beobacht. über den sympathischen Gränzstrang.

Arch. f. mik. Anat. 1866. Vol. II, p. 13.

*Courvoisier, L. G.*, Ueber die spinalen und sympathischen Zellen des

Frosches. Centralbl. f. d. med. Wiss. 1867, p. 897.

*Courvoisier, G.*, Ueber die Zellen der Spinalganglien, sowie des

Sympathicus beim Frosch. Arch. f. mikrosk. Anat. 1868. Vol. IV,

p. 125.

*Cuvier*, Leçons d’anatomie comparée. 2nd Edit. Vol. II.

*Cyon*, Ueber die Nerven des Peritoneum. Berichte über die

Verhandlungen d. königl. sächs. Gesellsch. d. Wissensch. zu

Leipzig. Math.-physik. Classe. 1868. Vol. XX, p. 119.

*Czermak, J. N.*, Ueber die Hautnerven des Frosches. Arch. f. Anat.

u. Physiol. 1849, p. 252. Leipzig, 1869.

*Czermak, J. N.*, Ueber die Herznerven des Frosches, Rana temporaria.

Arch. f. Anat. u. Physiol. 1851, p. 160.

*v. Darkschewitsch, L.*, Zur Anatomie der Glandula pinealis. Neurol.

Cent. Bl. 1886. Vol. V, p. 29.

*v. Deen*, De differentia et nexu inter nervos vitae animalis et

vitae organicae. Diss. inaugur. Lugduni-Bat., 1834.

*v. Deen*, Over de zijdelingsche Takken dar swervende zenuw van

den Proteus anguineus, aus Bijdragen tot de Naturkundige.

Wettenschappen, 1834.

*v. Deen*, Ueber den Ramus lateralis n. vagi bei den Batrachiern.

Arch. f. Anat. u. Physiol. 1834, p. 477 (abstract from Diss.

inaugur. De differentia et nexu inter nervos vitae animalis et

vitae organicae). Lugduni-Bat.

*Deiters, O.*, Untersuchungen über Gehirn und Rückenmark.

Braunschweig, 1865.

*Denissenko, G.*, Zur Frage über den Bau der Kleinhirnrinde bei

verschiedenen Klassen von Wirbelthieren. Arch. f. mikrosk. Anat.

1877. Vol. XIV, p. 203.

*Dietl, M. J.*, Beobachtungen über Theilungsvorgänge an Nervenzellen.

Wiener Sitzungsber. 1874. Vol. LXIX, Pt. III, p. 71.

*Dietl, M. J.*, Casuistische Beiträge zur Morphologie der

Nervenzellen. Wiener Sitzungsber. 1874. Vol. LXIX, Pt. III, p. 80.

*Dogiel, J.*, Die Ganglienzellen und Nerven des Herzventrikels beim

Frosche. Arch. f. mikrosk. Anat. 1877. Vol. XIV, p. 470.

*Dogiel, J.*, Die Ganglienzellen des Herzens bei verschiedenen

Thieren und beim Menschen. Arch. f. mikrosk. Anat. 1877.

Vol. XVII, p. 471.

*Dogiel, J.*, Die Nervenzellen und Nerven des Herzventrikels beim

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1851–1854. Vol. V, p. 107.

THE NERVOUS SYSTEM.

I. THE CENTRAL NERVOUS SYSTEM.

_(Re-written by the translator.)_

A. THE SPINAL CORD (Fig. 97).

*1.* *External form.* The spinal cord is, in comparison with the

brain, somewhat small; the two organs are directly continuous, and

present no distinct line of demarcation: the point of origin of

the first spinal nerve is, therefore, arbitrarily accepted as the

anterior limit of the spinal cord; posteriorly it terminates in the

_filum terminale_.

The spinal cord is flattened dorso-ventrally, and is constricted at

a point (_pars media_, Reissner) somewhat anterior to its middle: in

consequence of this constriction the cord has two enlargements; an

anterior smaller, and a posterior larger (_intumescentiae anterior v.

posterior_), from which arise the nerves of the brachial and lumbar

plexuses respectively. At about the sixth or seventh vertebra, the

hinder enlargement diminishes rather abruptly to form the so-called

_conus medullaris_; this is continued into the _filum terminate_,

which enters the cavity of the urostyle. The hinder portion of the

lumbar nerves forms a _cauda equina_, the constituent nerves of which

surround the _filum terminale_.

A *dorsal longitudinal fissure* (_sulcus longitudinalis superior_)

is well marked in the middle line of the dorsal surface of the

posterior enlargement; anteriorly and posteriorly it rapidly fades

away, its position being merely indicated by a small amount of

connective-tissue and a blood-vessel.

The *ventral longitudinal fissure* (_sulcus longitudinalis inferior_)

is well marked throughout the length of the cord. Neither fissure

can be traced in the hinder part of the _conus medullaris_ or in the

_filum terminale_.

Ten pairs of nerves arise from the spinal cord, each nerve arising

by two roots, a ventral and a dorsal (anterior and posterior), from

points near the ventral and dorsal longitudinal sinuses respectively:

the two roots unite at their point of exit from the vertebral canal

through the intervertebral foramen; each dorsal root possesses a

ganglion.

[Illustration: Fig. 97.

The nervous system of _Rana esculenta_, from the ventral surface.

(From Icones physiologicae by A. Ecker. Pl. XXIV, Fig. 1.)

_F_ Facial nerve.

_G_ Ganglion of pneumogastric nerve.

_He_ Cerebral hemisphere.

_Lc_ Optic tract.

_Lop_ Optic lobe.

_M_ Boundary between medulla oblongata and spinal cord.

_M1–10_ Spinal nerves.

_MS_ Connection between fourth spinal nerve and

sympathetic chain.

_N_ Nasal sac.

_Ni_ Sciatic nerve.

_No_ Crural nerve.

_o_ Eyeball.

_S_ Trunk of sympathetic.

_S1–10_ Sympathetic ganglia.

_Sm_ Rami communicantes of sympathetic.

_Sp_ Continuation of sympathetic into head.

_I_ Olfactory nerve.

_II_ Optic nerve.

_III_ Motor oculi nerve.

_IV_ Trochlear nerve.

_V_ Trigeminal and facial nerves.

_Va_ Ramus ophthalmicus of trigeminal.

_Vc_ Ramus maxillo-mandibularis of trigeminal.

_Vd_ Mandibular branch of trigeminal.

_Ve_ Hyomandibular branch of facial.

_Vg_ Gasserian ganglion.

_Vs_ Upper end of sympathetic trunk in connection with

Gasserian ganglion.

_VI_ Abducens nerve.

_VII_ Facial nerve.

_VIII_ Auditory nerve.

_X_ Glossopharyngeal and pneumogastric nerves.

_X1_ Ramus anterior of glossopharyngeal.

_X2_ Ramus Posterior of glossopharyngeal.

_X3–4_ Branches of pneumogastric.

]

*2.* *Internal structure.* As in other vertebrates, the spinal cord

consists of white and grey matter, the latter being surrounded by

the former; the relative amount of the one to the other varies in

different parts of the cord.

A *transverse* section of the cord presents the same general

characteristics as a similar section from a bird or mammal. The grey

matter lies in the middle, surrounding the central canal, and is

prolonged into each half of the section by ventral and dorsal horns

or _cornua_. The grey matter is surrounded by the white, but is not

so sharply marked off from this in amphibia as in birds and mammals.

A section through the *anterior enlargement* is almost quadrangular

in outline, and somewhat broader below than above; the ventral

longitudinal fissure is well marked and deep, while the dorsal is

indistinct. The dorsal horns are narrow and short, the ventral longer

and much broader. The space between the dorsal horns is narrow and

deep, that between the ventral shallow; the outer boundary of the

grey matter is almost straight, and the central canal is placed above

the centre of the section.

In the *constricted portion* of the cord (_pars media_) the relative

proportion of the white matter is increased at the expense of the

grey; the horns are less distinct, the dorsal being directed outwards

and the ventral somewhat flattened. The central canal is in the

centre of the section.

The *posterior enlargement* shows the ventral horns projecting

outwards, very much enlarged, and approaching the periphery; the

space between them is semilunar. The dorsal horns are well developed,

especially at their upper parts; the space between them is narrow and

deep. The outer margin of the grey matter is again nearly a straight

line. The central canal is below the centre of the section.

The arrangement of the parts in the _conus medullaris_ approaches

that in the _pars media_: the horns are much diminished in size, and

posteriorly they entirely disappear, the ventral horns persisting

longer than the dorsal. The central canal approaches the lower

surface.

No cornua can be recognised in the _filum terminale_ behind the

origin of the last pair of spinal nerves. The grey matter has here

a circular outline, with the exception of a very slight indentation

below; the white matter is almost absent: the central canal is on the

lower border of the section in the anterior part, while posteriorly

it occupies almost the entire space below the _pia mater_.

The *Central Canal* lies in the median line, and always presents a

distinct lumen; in the two enlargements this lumen forms a vertical

ellipse[51], while in the _pars media_ it is circular. The canal is

lined by a single layer of ciliated columnar epithelium, the cells

of which are usually conical, with their bases directed towards the

lumen; but other forms may be noticed. These cells average about

0·040 mm. in length and about 0·002 mm. in width at their broadest

part. The peripheral extremities of the cells have processes, which,

in the case of the cells above and below the cord, may often be

traced as far as the _pia mater_; the processes of the lateral cells

are less distinct and seem to be shorter. Anteriorly the central

canal opens into the floor of the fourth ventricle. The termination

of the central canal, posteriorly, has been described by Masius and

Vanlair. According to these observers, the canal towards the hinder

end of the _filum terminale_ increases very gradually in width, until

its epithelial cells are immediately underneath the _pia mater_,

its walls then converge somewhat more sharply to close the cavity.

From the point where the canal commences to increase in width, a

change takes place in its epithelial wall, which, instead of having a

single layer of cells, now possesses two to three layers; towards the

posterior end it is again thinned out to a single layer.

[Footnote 51: v. Kölliker states that the lumen is everywhere

circular.]

The *grey matter* varies in different parts of the cord and

in different parts of the same section. In a section through

the anterior enlargement a portion of the grey matter, placed

immediately above the central canal, is easily distinguished by its

vertically-placed, elliptical outline, and by its transparency: it

is known as the _substantia reticularis_. It consists of a mesh-work

of branched cells, through which course distinctly contoured fine

commisural fibres which are derived chiefly from the dorsal cornua,

the vertical from the epithelial cells of the roof of the central

canal. In a series of sections the _substantia reticulosa_ may be

seen to originate indistinctly in the most anterior part of the cord;

in the anterior enlargement it already possesses the elliptical form

mentioned, and in the _pars media_ it commences to send off lateral

processes to either side. In the posterior enlargement it reaches

its greatest absolute development, as it here surrounds the central

canal; while further backwards it encroaches until, together with the

remaining grey matter, it constitutes almost the whole of the _filum

terminale_.

Immediately underneath the central canal is a narrow band, which

immediately connects the grey matter of the two sides; above it is

bounded by the epithelium of the central canal, below by medullated

fibres. This _septum medium_ (Reissner) may be traced backwards from

the anterior extremity of the cord to the posterior enlargement. It

consists chiefly of transverse fibres, together with vertical fibres

derived from the epithelial cells on the floor of the canal, and

like the _substantia reticulosa_, is distinguished by its greater

transparency from the rest of the grey matter.

The grey matter consists of connective-tissue and nervous elements,

which are very intimately united; each of these elements including

its special cells and fibres. The *nerve-cells* vary much in size and

appearance. A prominent group of large cells in the ventral cornua,

the *lateral group* (Stieda), is very conspicuous. The individual

cells are seldom rounded, but are usually spindle-shaped or angular,

and each possesses one to five processes, which may often be traced

through considerable distances. These cells have an average length of

0·040 mm., and are about 0·016 mm. broad; they are somewhat larger

in the anterior enlargement than in the posterior, and are also

more numerous in a section from the anterior enlargement than in a

section from the posterior. In the terminal filament they gradually

disappear. Some of their processes are continued into the lateral

columns (Köppen).

Smaller, spindle-shaped or triangular cells are scattered irregularly

throughout the grey matter, without forming distinct groups, though

for descriptive purposes those of the centre have been named the

*central group*. In a stained, transverse section of the cord are

seen numerous *nuclei*; these undoubtedly belong both to small

nerve-cells and to connective-tissue cells, but except under the

most favourable conditions the two are indistinguishable. The larger

nerve-cells present the usual characters of nerve-cells: more or

less granular contents, nucleus, well-marked nucleolus, more or less

marked fibrillation; their processes are usually more homogeneous.

The cells are frequently pigmented.

The *processes* of the cells belonging to the lateral group radiate

in all possible directions, but certain well-marked processes

directed towards the middle line can always be made out. From

the lowest part of this group a few processes may be traced into

the ventral roots. The small cells seem to be chiefly arranged

vertically, although their processes radiate in all directions

(Stieda).

The *Fibres of the grey substance* are nearly all non-medullated.

They may be traced in all directions, but the best-marked groups are

either vertical or transverse. The vertical fibres appear to arise

from the central grey matter, and to ascend in larger or smaller

bundles towards the periphery. The transverse fibres are arranged

chiefly in the two commissures: the dorsal commissure (_commissura

superior_) is the smaller; the fibres are parallel, and show no trace

of decussation; externally they radiate in various directions to

the dorsal horns. The ventral commissure is composed of two layers,

the upper grey (_commissura inferior grisea_), the lower white

(_commissura inferior alba_): both are interrupted by the _septum

medium_. A well-marked decussation of the fibres is seen in the

middle line; the exact mode of termination of these fibres has not

been made out, but many appear to communicate with the large cells of

the ventral horns.

The *white matter* consists chiefly of longitudinal, medullated

fibres, in which various columns may be distinguished. The dorsal

columns are separated from the lateral by a process of neuroglia,

continued from the general investment of neuroglia lying under the

_pia mater_. The line of separation between the lateral and ventral

columns is not well marked; it is about the line which would be

formed by prolonging the ventral horns to the surface.

The fibres vary in size, and fibres of all sizes may be found in any

particular part of a transverse section; still the fibres of the

ventral columns have an average greater diameter than those of the

lateral columns, and the fibres of the dorsal columns are finer than

those of the lateral. The largest fibres of the ventral column are

placed near the ventral fissure and on its lower border; they attain

their greatest development in the posterior enlargement; these fibres

frequently contain two or three axis-cylinders each. Köppen suggests

that they may represent the _formatio reticularis_ of higher animals.

In the lateral columns the larger fibres are placed close to the

grey matter, these columns also receive fibres from the cells of the

lateral group.

In the dorsal columns the radiating root-fibres never reach the

median plane, but leave an area of purely longitudinal fibres on each

side of the dorsal fissure; these represent Goll’s columns, and have

a club-shaped outline in transverse section.

The white matter is pierced in all directions by fine

connective-tissue fibres and bundles of fibres which radiate from

the grey matter; some branch and join with others to form a network,

others pass almost uninterruptedly to the periphery. From the

connective-tissue cells of the _pia mater_, processes pass into the

white matter and assist in completing the connective-tissue matrix

for the nervous elements.

*Dorsal roots* of the spinal nerves. Each root consists of a single

bundle of nerve fibres, which suddenly bends at the periphery of the

cord in order to descend vertically through the white matter towards

the dorsal horn; the fibres divide into three sets, one to the dorsal

commissure, a second to the upper horn itself, and the third helps to

form the dorsal columns.

*Ventral roots* of the spinal nerves. Each ventral root consists of

three or four delicate threads, which may be traced to the ventral

cornua, which they reach after a vertical or very slightly oblique

course through the white matter. Other fibres arise from the ventral

columns, but these oblique fibres never extend to the median plane.

*Pigment* is found distributed irregularly through a section from any

part of the spinal cord; it is increased in amount in those parts in

which there is an increased amount of grey matter. The pigmentation

is always found more marked in the lower parts of the ventral horns

than in other parts.

B. THE BRAIN (Figs. 98, 102, 103, and 105).

*1.* *General description.* From behind forwards, the *dorsal*

surface of the brain presents the following parts for examination:

the _medulla oblongata_, the _cerebellum_, the optic lobes, the

thalamencephalon, the cerebral hemispheres, and the olfactory lobes.

The _medulla oblongata_ is a direct continuation of the spinal cord;

it is wider anteriorly than posteriorly, and is separated in front

from the optic lobes by a vertical plate of nervous matter, the

_cerebellum_. The optic lobes are two symmetrical ovoid bodies touching

each other in the median plane, and together forming the widest part

of the brain. In front of the optic lobes is the thalamencephalon,

with a thick vascular membrane, the choroid plexus, lying on its upper

surface, and connected with the pineal gland; the thalamencephalon

extends forwards between the posterior ends of the cerebral

hemispheres. The cerebral hemispheres are two symmetrical ovoid bodies,

narrow in front, wider and slightly diverging behind: each hemisphere

is prolonged forwards to form an olfactory lobe.

The *ventral* surface has in front the olfactory lobes, then the

cerebral hemispheres, behind these the _lamina terminalis_, the _tuber

cinereum_, the optic chiasma, the pituitary body, the _crura cerebri_,

and lastly the _medulla oblongata_, in the order here given.

The various *cranial nerves* (Figs. 102 and 103) may be seen to arise

as follows:--The olfactory nerve (_I_, _I_′, _L.ol_′) arises directly

from the anterior end and outer side of the corresponding olfactory

lobe, and from the cerebral hemisphere. The optic nerve (_To_ and

_II_) arises, as the optic tract, from the side of the brain below the

optic lobe, whence it passes to the chiasma on the under surface of

the brain. The oculo-motor (_III_) takes its origin from the ventral

surface close to the median line and between the _crura cerebri_.

The pathetic or trochlear nerve (_IV_) is attached to the dorsal

surface between the optic lobes and the _cerebellum_. The trigeminal

nerve (_V_) arises from the side and anterior part of the _medulla

oblongata_. The _abducens_ (_VI_) arises behind the pituitary body

close to the median line from the ventral surface of the medulla

halfway between the _sulcus_ and the origin of the _vagus_. The

facial and auditory (_VII_ and _VIII_) nerves arise from the _medulla

oblongata_ behind the trigeminal nerve, the facial being in front of

the auditory nerve. The glossopharyngeal (_IX_) nerve arises, in common

with the pneumogastric nerve (_X_), behind the auditory nerve.

*2.* *The several parts of the brain.*

*a.* The _medulla oblongata_ (Figs. 98, 102, and 103 _Mo_)--α.

*External form.* The _medulla_ is limited behind by the origin of the

first pair of spinal nerves, at which point a very faint constriction

is sometimes found: it extends forwards as far as the _cerebellum_. It

gradually widens as it passes forwards until just before it reaches its

anterior limit, where it presents a shallow but sharp constriction. The

*dorsal surface* is characterised by the presence of a deep, triangular

fossa, the *fourth ventricle* (Fig. 98 _S.r_), (_ventriculus quartus_,

Stieda; _sinus rhomboideus s. sinus triangularis_, Reissner; _fossa

rhomboidalis_); the sides of the triangle are, however, not quite

straight, but are slightly bent outwards just before they converge

towards the posteriorly-directed apex; the base of the triangle is

formed by the _cerebellum_. By careful examination, the ventricle is

seen to be continued for a short distance under the _cerebellum_,

where it opens into the Sylvian aqueduct. In the floor of the fourth

ventricle is a well-marked median longitudinal fissure (_sulcus

centralis_), (Fig. 98 _S_). Into the posterior part of the ventricle

opens the central canal of the spinal cord. As the fourth ventricle is

formed by the white matter passing to either side, and the simultaneous

flattening of the grey matter, the floor of the fourth ventricle is

composed of grey matter.

The fourth ventricle is closed in by a highly vascular membrane,

the *choroid plexus* of the fourth ventricle (_plexus choroideus

ventriculi quarti_, Reissner; _velum medullare posterius_). The

blood-vessels of the plexus will be described together with the

other vessels of the brain (p. 162). They are supported by a

connective-tissue matrix, and the whole covered with flattened

epithelium, which in the fourth ventricle is ciliated and often

pigmented.

The *ventral surface* of the _medulla oblongata_ (Fig. 102 _Mo_) has

a median ventral longitudinal fissure, a direct continuation of that

of the cord; in the anterior part of the _medulla oblongata_ there is

also to either side of this a lateral fissure, continued on to the

_crura cerebri_; these fissures correspond to the positions of the two

_rami posteriores_ of the internal carotid arteries; they are always

well seen in microscopical sections. The _medulla oblongata_ is so

intimately connected with the _pars commissuralis_ (pp. 149, 150) that

the minute anatomy of the two is best described at the same time.

β. *Internal structure.* Examined by means of serial sections, the

_medulla oblongata_ is seen to have, in comparison with the cord, an

increased amount of grey matter; this is especially the case in its

anterior part. The floor and inner parts of the walls of the ventricle

are formed of grey matter, in which the largest-sized cells have

disappeared, to be replaced by medium-sized cells. Traced from behind,

the ventral horns of the cord are seen to increase in size and to be

more widely separated until they form two isolated masses, while the

dorsal horns gradually diminish; at the same time they are forced

outwards and upwards, until they lie under the floor of the ventricle,

and so extend to the _pars peduncularis_.

[Illustration: Fig. 98.

Dorsal view of brain of _Rana esculenta_.

_Ad_ Choroid plexus.

_C_ Cerebellum.

_f_ Groove between cerebral hemispheres and olfactory lobes.

_G_ Opening in the roof of the third ventricle.

_Gp_ Pineal body.

_Hc_ Cerebral hemispheres.

_I_ Olfactory nerve.

_Li_ Wall of fourth ventricle.

_L.ol_ Olfactory lobe.

_L.op_ Optic lobe.

_Mo_ Medulla oblongata.

_S_ Longitudinal fissure of the fourth ventricle.

_S.r_ Fourth ventricle.

_Tho_ Thalamencephalon.

]

*Grey matter.* The _substantia reticularis_ is not present, but the

_septum medium_ extends forwards as far as the _pars peduncularis_.

The central canal extends upwards at the expense of the tissue above

it, and is here pear-shaped; at the same time the dorsal longitudinal

fissure deepens until the two meet in the fourth ventricle; beyond this

point one cannot speak of dorsal and ventral horns.

[Illustration: Fig. 99.

Transverse section through hinder end of _Medulla oblongata_ (magnified

30–80)[52], from Stieda.

[Footnote 52: In these diagrams, from Ludwig Stieda’s Studien über das

centrale Nervensystem der Wirbelthiere, the outline of the diagram is

magnified thirty times, while the details are magnified eighty times.]

_b_ Inferior commissure.

_f_ Dorsal horns.

_g_ Ventral horns.

_h_ Fourth ventricle.

_i_ Nucleus centralis.

_k_ Isolated mass of grey matter in which longitudinal

fibres of the pneumogastric nerve course.

]

Small nerve cells are irregularly distributed throughout the whole of

the grey matter and cannot be grouped; the larger cells, on the other

hand, are arranged in distinct groups which have special relations with

the nerves arising from the part. Occasionally these groups are not so

isolated as usual; in this case processes of one group can be traced

into another group (Reissner). Of these groups the chief are:

1. The _nucleus centralis_ (upper inner group, Reissner), _nucleus

medullae oblongatae_, Stieda (Fig. 99 _i_), is a group of cells found

towards the hinder end of the _medulla oblongata_, on either side of

and below the central canal; the group can be traced under the floor

of the fourth ventricle to about its middle. The cells are rounded or

spindle-shaped, the processes directed upwards, downwards, or outwards;

their average size is 0·040–0·048 mm. long by 0·020 mm. broad.

2. The *auditory nucleus* (_nucleus acusticus_, Reissner, Stieda),

(Fig. 100 _n_) is a large group of cells found in the wall of the

fourth ventricle opposite the point of origin of the auditory

nerve. The cells are rounded, pear-shaped, or of spindle form, and

interspersed between the nerve fibres; these cells have an average

length of 0·040 mm., and are about half as broad. The fibres of the

auditory nerve radiate from their superficial origin in all directions

through the grey matter towards these cells, and evidently communicate

with them (Fig. 101 _p_). One small group (Fig. 101 _r_) passes to

a lower level than the rest, and is regarded by Stieda as the true

auditory centre. Köppen considers that the auditory nerve has a

threefold origin: (1) from small cells on the median surface of the

auditory area; (2) from the large cells between the above; (3) from a

group of free nuclei on the dorsal surface of the auditory area.

[Illustration: Fig. 100.

Transverse section through the _Medulla oblongata_ at the point of

origin of the abducens nerve, from Stieda. (Magnified 30 x 80.)

_h_ Fourth ventricle.

_m_ Abducens nerve.

_n_ Auditory nucleus.

_o_ Abducens nucleus.

]

3. The *trigeminal nucleus* (_nucleus trigeminus_), (Fig. 101 _q_) lies

in part beneath the auditory nucleus but extends further forwards. It

forms a rounded group of cells placed under the outer angle of the

grey matter. The cells are somewhat crowded together, and are chiefly

of an elongated spindle-form, with their processes directed obliquely

downwards and outwards. The fibres of the trigeminal nerve separate

into two groups; the upper group is best traced in a horizontal

section, the fibres curving round to join the longitudinal fibres

continued from the dorsal columns of the cord. The fibres of the lower,

smaller group pass transversely inwards to the trigeminal nucleus.

According to Reissner the latter fibres are motor, the former sensory.

Probably other nerves are connected with the hinder part of this group.

4. The *abducens nucleus* (Fig. 100 _o_). From its superficial origin,

the fibres of the abducens nerve may be traced vertically upwards to a

small, rounded, grey mass; at this point the mass is somewhat isolated,

but further forwards it may be traced as belonging to the central grey

matter; it contains small spindle cells.

[Illustration: Fig. 101.

Transverse section of the _Medulla oblongata_, at the point of origin

of the auditory nerve, from Stieda. (Magnified 30 x 80.)

_h_ Fourth ventricle.

_n_ Auditory nucleus.

_o_ Abducens nucleus.

_p_ Auditory nerve.

_p_′ Ganglion of auditory nerve.

_q_ Hinder portion of trigeminal nerve.

_r_ Bundle of fibres arising from trigeminal nucleus

and joining the auditory nerve.

]

5. The *pneumogastric nucleus*. The pneumogastric, with its numerous

irregular roots, arises from the side of the _medulla oblongata_. The

hindermost fibres can be traced as a small bundle, passing almost

transversely through the white matter to the outer margin of the grey

matter. The larger portion of the fibres is placed in front of these;

part of this seems to be directly continuous with the longitudinal

fibres of the white matter; a second part, however, can be traced from

the surface transversely through the white matter to the grey matter.

These latter fibres, together with those of the group first described,

do not arise from the grey matter in this part of the _medulla

oblongata_, but curve round and run backwards longitudinally through

the grey matter, thus forming a rounded bundle of fibres (Fig. 99 _k_).

Between these fibres are interspersed small nerve-cells and nuclei

which disappear as the fibres approach the white matter. The vagus

undoubtedly receives fibres from the grey matter throughout a long

course, and again receives a large bundle just before leaving the grey

matter. The more exact origin of the various fibres has not been traced.

6. The _nucleus magnus_ (Reissner and Stieda) is a very peculiar group

of cells placed on either side, in the most anterior portion of the

_pars commissuralis_, that is, immediately underneath the _valvula

cerebelli_. The large cells are arranged in a transverse section in a

single row so as to enclose a pear-shaped space on either side, which

has its long axis directed from above, downwards and outwards, the

narrower end being above. In longitudinal section the line of cells is

seen to be open in front. The space enclosed by these cells is occupied

by a granular ground-substance which contains only few nuclei. Bellonci

is of opinion that these nuclei represent the _corpora quadrigemina

posteriora_ of higher animals.

*White matter.* In the hinder part of the _medulla oblongata_ the

arrangement of the white matter resembles that of the white matter

of the spinal cord; further forwards the white matter of the dorsal

surface commences to pass to either side, and ultimately it forms the

outer part of the walls of the fourth ventricle. The fibres of the

white matter of the ventral surface are unchanged in direction as they

proceed forwards. The fibres of the anterior part of the medulla are

thinner than those of the posterior portion (Stieda), according to

Reissner they gradually thin as they pass forwards. The fibres are

nearly all longitudinal, such transverse and oblique fibres as are

present being chiefly in connection with the various nerve-roots and

the commissures.

The _commissura superior_ is naturally lost in consequence of the

opening of the central canal into the fourth ventricle; the _commissura

inferior_ is increased in the anterior half and decreased in the

posterior half of the _medulla oblongata_; in the latter the fibres

become more and more oblique, and decussate very freely; ultimately

they seem to be either continued as longitudinal fibres or to join the

ganglia.

Near the _pars commissuralis_ is a transverse arched band of fibres,

passing from the under surface of one half of the cord over the ventral

longitudinal fissure through the _septum medium_ to the under surface

of the opposite half; part of the fibres are continued upwards along

the periphery to the _cerebellum_, part to the _nucleus magnus_.

Vertical, straight, or slightly arched fibres are found in the walls of

the fourth ventricle.

A section from the _medulla oblongata_ has a larger amount of *pigment*

than a section from the spinal cord, and the anterior portion of the

_medulla oblongata_ contains more than the posterior portion. The

pigment is chiefly found in a curved line, placed in the lower and

outer parts of the grey matter; the amount present varies in different

specimens.

*b.* The _Cerebellum_ and _Valvula cerebelli_.

α. *External form.* The _cerebellum_ is a thin, semilunar plate,

which projects between the optic lobes and the fourth ventricle, its

base covering the most anterior part of the ventricle; the posterior

surface possesses a very faint median fissure (Reissner). The _valvula

cerebelli_ (_Velum medullare anterius_, Reissner) is the thin lamella

which connects the anterior surface of the _cerebellum_ with the optic

lobes.

β. *Minute structure.* By means of longitudinal, vertical sections,

the posterior surface of the _cerebellum_ is seen to be covered with

epithelium; in the lower part of the surface this is columnar or

conical, above it is flattened: immediately beneath, that is in front

of this is a finely granular layer, with very closely packed and

granular nuclei. In front of these is a stratum of nerve-fibres forming

the second layer of the cerebellum.

Still more anteriorly is the third layer of the cerebellum, an

irregular double layer of large cells (Purkinje’s cells, Denissenko);

the cells have an average length and breadth of 0.040 mm. and 0.015 mm

respectively; they are pear-shaped or of spindle-form, and possess

usually two well-marked processes, one passing into the layer behind,

the other forwards into the anterior layer to be immediately described,

while other less distinct processes radiate irregularly in all

directions. The fourth and most anterior layer of the cerebellum is a

thick stratum of nerve fibres with numerous nuclei (0.006 to 0.008 mm.

diameter). The fibres are for the most part arranged transversely, but

some course in various directions. These fibres underlie the flattened

epithelium which covers the anterior surface of the cerebellum.

The fibres of the second layer course, for the most part, in an almost

vertical plane; they connect the _cerebellum_ with the optic lobes

(_processus cerebelli ad corpora bigemina_) and with other parts of the

brain.

The fibres of the fourth layer receive numerous long processes from the

large cells of Purkinje; they form a large commissural system, which

can be followed ventrally on each side into the _pars commissuralis_.

A part of the fibres ends here in the grey matter, a second portion

enters the auditory area and forms a descending auditory root, a third

part joins the lateral columns (in the _medulla oblongata_), and

more anteriorly some join the ventral columns. The descending fibres

from the _cerebellum_, together with the _fibrae arcuatae_ found in

the ventral columns, indicate the presence of a _pons Varolii_. The

fibres of this ventral commissure decussate only on its dorsal surface

(Köppen).

The _Valvula cerebelli_ contains a few medullated fibres and the roots

of the trochlear nerves; these pass from the _medulla oblongata_ into

the _valvula cerebelli_, cross in the median line, and then proceed

forwards as the trochlear nerves.

The *pigment* in the _pars commissuralis_ is arranged in a curved line

similar to that found in the _medulla oblongata_, but the line is

shortened at either extremity, and consequently does not extend into

the _cerebellum_.

*c.* The *optic lobes* and _Crura cerebri_. (_Corpora geminata_ and

_Pars peduncularis_, Reissner; _Lobus opticus_, Stieda; Vierhügel,

Tiedemann; Vierhügel (Zweihügel) and _Pedunculi cerebri_, Schiess;

Mesencephalon, Huxley.)

[Illustration: Fig. 102.

Ventral view of brain of _Rana esculenta_.

_Cho_ Optic chiasma.

_Hc_ Cerebral hemispheres.

_Hy_ Pituitary body.

_L.ol_ Olfactory lobe.

_L.ol^1_ Origin of olfactory nerve from

the cerebral hemisphere.

_Lt_ Lamina terminalis.

_Mo_ Medulla oblongata.

_To_ Optic tract.

_Tu.c_ Tuber cinereum.

_I_ 1st }

_I^1_ 2nd } root of the olfactory nerve.

_II_ Optic nerve.

_III_ Oculo-motor nerve.

_IV_ Trochlear nerve.

_VI_ Abducens nerve.

_V_, _VII_, _VIII_ Trigeminal, facial, and

auditory nerves.

_IX_, _X_, _XI_ Glossopharyngeal, pneumogastric,

and accessory nerves.

]

α. *External form* (Figs. 98, 102, 103 _L.op_). The optic lobes are two

prominent ovoid bodies placed immediately in front of the _cerebellum_,

and connected with it by the _valvula cerebelli_; posteriorly they

touch each other in the median plane, while anteriorly they diverge

and thus constitute the widest part of the brain; in the angle thus

formed is the thalamencephalon. The optic lobes are always more darkly

pigmented than any other part of the central nervous system.

[Illustration: Fig. 103.

Lateral view of brain of _Rana esculenta_.

_Ad_ Choroid plexus.

_C_ Cerebellum.

_Gp_ Pineal body.

_Hc_ Cerebral hemisphere.

_Hy_ Pituitary body.

_L.ol_ Olfactory lobe.

_L.ol_^1 Disc at origin of second root.

_L.op_ Optic lobe.

_Mo_ Medulla oblongata.

_Tho_ Thalamencephalon.

_To_ Optic tract.

_Tu.c_ Tuber cinereum.

_I_ 1st }

_I^1_ 2nd } root of olfactory nerve.

_II_ Optic nerve.

_IV_ Trochlear nerve.

_V_ Trigeminal nerve.

_VII_ Facial nerve.

_VIII_ Auditory nerve.

_IX_, _X_, _XI_ Glossopharyngeal, pneumogastric,

and accessory nerves.

]

The _Crura cerebri_ are two columns of white matter, placed beneath

the optic lobes, and partly hidden by the pituitary gland. At their

junction with the _medulla oblongata_, or rather with the _pars

commissuralis_, is a very slight transverse fissure; at the same point

the ventral longitudinal fissure is interrupted by an extremely small

grey tubercle (Stieda).

β. *Internal structure.* From the anterior extremity of the fourth

ventricle a canal, the Sylvian aqueduct (_Aqueductus Sylvii_, _iter

a tertio ad quartum ventriculum_), may be traced forwards under the

cerebellum, in the median line of this section of the brain. At about

opposite the middle of the length of the optic lobes the canal is

dilated and communicates with the cavities or ventricles (_Ventriculi

lobi optici_, Stieda) enclosed by these; a general cavity is formed,

which in transverse section has something of the form of the letter T.

The roof of the cavity is thinner than the floor; this is especially

the case in the median plane opposite the superior longitudinal fissure

between the optic lobes; the floor is thinned in the middle line by the

descending portion of the cavity. The cavity of each optic lobe extends

both forwards and backwards beyond its point of communication with the

dilated Sylvian aqueduct, hence in a transverse section taken in front

of this point (Fig. 104 _h_′) the cavity of either side appears to be

isolated; in a horizontal and longitudinal section (Fig. 105 _Aq_) the

general arrangement of the parts may be well seen.

The grey matter is chiefly arranged in a layer so as to surround the

cavity (Fig. 104), this layer being deeper on either side of the

descending portion of the cavity than elsewhere. For the rest the grey

matter is much interspersed among the white matter, except at the

circumference of the section, which is entirely formed of white matter.

It contains a large number of small cells, of which the nuclei are

alone visible; in the parts mentioned where the layer is most marked

these cells are arranged in oblique rows, between which pass fine

bundles of medullated fibres (Fig. 104, between _u_ and _h_′). A group

of large cells (Fig. 104 _u_) is found on either side of the middle

line and under the floor of the cavity; the cells are about 0.032 mm.

long, and 0.016 mm. broad; the oculo-motor nerve may be traced to

this group, which is the *oculo-motor nucleus*. A small commissure of

decussating fibres connects the nuclei of opposite sides (Köppen).

[Illustration: Fig. 104.

Transverse section through the anterior portion of the optic lobes

opposite the origin of the motor-oculi nerve; from Stieda (magnified

30–80).

_h_ Lower portion of cavity.

_h_′ Lateral portion of cavity.

_u_ Ganglia of oculi-motor nerve.

_v_ Oculo-motor nerve.

_w_ Large cells of the optic lobe.

_x_ Roof of optic lobe.

_y_ Posterior commissure.

_z_ Anterior diverticulum of the cavity.

]

The substance of the roof of the optic lobes (Fig. 104 _x_) is arranged

in very distinct layers: above is a layer free from cells; the fibres

of this layer are extremely fine and delicate, and have not been

accurately traced; under this is a nuclear layer; a second layer of

fine fibres follows, which is succeeded by a second layer of nuclei in

a granular matrix, and lastly epithelium (Stieda). (Reissner describes

three nuclear layers in _Bufo variabilis_, and this is also the case in

_R. temporaria_, G. H.; according to Köppen, the number is variable.)

In the anterior portion of the roof a distinct bundle of fibres may be

made out; externally they bend downwards, and can be traced as far as

the _crura cerebri_.

Fibres corresponding with the commissural and arched fibres of the

_medulla oblongata_ are continued into the hinder portion of the _crura

cerebri_, the change from _pars commissuralis_ to _crura cerebri_ being

very gradual. The longitudinal white fibres are much increased in

number in the _crura cerebri_, and a portion of them can be traced to

the _nucleus magnus_.

A peculiar irregular group of large cells (Fig. 104 _w_) is found where

the roof meets the _crura cerebri_; these vary in diameter from 0.024

to O.040 mm., and their processes are very indistinct.

The fibres of the optic tracts arise, according to Köppen, from two

different origins: the one lies on the hindermost part of the optic

lobe; from this point the fibres curve downwards and forwards to form

longitudinal fibres; this root Köppen names the ventral ascending root,

it can be traced through the entire length of the organ. The second

root arises in the _tectum opticum_ near the longitudinal fissure; it

is smaller than the foregoing, and has been named the dorsal ascending

optic root. The fibres of these two roots unite anteriorly near the

posterior commissure, at which point they receive additional fibres

(Köppen). Bellonci traces a large proportion of the fibres of the optic

tract to the _nucleus magnus_, which pair of nuclei, as already stated,

he regards as the posterior pair of _corpora quadrigemina_ of higher

animals.

The _pars peduncularis_ is the continuation of the _pars commissuralis_

underneath the optic lobes; a gelatinous mass lying in the median plane

and containing numerous isolated nuclei (_Ganglion interpedunculare_)

divides it into two lateral halves. The longitudinal fibres are

ungrouped posteriorly, but arranged in rounded strands in the middle,

especially dorsally; anteriorly the grouped arrangement is lost and the

number of fibres diminished.

The *pigment* of this region has, in a transverse section, an outline

which has something the form of a lyre; commencing on either side of

the median line, and underneath the deepest portion of the cavity,

the pigment line passes, first, directly outwards; then suddenly turns

upwards and slightly outwards parallel with the wall of the deeper part

of the cavity; it then curves outwards to pass below the cavity of the

optic lobe, where it divides, one portion passing outwards, the other

between the Sylvian aqueduct and the ventricle of the optic lobe.

*d.* The *Thalamencephalon* (Huxley), (_Lobus ventriculi tertii_,

Stieda; _Thalami optici_, Reissner; _Thalamus opticus s. Lobus

ventriculi tertii_, Stannius; Ganglien der Haemisphaeren, Carus).

α. *External form.* From above (Fig. 98 _Tho_) the thalamencephalon

is seen as a lozenge-shaped mass lying in front of the optic lobes,

and behind and between the diverging posterior ends of the cerebral

hemispheres; it is covered by a thick vascular membrane, the *choroid

plexus*, through which passes the pedicle of the *pineal body*

(_Glandula pinealis_). On removing the choroid plexus a small aperture

is seen in the roof of the thalamencephalon, connecting the hollow

pedicle of the pineal gland with the *third ventricle*. The ventricle

appears as a narrow slit in the median line, its walls being formed

by the optic thalami. By pressing aside the cerebral hemispheres the

*posterior commissure* (_Commissura posterior_) may be seen lying quite

in front and deep in the cleft of the ventricle. Immediately behind the

pedicle of the pineal body is a slight but well-marked depression (Fig

98 _G_), the origin of which has not been investigated (Wiedersheim).

The choroid plexus is continued forwards between the cerebral

hemispheres (Figs. 98, 103 _Ad_) for some distance, and terminates in a

fine thread of connective-tissue.

The under surface of the thalamencephalon (Fig. 102) is divided

into two parts by the *optic chiasma* (_Cho_): the anterior portion

(_Lt_) is the _lamina terminalis_ (_Substantia cinerea anterior_);

the posterior (_Tu.c_) the _tuber cinereum_. The _lamina terminalis_

is bounded on either side by the cerebral hemispheres. The _tuber

cinereum_ (Figs. 102, 103 _Tu.c_) is a small median swelling

immediately behind the optic chiasma, and caused by the depression

of the floor of the third ventricle to form the _infundibulum_

(_Diverticulum infundibuli_, Reissner).

[Illustration: Fig. 105.

Horizontal section through the brain to show the ventricles.

_Aq_ Ventricles of optic lobes and the aqueduct of Sylvius.

_Dv_ Third ventricle.

_MF_ Foramen of Monroe.

_Sv_ Lateral ventricle.

_Vv_ Fourth ventricle.

]

The *pituitary body* (_Hypophysis cerebri_) is a flattened sac, placed

behind the _tuber cinereum_ and continuous with it by means of the

_infundibulum_.

β. *Internal structure.* The aqueduct of Sylvius, after communicating

with the ventricles of the optic lobes, again contracts (Fig. 105),

but still remains somewhat larger than before. In the thalamencephalon

the Sylvian aqueduct opens into the third ventricle, which gradually

assumes the form of a vertical slit with the walls bulging slightly

outwards in their upper parts. The thin roof of this ventricle, where

complete, contains a band of transverse fibres. The floor is depressed

both before and behind the part immediately above the optic chiasma,

the posterior depression lying above the _tuber cinereum_, which here

descends towards the _infundibulum_: a transverse section through this

portion of the third ventricle has the form of a square standing on one

angle, the superior angle being produced into the vertical slit of the

general cavity. The anterior depression is formed by the general cavity

being prolonged downwards and forwards to the _lamina terminalis_ in

the form of a narrow and shallow slit.

The white and grey matter of the thalamencephalon are only so far

distinct in that the portion immediately surrounding the cavity is

darker than the rest of the section. In the immediate neighbourhood

of the cavity are many small cells and nuclei, which become scarcer

further from the ventricle; they are arranged in rows, separated by a

fibrillated matrix. On either side is a distinct bundle of longitudinal

fibres, the ‘round bundle’ of Köppen, which come from the optic lobes

but appear to arise from either the _pars commissuralis_ or the

_medulla oblongata_, and to receive additional fibres from the optic

thalami; they pass forwards to the base of the cerebral hemispheres

(Stieda). A second set of longitudinal fibres arises in the substance

of the _tuber cinereum_ and passes forwards to the hemispheres (strand

of the _Tuber-cinereum_ and _Thalamus_, Köppen); this band, together

with the ‘round bundle,’ form a _crus cerebri_ (Köppen).

The commissural fibres of the thalamencephalon are: (1) a _commissura

transversa Halleri_ in the posterior portion of the organ; (2) an optic

commissure, consisting of fibres arising from the thalamencephalon

(thalamencephalic root) and crossing the median line to join the optic

tract of the opposite side; (3) a probable commissure between the optic

nerves just in front of the _chiasma opticorum_; the existence of the

latter is not yet proved beyond doubt; (4) the large commissure of

transverse fibres found in the roof of the third ventricle; whether the

fibres decussate or not is uncertain (Köppen).

The fibres of the large commissure of the roof are, in part, continued

into the strand of the _Tuber-cinereum_ above mentioned, and thus

conducted to the posterior parts of the hemispheres (Köppen). A group

of fibres (Meynert’s band, Köppen) is found in each lateral wall of

the third ventricle; they pass from the region of the _nucleus parvus_

downwards in a curved course almost parallel with the external border

of the thalamencephalon.

A distinct group of cells may be noted in this section of the brain,

an arciform or circular group (_Nucleus parvus_, Reissner; ganglion

of the _habenula_, Köppen) of large spindle cells (average diameter

0.016 mm.), placed under the upper border close to the third ventricle;

the group extends alongside the whole length of the ventricle. A second

group lying in the middle and posterior parts beneath the ventricle,

Köppen names the ‘ventral nucleus’ of the thalamencephalon.

The *pigment* in the posterior part of the thalamencephalon is arranged

in a manner similar to that in the optic lobes and _crura cerebri_;

anteriorly where the third ventricle is prolonged forwards and

downwards the arrangement is different; the pigment lies in a curved

line above the process of the ventricle, with its concavity directed

downwards, each end bifurcating, in order that one branch may pass

upwards, the other downwards.

The *pineal body*[53] is a small vesicle placed underneath the skin

above the fronto-parietal bones; in the embryo it is connected with

the third ventricle by means of the pedicle[54] already mentioned; the

skin covering the body is always paler than the surrounding skin, and

the usual cutaneous glands are absent in this part; the paler spot on

the head may always be found, but is more distinct in _Rana temporaria_

than in _Rana esculenta_. The structure on the roof of the third

ventricle, which is usually known as the pineal body, is nothing more

than a thickened portion of the choroid plexus, and consists of a group

of convoluted vessels surrounded by _pia mater_, which is described

by Wyman as being covered with ciliated epithelium (_R. pipiens_).

The true pineal body is a small body with an outer connective-tissue

capsule, derived from the _pia mater_; this encloses an irregular

mass of epithelial cells; according to de Graaf a twig of the _ramus

supramaxillaris_ reaches it subcutaneously, and a blood-vessel

accompanies the pedicle through the _foramen parietale_. According to

Darkschewitsch, the pedicle contains medullated nerve-fibres derived

directly from the brain.

[Footnote 53: Compare Ehlers, Ueber die Zirbel der Haifische, Zeit.

f. wiss. Zool. 1878, Vol. XXX; and Balfour, Development of the

Elasmobranch Fishes, chap. ix.]

[Footnote 54: Wiedersheim states that the pedicle is hollow, and

regards the part formerly known as the pineal body as a thickened

portion of this pedicle.]

The *pituitary body* (Figs. 102, 103 _Hy_) when examined with a lens

is seen to consist of two portions: an anterior, superior, and smaller

white portion, and a larger, inferior, posterior, and reddish portion.

The anterior portion has the form of a very small, flat disk, and is

enclosed in a connective-tissue capsule which sends in larger and

smaller processes. In either transverse or longitudinal section it

is seen to be formed of two horizontal layers separated by a line

of blood-vessels and connective-tissue. The upper layer consists of

a granular and reticular matrix, containing many nuclei (averaging

0.006 to 0.010 mm. diameter), and divided into irregular rounded or

polyhedral spaces by bands of tissue derived from the capsule. This

layer is more vascular than the lower. The lower layer consists of

a mass of clear, nucleated rounded or polyhedral cells (0.016 to

0.024 mm. in diameter; nuclei from 0.008 to O.012 mm. in diameter,

Reissner), pierced by very fine connective-tissue septa derived

from the capsule. The septa are, for the most part, vertical and

longitudinal (Reissner), the blood-vessels are very few.

The posterior larger portion of the pituitary body (Fig. 106) is also

compressed from above downwards, and in transverse section as an oval

outline. It possesses an external thin connective-tissue capsule,

which sends in fine processes to support a mass of convoluted tubes,

between which course a few blood-vessels; these tubes possess an outer

nucleated basement-membrane, and are lined with a single layer of

more or less cylindrical epithelium, which entirely fills the tube;

hence the tubes possess no lumen. The tubes are from 0.04 to 0.08 mm.

in diameter; the cells are clear or granular, and possess distinct,

rounded nuclei.

*e.* The *Cerebral Hemispheres* and *Olfactory Lobes*. The cerebral

hemispheres (_Lobi hemisphaerici_, Stieda; _Lobi cerebrales_, Reissner;

Centralmasse des Geruchssinns, Carus; Hemisphaeren des grossen Hirns,

Tiedemann; Grosse Hemisphaeren, Schiess; Prosencephalon, Huxley). The

olfactory lobes (_Tubercula olfactoria_, Stieda; _Lobi olfactorii_,

Reissner; Riechkolben, Schiess; Rhinencephalon, Huxley).

α. *External form* (Figs. 98, 102, 103 _Hc_ and _L.ol_). The two

cerebral hemispheres form together the largest section of the brain;

from above they are seen to be separated by a dorsal longitudinal

fissure, which is here well marked: each hemisphere is an ovoid

body with the smaller end directed forwards and continuous with the

corresponding olfactory lobe; the posterior end forms one half of the

anterior boundary of the thalamencephalon. The olfactory lobes are two

elongated, rounded bodies directly continuous with the corresponding

cerebral hemispheres, and likewise partially separated in the median

line by a dorsal longitudinal fissure: at the point of union of the

cerebral hemispheres and olfactory lobes is a faintly marked transverse

depression.

[Illustration: Fig. 106.

Section through the lower division of the pituitary body (magnified 360

times); from Stieda.

_a_ Tubes lined with epithelium.

_b_ Blood-vessels.

]

On the ventral surface the parts are again marked off from one another

by a corresponding ventral longitudinal fissure and a transverse

groove; the two longitudinal fissures being continuous, anteriorly,

between the olfactory lobes. The cerebral hemispheres appear to be

more widely separated behind (Fig. 102) than is the case on the dorsal

surface, and in the space so formed is the _lamina terminalis_ (_Lt_).

Seen from the side, the slight depression of the upper surface, between

the cerebral hemispheres and the olfactory lobes, is seen to be

continued downwards and slightly backwards to join the corresponding

groove on the inferior surface.

The longitudinal fissures are shallow and do not meet, except at one

point, at about the middle of the cerebral hemispheres (Fig. 105). The

olfactory bulbs arise superficially (Figs. 102, 103 _I_′ and _L.ol_)

from the whole length of the olfactory lobe, between the anterior

extremity (_I_′) and the posterior (_L.ol_), where they are also

attached to the cerebral hemispheres.

β. *Internal structure.* The cerebral hemispheres and olfactory lobes

are hollow, the common cavity of each side being known as the lateral

ventricle (_Ventriculus lateralis_); these ventricles communicate

with each other, and with the third ventricle (Fig. 105). The narrow

aperture by which the lateral ventricles communicate is known as the

*Foramen of Monro* _(MF_); it communicates with the third ventricle

posteriorly, and with the space between the cerebral hemispheres

anteriorly, and thus forms a common cavity (_Ventriculus communis

loborum hemisphaericorum_, Stieda).

In general terms, each ventricle may be said to be a semilunar cavity,

prolonged backwards and forwards (Fig. 107 _c_); the outer wall is

always more or less concave, while the inner varies according to

the part of the hemisphere examined. At the anterior and posterior

extremities the inner wall is convex and bulges into the cavity

(Fig. 107); in the middle portion of the cavity the inner wall

presents a longitudinal groove (_Ventriculi lateralis cornu internum_,

Reissner), (Fig. 108 _d_), and consequently the inner wall has here two

rounded masses, an upper and a lower (_Corpus striatum_, Wiedersheim),

projecting into it. By tracing them backwards and forwards, the lower

swelling is seen to increase at the expense of the upper, while at the

same time the lateral grooves disappear; the outline of the cavity

shown in Fig. 107 is then obtained. The roof of the ventricle is arched

and broader than the floor, which, in the middle part, exists only as

a vertical slit (Fig. 108), (_Ventriculi lateralis cornu inferius_,

Reissner): towards the anterior and posterior extremities it widens and

becomes shallower (Fig. 107).

[Illustration: Fig. 107.

Transverse section through the hinder portion of the cerebral

hemispheres; from Stieda.

_c_ Lateral ventricle.

_c_′ Common ventricle of Stieda.

_d_ Longitudinal fibres.

_f_ Anterior prolongation of the third ventricle.

]

The cerebral hemispheres and olfactory lobes are composed of a fine

granular matrix, enclosing spindle-shaped, rounded, or pear-shaped

nerve cells and nuclei, and containing very fine fibres. The cells

(Fig. 109 _b_) are more numerous towards the ventricle, and somewhat

sparse towards the superficial surface. The cells are of two chief

sizes, the smaller and more numerous average 0.004 mm. to 0.008 mm. in

diameter; they are found chiefly in the deeper portions of the section,

but also form a very thin irregular layer beneath the _pia mater_: the

larger cells have an average diameter of 0.010 mm. to 0.012 mm., and

are placed towards the periphery, especially in the dorsal part of the

inner wall.

[Illustration: Fig. 108.

Transverse section near the middle of the cerebral hemispheres.

_c_ Lateral ventricle.

_d_ Groove on the inner wall.

]

In this irregular collection of cells the following centres have been

described: (1) The nucleus, through which the _corpus callosum_ passes

(Köppen); (2) the lower internal or median cell-area (Osborne),

situated above the foregoing nucleus in the posterior and middle

portions of the hemispheres; (3) the upper internal cell-area (Osborne)

is the area of large cells in the dorsal part of the inner wall; (4)

the _Corpus striatum_ (Osborne) is a mass of cells between the _corpus

callosum_ and the _commissura anterior_; Köppen doubts the correctness

of Osborne’s opinion, and suggests that a group of cells found in the

wall of the third ventricle in front of ‘Meynert’s band’ may perhaps be

a _corpus striatum_.

[Illustration: Fig. 109.

From a transverse section through one of the cerebral hemispheres; from

Stieda. (Magnified 360 times.)

_a_ Epithelium of lateral ventricle.

_b_ Nerve cells.

_c_ Connective-tissue processes from the Pia mater.

]

A bundle of longitudinal medullated nerve-fibres, the ‘round bundle,’

is found on either side of the median line (Fig. 107 _d_), and near

the lower border in the posterior portion of this region; these fibres

can be traced from the posterior section of the thalamencephalon; they

diminish in number as they course through the base of the cerebral

hemispheres, and are ultimately lost in the lower anterior part of the

outer walls. A second strand of longitudinal fibres is the continuation

of that of the _Tuber-cinereum_ above described, which can be followed

to the outer wall of the posterior part of the ventricles, and to the

anterior commissure.

The commissures are: (1) The _corpus callosum_, a large bundle of

transverse fibres, seen best in a transverse section, at the point of

junction between the _lamina terminalis_ and the cerebral hemispheres,

forming an arch over the roof of the anterior prolongation of the third

ventricle. The fibres course to the inner and anterior parts of the

hemispheres, and are situated chiefly behind the _foramen Monroi_.

To this commissure must be added those fibres which unite the two

olfactory lobes, and possibly the fibres (_Commissura posterior_)

found in the roof of the third ventricle (Köppen). (2) The _Commissura

anterior_ (Stieda), a smaller set, found immediately under the floor

of the common ventricle, forming in their course outwards a curve,

with the concavity directed downwards. This commissure connects the

‘round bundles’ of opposite sides, and those fibres coursing with the

‘round bundles’ to the olfactory lobes constitute the _pars olfactoria_

of the _commissura anterior_. To this commissure must also be added

some fibres found on the ventral surface of the _commissura anterior_

and connecting the two strands of the _Tuber-cinereum_; an unusually

coarse strand of these fibres can be traced to the inner wall of the

ventricle, and is termed the _pars olfactoria interna_ by Osborne.

The general structure of the olfactory lobes resembles that of the

hemispheres; the olfactory nerves arise each by two roots, an outer

and inner. The outer root arises from the outer wall near the groove

between the corresponding hemisphere and olfactory lobe; the inner or

anterior root arises from the anterior surface of the olfactory lobe.

Both roots have a peculiar method of origin from the extremely fine

fibrillar network of the matrix (Nerve-fibre-conglomerate, Köppen), in

which are rounded dark bodies known as ‘glomeruli;’ in the ‘glomeruli’

dark points and nuclei are seen, between larger and smaller bands of

nerve-fibres. Köppen holds that all the sensory nerves of the brain

arise in a similar manner.

A decussation takes place between the two inner roots of the olfactory

nerves; possibly the external roots are connected by means of the

_commissura anterior_.

Very little *pigment* exists in the cerebral hemispheres or olfactory

lobes, the greater portion is found in the upper part of the inner

walls of the cerebral hemispheres.

The *epithelium* of the *ventricles of the brain*, like that of the

central canal of the *spinal cord*, consists of conical cells with

their bases directed towards the cavity, and their apices directed

peripherally and prolonged into distinct processes (Figs. 104, 109

_a_). In such situations, as the choroid plexuses, where nervous

tissue is absent and the cavity is completed by _pia mater_ alone, the

epithelial cells are flattened. Everywhere else it is ciliated[55]

and possesses distinct round nuclei which are as broad as the cells

themselves. The epithelium is somewhat irregularly and sparsely

pigmented; the ventral parts of the central canal of the spinal cord,

of the fourth ventricle, and of the Sylvian aqueduct are always more

pigmented than the dorsal parts.

[Footnote 55: Schmidt (_l. c._) states that the epithelium of the

central canal of the spinal cord is not ciliated.]

PLAN OF THE ARRANGEMENT OF THE NERVE-FIBRES OF THE CENTRAL NERVOUS

SYSTEM, ACCORDING TO KÖPPEN.

NERVE FIBRE NERVE-FIBRE COMMISSURES. NERVE-FIBRES

STRANDS IN STRANDS IN WHICH CONNECT

THE WHITE THE GREY THE STRANDS OF

MATTER. MATTER. WHITE FIBRES.

{1. The ventral 1. Longitudinal 1. The ventral

{ columns, fibres in commissure.

{ including the dorsal 2. The dorsal

{ the strand portion of commissure

{ of coarse the grey (rudimentary).

The { fibres. matter.

spinal {2. The lateral

cord. { columns.

{3. The dorsal,

{ columns

{ including

{ Goll’s

{ column.

{1. The ventral 1. Longitudinal 1. The ventral 1. The fibrae

{ columns, fibres of the commissure: arcuatae.

{ including Substantia _a._ Dorsal

{ the strand gelatinosa portion.

{ of coarse Rolandi, and _b._ Ventral

{ fibres which give portion.

{ dorsal off ascending 2. (The dorsal

{ longitudinal vagal and commissure

{ strand. trigeminal is wanting.)

{2. The lateral roots.

{ columns,

The { which give

medulla { off an as-

oblongata.{ cending

{ (recurrent)

{ vagus-root.

{3. The dorsal

{ columns.

{ _a._ Ascending

{ trigeminal

{ root.

{ _b._ Auditory

{ root.

{ _c._ Descending

{ auditory

{ root.

{1. The ventral 1. Longitudinal 1. The ventral 1. The fibrae

{ columns, fibres of the commissure. arcuatae.

{ including grey matter. 2. The dorsal

{ the dorsal Small ascend- commissure,

{ longitudinal ing trigeminal cerebellar

{ strand (the root. commissure

{ strand of and trochlear

{ coarse fibres decussation.

The { has dis-

cere- { appeared).

bellum {2. The lateral

and the { columns (each

pars { receives a

commis- { strand from

suralis. { the cere-

{ bellum).

{3. The dorsal

{ columns.

{ _a._ Large as-

{ cending tri-

{ geminal root.

{ _b._ Descending

{ auditory

{ root.

{1. The ventral (Wanting.) 1. The ventral 1. The fibrae

{ columns, commissure. arcuatae.

{ including the 2. The dorsal

{ dorsal longi- commissure.

{ tudinal _a._ Commissura

{ strand. posterior.

{2. (The lateral _b._ Marginal

The { columns can- commissure.

optic { not now be

lobes. { distinguished

{ from the ven-

{ tral columns.)

{3. The dorsal

{ columns: two

{ ascending

{ opticus-roots.

{1. The ventral (Wanting.) 1. The ventral 1. The fibrae

{ and dorsal commissure: arcuatae.

The { columns. _a._ Commissura

thalam- { _a._ Fibres from transversa

enceph- { the optic Halleri.

alon and { lobes. _b._ Commissura

the cere- { _b._ The ‘round anterior.

bral hemi-{ bundle.’ 2. The dorsal

spheres. { _c._ The ‘Thal- commissure,

{ amus-Tuber- the corpus

{ cinereum callosum.

{ strand.’

C. THE COVERINGS OF THE BRAIN AND SPINAL CORD.

1. The _Pia mater_ closely invests the whole of the brain and spinal

cord; it may, in fact, be regarded as the flattened, outermost layer

of the connective-tissue material which supports the nervous elements;

fine processes (Fig. 109 _c_) pass radially from its inner surface to

join the general connective-tissue matrix or neuroglia. It is usually

pigmented, those portions covering the optic lobes and spinal cord

being especially deeply pigmented: on the cerebral hemispheres it

contains very little pigment; at times, indeed, in _R. temporaria_,

pigment is absent from this part.

The _pia mater_ is continued on to the choroid plexuses and pituitary

body, and on to the pineal body by means of the pedicle; the membrane

is very vascular, and forms, especially for the cerebral hemispheres

and the olfactory lobes, a very important source for the direct supply

of blood-vessels.

2. The _Dura mater_, or lining membrane of the cranial cavity and

vertebral canal, is a connective-tissue membrane containing many

very much branched, pigmented cells. This membrane is not so deeply

pigmented as the _pia mater_, except in that portion covering the

cerebral hemispheres and the olfactory lobes, which is much darker than

the corresponding portion of the _pia mater_.

3. The *arachnoid membrane* is the layer of endothelial cells covering

the inner surface of the _dura mater_: by means of the blood-vessels

and nerves, etc., it is continued on to the _pia mater_ of the spinal

cord and brain, which it in like manner closely invests. Masses

of *calcareous crystals* are found between the epineurium and the

arachnoid (Wiedersheim) on each spinal nerve at its exit from the

intervertebral foramen, also on the trigeminal nerves in the cranium.

Additional smaller and more irregular masses are found on the dorsal

part of the _dura mater_ of the spinal cord.

D. THE VESSELS OF THE BRAIN AND SPINAL CORD. (Fig. 110.)

The arteries of these organs are derived from the internal carotids and

the _arteriae vertebrales_. As soon as the internal carotid arteries

reach the cranial cavity each divides into two branches, an anterior

(_Ramus anterior_, Schöbl[56]) and a posterior (_Ramus posterior_,

Schöbl); the anterior branches course forwards on the lower part

of the surface of the brain as far as the anterior portion of the

thalamencephalon, where they again divide; one division, _arteria lobi

hemisphaerici inferior externa_ (Schöbl), continues forwards along the

outer surface of the cerebral hemisphere and of the olfactory lobe

as far as the olfactory bulb, where it may still be distinctly seen;

it gives off branches to the adjacent parts in its course forwards.

The vessels of the two sides communicate with each other by means of

delicate transverse vessels (_Arteriae communicantes anteriores_).

The second division, _arteria lobi hemisphaerici superior interna_

(Schöbl), of the _ramus anterior_ courses on the thalamencephalon to

the dorsal surface, gives a large branch to the choroid plexus of the

fourth ventricle, and runs forwards in the dorsal longitudinal fissure;

it supplies vessels to all the neighbouring parts.

[Footnote 56: These names are those adopted by Schöbl for corresponding

vessels in the newt, the translator has accepted them as being

suitable, with slight modification, to the vessels of the frog.]

The _rami posteriores_ converge as they course backwards, and

ultimately unite to form an _arteria basilaris_, which is continued in

the median line of the under surface of the spinal cord as the _arteria

spinalis anterior_. The _ramus posterior_ gives off, in its course,

many small vessels to the neighbouring parts, and two larger vessels

on either side, one of which, the _arteria lobi optici_ (Schöbl), is

distributed to the optic lobes, while the other passes to the pituitary

body.

The branches of these vessels form a network in the _pia mater_, from

which the nervous tissue is supplied; they also send numerous branches

directly into the brain and cord, and these have a similar arrangement

for all parts of the brain with the exception of the cerebral

hemispheres and olfactory lobes; more or less vertical branches arise

from the posterior parts of the upper borders of the _rami anteriores_,

from the whole of the _rami posteriores_, and from the _arteria

basilaris_; these course upwards from their place of origin into the

corresponding part of the brain, give off a few branches in the white

matter, and then branch freely, and at somewhat sharp angles, in the

grey matter. In the _pars commissuralis_ a large branch may be traced

from the _ramus posterior_ on either side upwards into the cerebellum

almost to its upper border. The vessels of the cerebral hemispheres

and olfactory lobes seem to possess no other definite arrangement than

that described above. The grey matter seems, on the whole, to be more

vascular than the white; the vessels of the latter are chiefly arranged

radially to the surface, and run in courses which are more or less

straight; the vessels of the grey matter are more irregular and sinuous.

The _Arteria spinalis anterior_ courses along the whole length of the

spinal cord, giving off lateral branches and communicating with _rami

spinales_ (branches of the _Arteria vertebralis_). These branches form

a plexus in the _pia mater_, from which vessels pass at irregular

points into the cord; one set of small vessels, described by Reissner,

pass in a straight course from the superior longitudinal sinus towards

the _substantia reticularis_, where they divide. Other branches pass

directly from the _arteria spinalis anterior_, through the ventral

longitudinal fissure, and there divide; the twigs as a rule avoiding

the _septum medium_ and passing in greater part towards the ventral

horns. The vessels in the white matter are for the most part radial and

straight, while in the more vascular grey matter they are irregular and

more sinuous in their course.

[Illustration: Fig. 110.

Diagram to show the _Vena spinalis posterior_, the _Venae spinales

superiores_, and the origin of the _Vena jugularis interna_.

_H_ Hemispheres.

_ji_ Internal jugular.

_op_ Optic lobes.

_mo_ Medulla oblongata.

_spp_ Vena spinalis posterior.

]

The choroid plexus of the third ventricle (_Plexus choroideus

ventriculi tertii_, Reissner) lies, as already described, on the roof

of the third ventricle; it is somewhat triangular in form (_Plexus

venosus triangularis_, Schöbl), and is evidently a prolongation of

the _pia mater_, with an increased supply of vessels. It receives,

at its anterior angle, veins from the cerebral hemispheres and the

adjacent parts. The under surface is covered with a layer of ciliated

pavement epithelium. At the posterior angles of the plexus the veins

communicate on either side (Fig. 110) with the anterior lateral

prolongation of the _vena spinalis superior_, and with the internal

jugular vein. The small body usually named the pineal body, and placed

on the roof of the ventricle, is only a small thickened portion of

the plexus, and consists of a group of convoluted blood-vessels. This

plexus sends a considerable bunch of vessels into the upper part of the

third ventricle. The choroid plexus of the fourth ventricle (_Plexus

choroideus ventriculi quarti_, Reissner) is a triangular membrane,

slightly attached to the borders of the fourth ventricle. Its upper

surface is flat and bounded at either side by a large vein, the _vena

spinalis superior_; anteriorly it is bounded by the _cerebellum_.

The under surface is not flat; in the middle line is a slight furrow

corresponding to the position of a median vessel, which may, when

injected, be seen from the dorsal surface; from it a number of short

vessels pass outwards and slightly backwards to the outer border. The

connective-tissue between these vessels is pushed down into the cavity;

in this manner one obtains a double row of flattened villous-looking

bodies, which frequently have their tips pigmented. The under surface

of the plexus is clothed with flattened, ciliated epithelium.

On either side of the pituitary body is another small venous plexus

(_Plexus lateralis_, Schöbl); it communicates above with the posterior

angle of the choroid plexus of the third ventricle; externally, with

the internal jugular vein; and internally with its fellow of the

opposite side by more or less irregular transverse vessels, which,

together with a median vein from the fissure between the cerebral

hemispheres, form an irregular _circulus venosus_ around the pituitary

body. These plexuses receive blood from the lower part of the cerebral

hemispheres, the thalamencephalon, the optic lobes, and from the

pituitary body. The veins of the spinal cord open into the dorsal,

_vena spinalis posterior_ (Fig. 110 _spp_), which bifurcates at the

posterior angle of the fourth ventricle, one division passing to either

side and forming, as already described, the outer border of the choroid

plexus; anteriorly it opens into the internal jugular vein (Fig. 110

_ji_).

The blood-vessels, both arteries and veins, are irregularly pigmented,

both on the surface and in the interior of the central nervous system;

those of the cerebral hemispheres and olfactory lobes have very little

or no pigment.

DESCRIPTION OF THE FIGURES ON PLATE I.

Fig. 111. Dorsal view of the orbit, etc.; deep dissection.

_c_ Course of the palatine nerve.

_db_ Fibrous plate.

_dm_ M. depressor maxillae.

_ef_ Terminal branches of the ophthalmic nerve.

_F_ Facial with R. anterior of the glossopharyngeal.

_g,g,g_ Terminal twigs of the ophthalmic nerve.

_h_ Muscular twigs of the Ram. maxillaris.

_ics_ M. intertransversar. capitis superior.

_ii_ Twigs of upper eyelid.

_K_ Terminal twigs of the Ram. maxillaris.

_la_ M. levator anguli scapulae.

_pt_ M. pterygoideus.

_sc_ M. sterno-cleido-mastoideus.

_Sp_ Sympathetic nerve.

_t_ M. temporalis.

_tr_ Nasal branch.

Union of facial nerve with Ram. anterior of the glossopharyngeal

nerve.

_I_ Olfactory nerve.

_II_ Optic nerve.

_III_ Motor oculi nerve.

_IV_ Trochlear nerve.

_Vb_ R. palatinus }

_Vc_ R. maxillo-mandibularis } of the trigeminal

_Vc_′ R. maxillaris } nerve.

_Vd_ R. mandibularis }

_Ve(VII)_ Facial nerve.

_Vg_ Gasserian ganglion.

_VN_ Trigeminus.

_VS_ Sympathetic nerve.

_VI_ Abducens nerve.

_VII_′ Facial nerve.

_VIII_ Auditory nerve.

_X1_ Ram. anterior of the glossopharyngeal.

_X2_ Glossopharyngeal nerve.

_X3_ Pneumogastric nerve.

_XG_ Ganglion nervi vagi.

_XI_ Accessory nerve.

Fig. 112. Dorsal view of the orbit, etc.; superficial dissection.

_e_ External branch of the ophthalmic nerve.

_f_ Internal branch of the ophthalmic nerve.

_g,g,g_ Terminal twigs of the ophthalmic nerve.

_II_ Optic nerve.

_III_ Motor oculi nerve.

_IV_ Trochlear nerve.

_Va_ Ophthalmic nerve.

_Vc_ R. maxillo-mandibularis.

_Vc_′ R. maxillaris of the trigeminal nerve.

_Vd_ R. mandibularis of the trigeminal nerve.

_VI_ Abducens nerve.

_VI_′ Ciliary nerves.

Fig. 113. View of roof of mouth; mucous membrane, etc., removed.

_a_ Branch of the Ram. palatinus to the Harderian gland.

_b_ Branch of the Ram. palatinus to the intermaxillary gland.

_c_ Inosculating-branch of the Ram. palatinus.

_Ch_ Internal naris.

_d_ Cutaneous branch of the Ram. palatinus.

_db_ Fibrous plate.

_F_ Ramus mandibularis of the facial nerve.

_Gl_ Intermaxillary gland.

_HD_ Harderian gland.

_l_ Eyeball.

_m_ M. masseter.

_Ms.ob.i._ M. obliquus inferior.

_OK_ Upper jaw.

_r_ M. retractor bulbi.

_ri_ M. rectus internus.

_ri_′ M. rectus inferior.

_sc_ M. sterno-cleido-mastoideus.

_uk_ Mandible.

† Terminal branch of Ram. palatinus.

* Union of facial nerve with Ram. anterior of the

glossopharyngeal.

_III_ Motor oculi nerve.

_Vb_ Ramus palatinus of trigeminal nerve.

_Vc_ Ramus maxillo-mandibularis of the trigeminal nerve.

_Vc_′ Ramus maxillaris of the trigeminal nerve.

_Vd_ R. mandibularis.

_Ve(VII)_ Facial nerve.

_X_ Pneumogastric nerve.

_X1_ Glossopharyngeal nerve.

Fig. 114[57]. Lateral dissection of head, etc., to show the cranial

nerves, etc.

[Footnote 57: The branches of the vagus which are represented in this

figure as crossing the petrohyoid muscles ought to be underneath

them.--_Trans._]

_c_ Cardiac nerve.

_F_ Union of facial and Ram. anterior of the

glossopharyngeal nerve.

_h_ Ram. hyoideus of the glossopharyngeal nerve.

_la_ M. levator angi scapulae.

_lgd_ M. longissimus dorsi.

_m_ M. masseter.

_mt_ Ram. mandibularis of the glossopharyngeal nerve.

_M1_ Hypoglossal nerve.

_M2_ Brachial nerve.

_Oh_ Anterior cornu of the hyoid bone.

_oi_ M. obliquus internus.

_ph1_ M. petrohyoideus I.

_ph2_ M. petrohyoideus II.

_ph3_ M. petrohyoideus III.

_ph4_ M. petrohyoideus IV.

_pp_ Pharyngeal branches of the pneumogastric nerve.

_tt_′ M. temporalis.

_us_ Muscular twigs of the Ram. mandibularis of the

trigeminal nerve.

_Z_ Tongue.

_Va_ Terminal twigs of the ophthalmic nerve.

_Vc_ Ram. maxillaris of the trigeminal nerve.

_Vd_ Ram. mandibularis of the trigeminal nerve.

_Ve(VII)_ Facial nerve.

_X1_ Ram. anterior of the glossopharyngeal nerve.

_X2_ Glossopharyngeal nerve.

_X3_ Pneumogastric nerve.

_X3g_ Ram. gastrici of the pneumogastric nerve.

_X3l_ Ram. laryngeus.

_X3p_ Ram. pulmonales of the pneumogastric nerve.

_XI_ Accessory nerve of the pneumogastric nerve.

Fig. 115. Dissection of the floor of the mouth.

_dm_ M. depressor maxillae.

_F_ United facial nerve and Ram. anterior of the

glossopharyngeal nerve.

_gh_ } M. geniohyoideus.

_gh_′ }

_h_ Ram. hyoideus of the glossopharyngeal nerve.

_H_ Heart.

_Lg_ Lung.

_m_′ Ram. mandibularis of the glossopharyngeal nerve.

_M1_ Hypoglossal nerve.

_M2_ Brachial nerve.

_Oh_ Anterior cornu of the hyoid bone.

_oh_ Momohyoideus.

_ph1_ M. petrohyoideus I.

_ph2_ M. petrohyoideus II.

_ph3_ M. petrohyoideus III.

_ph4_ M. petrohyoideus IV.

_sc_ M. sterno-cleido-mastoideus.

_sh_ M. sterno-hyoideus.

_sm_ M. mylohyoideus (submaxillaris).

_smt_ M. submentalis.

_TT_ Thyroid.

_uk_ Mandible.

_us_ Muscular twigs of the Ram. mandibularis.

_X2_ Glossopharyngeal nerve.

_X3_ Pneumogastric nerve.

_X3c_ Rami cardiaci of pneumogastric nerve.

_X3l_ Ram. laryngeus of the pneumogastric nerve.

_X3p_ Ram. pulmonalis of the pneumogastric nerve.

II. THE PERIPHERAL NERVOUS SYSTEM.

E. THE CRANIAL NERVES.

(To facilitate reference the original arrangement of this part has been

altered.)

[There are ten pairs of cranial nerves in the frog, which are numbered

in order from before backwards. The mode of origin of these nerves, and

their deeper relations with the respective parts of the brain to which

they belong, have already been described.]

*1.* The *olfactory nerve* (_N. olfactorius_), [(Figs. 97, 98, 102,

103, 111 _I_) runs a very short course only a few lines in length,

and escapes from the cranium by an opening in the cartilage of the

sphenethmoid into the nasal cavity, where it divides into two branches,

each of which breaks up into a brush of filaments, to be distributed

in the olfactory mucous membrane. The nerve contains no white fibres].

(See organ of smell, p. 385.)

*2.* The *optic nerve* (_N. opticus_), (Figs. 97, 102, 103, 111 _II_)

[arises, as already described, by the optic tract, and joins with its

fellow at the optic chiasma or commissure, where part of the fibres

pass over to the opposite side (according to Michel all the fibres

cross). Each optic nerve then courses outwards, piercing the cartilage

of the cranium and so reaching the eyeball]. (See organ of sight,

p. 408.)

*3.* The *motor oculi* (_N. oculomotorius_, _Oculo-motor_, _Motor

communis_), (Figs. 102, 111, 112, 113 _III_). From its origin it

courses outwards and forwards, perforating with a slight obliquity

the cartilaginous wall of the cranium, just in front of the Gasserian

ganglion (Fig. 116 _III_); on reaching the orbit it divides into two

branches, between the _Levator bulbi_ and the _Rectus internus_ and

_inferior_. One branch, _Ramus superior_, runs over and parallel to

the _Ramos ophthalmicus Trig._ and enters the under surface of the

_Rectus superior_; the second, lower branch, _Ramus inferior_, supplies

filaments to the _Rectus internus_ and _inferior_ and to the _Obliquus

inferior_. Apparently it exchanges fibres with the ophthalmic division

of the trigeminal nerve.

[That portion of the motor-oculi between its branch to the _Rectus

superior_ and to that to the _Rectus internus_ contains a number of

nerve cells; according to Schwalbe (_l. c._, p. 235 and Pt. XII,

Fig. 4) these cells are arranged in four groups or ganglia; the second

group forms a very slight prominence, and is covered by a very fine

layer of nerve fibres, it also gives off some extremely fine nerves to

the eyeball; this is perhaps a ciliary ganglion (_Ganglion ciliare_,

Schwalbe). Beard, however, names it ‘mesocephalic ganglion.’ The other

three groups are simply clusters of cells between the fibres of the

nerve.]

*4.* The *pathetic* or *trochlear nerve* (_N. trochlearis_,

_patheticus_), (Figs. 102, 103, and 112 _IV_). From its origin it

courses forwards with and then crosses the motor-oculi; it perforates

very obliquely the cartilaginous lateral wall of the cranium, in

front of the motor oculi but above and very slightly behind the optic

foramen (Fig. 116); it runs parallel to and in company with the _Ramus

opthalmicus trigemini_, and appears to exchange a few fibres with it,

which, however, according to de Watteville, Stannius, Cuvier, and

Wyman, is not really the case. In this course it lies first to the

inner, then to the outer side of the ophthalmic, over which it passes

to supply the _Rectus superior_. [The pathetic and ophthalmic nerves

are sometimes enclosed in a common sheath (de Watteville).]

*5.* The *trigeminal nerve* (_N. trigeminus_), (Figs. 97, 102, 103,

111, 112, 113, and 114 _V_) is the largest of the cranial nerves in

the frog; from its origin it runs outwards and forwards to the skull

wall, and just before reaching this enters the large _Gasserian

ganglion_. It then passes through the cranial wall immediately in

front of the auditory capsule, and divides at once into two main

branches (Fig. 111 _Vg_, etc.), the _Ramus ophthalmicus_ and the _Ramus

maxillo-mandibularis_.

The Gasserian ganglion is a large, yellow, oval ganglion placed in

a depression in the outer wall of the cranium; it is covered by a

‘periganglionic gland,’ which is similar in structure to that of

a spinal ‘periganglionic’ gland (see p. 180), the fibrous capsule

being stronger. This ganglion receives three other nerves besides the

trigeminal, _viz._ the sixth and seventh nerves, and branches of the

sympathetic.

According to de Watteville, these nerves are arranged on the lower

surface of the ganglion, so that the sympathetic is below, the

sixth above it, then the seventh, while the fifth is uppermost;

the sympathetic splits into several bundles which join the various

branches of the ganglion; the sixth divides into two bundles, one of

which usually joins the ophthalmic, while the other makes its exit

independently; the seventh splits into two bundles, one of which

leaves the ganglion as the hyomandibular branch, the other gives some

fibres to the palatine branch of the trigeminal nerve, and is then

continued as the palatine branch of the facial.]

*a.* The _Ramus ophthalmicus_ (_Ramus nasalis_, Fischer; Ophthalmic

or Orbital Branch, Wyman; Orbito-nasal or Ophthalmic Nerve, de

Watteville), (Figs. 111 and 112 _Va_). After leaving the Gasserian

ganglion the nerve is directed forwards parallel to the side of the

cranium, between this and the eyeball. It lies beneath the _Rectus

superior_, but above all the other muscles of the eyeball and the optic

nerve. At the anterior end of the orbit it divides into two branches

(_e_ and _f_) which pass through apertures (_Foramen pro ramo nasali_)

in the cartilage of the sphenethmoid, to reach the nasal cavity, where

they lie between the cartilage and mucous membrane. In this position

the nerves supply branches to the mucous membrane and then pierce the

skull to appear on the surface (Figs. 111, 112 _g,g,g_, 114 _Va_), the

skin of which they supply. The branches inosculate with each other and

communicate with the anterior twigs of the _Ramus maxillaris_ (Fig. 114

_Vc_) and with those of the _Ramus palatinus_ (Fig. 113 _Vb_).

In its course through the orbit the _Ramus ophthalmicus_ gives off:--

(1) Near the Gasserian ganglion one, two, or three small branches,

which communicate with the trochlear nerve, and may then be traced to

the sclerotic of the eyeball; some of the twigs enter near the optic

nerve, others nearer the cornea. They are regarded as _Ciliary nerves_

(_Nervi ciliares_).

(2) A large branch, the *palatine nerve* (_Ramus palatinus_),

(Figs. 111 and 113 _Vb_, _b_). This nerve, after receiving the palatine

branch of the facial, descends on the inner border of the _Levator

bulbi_ to the mucous membrane of the mouth, to which it gives numerous

filaments; it then runs parallel and near to the median line, lying on

the base of the skull; and near the front of the orbit it divides into

three branches. The smallest (_a_) supplies the Harderian gland (_HD_);

the second (_b_) continues in the course of the original nerve, pierces

the vomer, and divides, giving filaments to the mucous membrane in its

whole course, while the terminal twigs supply the intermaxillary gland

(_Gl_) and the surrounding structures. One of these (†) ascends on the

anterior border of the sphenethmoid to the intermaxillary (internasal,

Born) space to supply the numerous glands and to inosculate by one or

two twigs with the nasal branch (_tr_) of the trigeminal. The third

branch of the palatine nerve (_c_) curves directly outwards, just

behind the palatine bone, to the inner surface of the maxillary bone;

it then courses backwards, pierces the fibrous plate (_db_) between the

eyeball and the pterygoid bone, and ultimately unites with superior

maxillary division of the trigeminal nerve (_Vc_′). Just before

piercing the fibrous plate it gives off a tolerably large nerve (_d_)

to the mucous membrane, and in its whole course it gives off twigs to

the mucous membrane and surrounding structures; many of these twigs

inosculate with other terminal twigs of the trigeminal.

[This nerve has been minutely described by Stirling and Macdonald;

these observers find that the branches form a very fine and

close plexus in the mucous membrane, etc.; fibres were traced to

blood-vessels, glands, etc., and ganglion cells were found scattered

along the nerves. The fibres are both medullated and non-medullated,

and many of the cells are described as ‘spiral cells’ (see page 201)

such as those described by Arnold in the sympathetic system of the

frog.]

*b.* The _Ramus maxillo-mandibularis_ (_Ramus maxillaris_, Ecker and

Hoffmann; Supra-maxillary, Humphry; Upper Maxillary Branch, Wyman),

(Figs. 111, 112, and 113 _Vc_). This nerve is the largest division

of the trigeminal; it runs directly outwards behind the eyeball, in

front of the auditory capsule, and between the temporal and pterygoid

muscles. After a very short course it divides into the maxillary and

mandibular nerves. The nerve gives off before its division a few fine

filaments (_i,i_) to the hinder half of the upper eyelid and to the

surrounding skin, also a branch (_h_) [not correctly drawn in Fig. 111]

which divides to supply twigs to the temporal and pterygoid muscles.

(1) The _Ramus maxillaris_ (_Ramus supramaxillaris_, Ecker; _Ramus

maxillaris superior_, Hoffmann; Upper Maxillary Branch, Wyman;

Supramaxillary, Humphry and de Watteville), (Figs. 111, 112, 113, and

114 _Vc_) runs outwards and then forwards, between the eyeball and

the outer wall of the orbit, to the margin of the upper jaw (_K_); a

large portion terminates in a number of small branches for the supply

of the skin of the lower eyelid, the upper lip, and of the parts

between the tympanic membrane and the external nares. Some of these

twigs inosculate with twigs of the palatine and ophthalmic nerves. The

second portion of the nerve unites with the palatine nerve, as already

described.

(2) The _Ramus mandibularis_ (_Ramus maxillaris inferior_, Hoffmann;

Mandibular or Lower Jaw Branch, Wyman; Inframaxillary, Humphry and de

Watteville), (Figs. 111, 112, 113, 114, and 115 _Vd_, _us_). As far as

the outer margin of the eyeball, this nerve runs parallel to and behind

the _Ramus maxillaris_, in which course it supplies branches to the

temporal and pterygoid muscles; it then curves backwards, outwards, and

downwards to the under surface of the squamosal bone, where it supplies

a twig to the _Depressor maxillae_, and then perforates the _Masseter_:

in this manner it reaches the outer surface of the mandible, just

behind the insertion of the temporal muscle; it then courses forwards,

under the skin, to the _Symphysis menti_. In this course the nerve

gives off numerous branches to the skin and surrounding parts, it also

supplies the mylo-hyoid and submental muscles; one branch (Figs. 114

and 115 _us_) is larger than the rest, and supplies the under surface

of the floor of the mouth and lower lip.

*6.* The *abducens nerve* (_Nervus abducens_, Ecker and Hoffmann;

included in the trigeminal, Wyman), (Figs. 111 and 112 _VI_). This very

slender nerve courses along the inner wall of the cranium from its

origin to the Gasserian ganglion (_q.v._) which it joins, and leaves

this in contact with the ophthalmic division of the trigeminal nerve:

the nerve then bifurcates in the orbit; the outer branch supplies

the _Rectus externus_, the inner inosculates with small twigs of the

ophthalmic division of the trigeminal, and then gives off a number of

small ciliary nerves, already described, and one special branch to the

_Retractor bulbi_.

*7.* The *facial nerve* (_N. facialis_, Ecker and Hoffmann; _Facialis_

(_Portio dura_), Wyman; Facial or Jugular Nerve, Stannius), (Figs. 111,

113, and 114 _Ve_ _VII_) arises immediately behind the trigeminal, and

runs forwards, first in company with the auditory nerve, then alone, to

the Gasserian ganglion (_q.v._), with which it unites; it again appears

at the posterior angle of the ganglion, and escapes from the cranium in

company with, and immediately behind, the _Ramus mandibularis_ of the

trigeminal, and divides at once into two branches.

*a.* The _Ramus palatinus_ at once unites with the palatine branch of

the ophthalmic division of the trigeminal nerve. Wiedersheim doubts

this arrangement, and holds that the two portions arise in common

from the Gasserian ganglion; neither has Wyman (_Rana pipiens_)

described this branch of the facial. [De Watteville states that the

_Ramus palatinus_ and the palatine branch of the ophthalmic nerve

are separated by the carotid artery. In urodeles the two nerves are

distinct.]

*b.* The _Ramus hyomandibularis_ (_Ramus jugularis_, Hoffmann;

Facial Nerve, Wyman), (Figs. 113 and 114, _Ve_ _VII_). The nerve is

directed outwards and backwards so as to pass around the bony wall

of the auditory capsule, it then crosses over the inner end of the

columella, with which it is in close contact, and is then joined, under

cover of the _sterno-cleido-mastoideus_ (_sc_), by a branch (_Ramus

communicans_) of the glossopharyngeal (see Fig. 111 *). The single

trunk (_F_) so formed is directed downwards in the posterior wall of

the Eustachian tube to just above the angle of the mandible, where it

divides into three branches, or sometimes into two, in which case a

third nerve is supplied by one of the others, usually by the _Ramus

hyoideus_.

(1) The _Ramus mandibularis_ (_Ramus mentalis_, Hoffmann; Third Trunk

of the Facial, Wyman), (Figs. 114 _mt_ and 115 _m_′) passes inside the

angle of the jaw and courses forwards, between the skin and mylo-hyoid

muscle, parallel to the _Ramus mandibularis_ of the trigeminal nerve,

as far as the symphysis. It supplies small twigs to the neighbouring

parts of the mucous membrane of the mouth.

(2) The _Ramus hyoideus_ (_Ramus jugularis_, Hoffmann; Second Trunk of

the Facial, Wyman), (Figs. 114, 115 _h_) is the largest division of

the facial; it courses forwards subcutaneously over the _deltoideus_

and the hinder fibres of the _mylo-hyoideus_ to the anterior cornu of

the hyoid bone, supplying its muscles and the skin of the throat and

sternal region.

(3) The _Ramus auricularis_ (First Trunk of the Facial, Wyman); is a

small branch, directed outwards; it supplies a branch to the walls of

the tympanic cavity, and is finally distributed in the skin under the

tympanic membrane and behind the angle of the mouth.

*8.* The *auditory nerve* (_N. acusticus_, Ecker, Hoffmann; Auditory

Nerve, Wyman). This nerve has a very short course in the cranium;

it reaches the auditory organ by a foramen (Fig. 116 _VIII_) in the

auditory capsule, and is then distributed in the ear, with which it

will be described.

*9.* The *glossopharyngeal nerve* (_N. glossopharyngeus_, Ecker;

_Ramus glosso-pharyngeus_, Hoffmann; Glossopharyngeal, Wyman;

Glossopharyngeal of the Vagus, Müller), (Figs. 102 and 103 _X_, 111,

113, 114, and 115 _X_^2) arises in common with the pneumogastric nerve,

and quits the skull with it through an opening (_Canalis nervi vagi_)

immediately behind the auditory capsule, and at once divides into two

branches; both of which lie under the _MM. intertransversarii capitis_

and are covered by the _Depressor mandibulae_.

*a.* The _Ramus anterior_ (Verbindungsast des Glossopharyngeus,

Wiedersheim; _Ramus communicans_, Hoffmann; Uniting Branch, Wyman;

Laryngeal Branch of the Vagus, Volkmann; _Communicans ad facialem_,

de Watteville), (Figs. 111, 113, and 114 _X^1_). This nerve curves

downwards and forwards around the auditory capsule and beneath the

_Depressor mandibuli_ to join the facial nerve, as already described.

*b.* The _Ramus posterior_ (_N. glossopharyngeus_, Wiedersheim; _Ramus

lingualis_, Hoffmann), (Figs. 111, 113, and 114 _X^2_). This nerve

runs downwards and forwards to the ventral surface of the pharynx,

dips underneath the _Petrohyoideus IV_, to appear again, after a short

interval, by piercing the _Petrohyoideus II_ or _III_, courses parallel

to and behind the anterior cornu of the hyoid bone, and thus reaches

the floor of the mouth. The nerve then runs forwards in a very sinuous

course, close to the median line, and between the _Geniohyoideus_

and _Hyoglossus_; in its course on the floor of the mouth it crosses

the hypoglossal nerve. It supplies the petrohyoid muscles, and gives

numerous small branches to the mucous membrane of the pharynx (_Ramus

pharyngeus_, Hoffmann).

*10.* The *pneumogastric* or *vagus nerve* (_Vagus_; _Ramus

intestinalis nervi vagi_, Fischer, Hoffmann; Vagal Trunk, Wyman;

Vaso-sympathetic, Gaskell), (Figs. 111, 113, 114, and 115 _X_^3). This

nerve arises in common with the glossopharyngeal; the two nerves leave

the skull together by an opening in the exoccipital bone (_Canalis

nervi vagi_); immediately outside the skull they acquire a ganglionic

enlargement (_Ganglion condyloideum_, _Ganglion nervi vagi_), (Fig. 111

_XG_); in this course the glossopharyngeal lies in front of the

pneumogastric, which it then leaves. The pneumogastric lies first upon

the _M. levator anguli scapulae_, then running backwards and downwards

along the hinder border of the _Petrohyoideus IV_ it comes to the side

of the pharynx; it is covered by the trapezius, and passes between the

hypoglossal nerve and the _Aorta ascendens_; arriving at the _Arteria

pulmonalis_, it gives off its terminal branches. [Gaskell has proved

that this nerve contains both sympathetic and vagal elements, and that

it is therefore really a vago-sympathetic.]

[Illustration: Fig. 116.

Right half of skull of _Rana esculenta_, seen from within.

_II_ Optic nerve.

_III_ Motor oculi nerve.

_IV_ Trochlear nerve.

_VN_ Root of the trigeminal nerve.

_Vg_ Gasserian ganglion.

_VS_ Sympathetic nerve.

_VI_ Abducens nerve.

_VII^1_ Facial nerve.

_VIII_ Auditory nerve.

_X^{1–3}_ Pneumogastric and glossopharyngeal nerves.

]

The branches and communications of the pneumogastric nerve are:--

*a.* Communications between the ganglion of the pneumogastric and the

sympathetic system (Fig. 111 _Sp_), which again connect it with the

Gasserian ganglion of the trigeminal nerve (Fig. 111 _VS_, _Vg_); (see

sympathetic system).

*b.* [The _Ramus cutaneus dorsalis_ (Fischer) is considered by Stannius

and Fürbringer to be the homologue of the _Ramus auricularis_; it

passes outwards between the _digastricus_ and _temporalis_ to be

distributed in the skin of the suprascapular region. This nerve is the

persistent portion of the _N. lateralis nervi vagi_ of the tadpole.]

*c.* During its course over the _petrohyoidei_ the vagus gives off a

few twigs (Fig. 114 _pp_), which form a fine plexus to supply these

muscles and the pharynx [also a twig to the trachea, according to

Hoffmann].

*d.* The _Ramus accessorius_ (Fürbringer, Hoffmann, Ecker), (Fig. 114

_XI_) is usually a single small nerve which runs between the

_intertransversarii_ muscles and the _trapezius_: it supplies the under

surface of the latter muscle.

*e.* The _Ramus scapularis_ (Hoffmann) is a very slender nerve arising

near the _Ramus accessorius_; it passes along the under surface of the

_trapezius_ to the inner surface of the _interscapularis_, which it

supplies.

*f.* The _Ramus laryngeus_ (_Recurrens vagi_), (Figs. 114 and 115

_X3l_) runs for some distance parallel to the pneumogastric, separated

from it by the _petrohyoideus IV_; arriving at the hinder cornu of the

hyoid bone the nerve loops round the _Arteria pulmonalis_, and divides

into two branches for the supply of the larynx.

*g.* The _Rami gastrici_ (Fig. 114 _X3g_) are usually two in number:

they pierce the partial diaphragm formed by the anterior fibres of the

_M. obliquus internus_ and terminate in the walls of the stomach.

*h.* The _Rami pulmonales_ (Figs. 114 and 115 _X3p_) also perforate the

partial diaphragm, and then course along the pulmonary arteries to the

lungs.

*i.* The _Ramus cardiacus_ (Figs. 114 and 115 _X3c_). This nerve is

usually smaller on the right side than on the left; it passes along the

dorsal surface of the pulmonary artery and _Vena cava superior_ to the

_Sinus venosus_; in this course it gives off two or three twigs to the

roots of the lungs: the two nerves communicate just before reaching the

heart and pass on to the auricular septum, whence they are distributed

to the heart.

*j.* A slender *Laryngeal nerve* arises from the vagus alongside the

_Ramus cardiacus_; it courses along the hinder end of the greater cornu

of the hyoid to the outer side of the pharynx, which it pierces to pass

to the larynx.

F. THE SPINAL NERVES.

I. *General description.* Ten pairs of nerves arise, as already

described (p. 135), from the spinal cord; each nerve has two roots, a

ventral or anterior, and a dorsal or posterior, which unite at their

points of exit from the intervertebral foramen: just before, and for a

short distance beyond this union, each dorsal root bears a ganglionic

enlargement.

*a.* The length and direction of the various *Nerve-roots* vary

greatly; the roots of the anterior spinal nerves run a very short

course, almost transversely outwards, from their points of origin to

the intervertebral foramina: the roots of the middle and posterior

nerves, in consequence of the vertebral column being considerably

longer than that part of the cord belonging to it, pass obliquely

backwards to the foramina, the hinder nerves of the _Cauda equina_

running for a considerable distance in the vertebral canal (Fig. 116).

The relations of these roots are as follows:--

(1) The roots of the first nerve (_N. hypoglossus_) arise from the

spinal cord at a point between the first and second vertebrae; they run

transversely outwards to escape between the first and second vertebrae.

The dorsal root is extremely slender.

[Illustration: Fig. 117.

The nervous system of _Rana esculenta_, from the ventral surface. (From

Icones physiologicae by A. Ecker. Pl. XXIV, Fig. 1.)

_F_ Facial nerve.

_G_ Ganglion of pneumogastric nerve.

_He_ Cerebral hemisphere.

_Lc_ Optic tract.

_Lop_ Optic lobe.

_M_ Boundary between medulla oblongata and spinal cord.

_M1–10_ Spinal nerves.

_MS_ Connection between fourth spinal nerve and

sympathetic chain.

_N_ Nasal sac.

_Ni_ Sciatic nerve.

_No_ Crural nerve.

_o_ Eyeball.

_S_ Trunk of sympathetic.

_S1–10_ Sympathetic ganglia.

_SM_ Rami communicantes of the same.

_Sp_ continuation of sympathetic into head.

_I_ Olfactory nerve.

_II_ Optic nerve.

_III_ Motor oculi nerve.

_IV_ Trochlear nerve.

_V_ Trigeminal and facial nerves.

_Va_ Ramus ophthalmicus.

_Vc_ Ramus maxillaris.

_Vd_ Mandibular branch of trigeminal.

_Ve_ Hyomandibular branch of facial.

_Vg_ Gasserian ganglion.

_Vs_ Upper end of sympathetic trunk in connection with

Gasserian ganglion.

_VI_ Abducens nerve.

_VII_ Facial nerve.

_VIII_ Auditory nerve.

_X_ Glossopharyngeal and pneumogastric nerves.

_X1_ Ramus anterior of glossopharyngeal.

_X2_ Ramus posterior of glossopharyngeal.

_X3, 4_ Branches of pneumogastric.

]

(2) The roots of the second nerve (_N. brachialis_) arise at the level

of the second vertebra and leave the vertebral canal between the second

and third vertebrae.

(3) The roots of the third nerve arise from the cord between the

second and third vertebrae, and pass out between the third and fourth

vertebrae.

(4) The fourth nerve arises by its two roots at the level of the fourth

vertebra, and quits the vertebral canal between the fourth and fifth

vertebrae.

(5) The fifth nerve arises opposite the fourth vertebra, and passes

outwards and slightly backwards to the foramen between the fifth and

sixth vertebrae.

(6) The roots of the sixth nerve are attached to the cord opposite the

fourth vertebra, and leave the vertebral canal between the sixth and

seventh vertebrae.

(7) The seventh nerve arises from the cord at a point between the

fourth and fifth vertebrae, and leaves the canal between the seventh

and eighth vertebrae.

(8) The eighth nerve quits the cord at a point opposite the

articulation between the fifth and sixth vertebrae, then runs backwards

to the eighth vertebra, and escapes between this and the ninth vertebra.

(9) The ninth nerve arises at the level of the sixth vertebra, and

passes out between the sacrum (ninth vertebra) and the urostyle.

(10) The tenth nerve (_N. coccygeus_) arises from the cord immediately

behind the ninth nerve and opposite the sixth vertebra, and courses

alongside the terminal filament to pass out through a foramen in the

urostyle (_canalis coccygeus_).

*b.* [The *Spinal ganglia* (_Ganglia intervertebralia_).

At their exit from the intervertebral foramen the two roots of each

nerve unite and bear a ganglionic enlargement, which is in intimate

connection with the dorsal root, but is merely in contact with

the ventral root. The ganglia lie[58] in the large intervertebral

depressions, upon the under surface of the _Proc. obliqui_, above and

behind, being in relation with the bodies of the vertebrae internally,

and the _Musculi intertransversarii_ externally (Fig. 119); ventrally

the ganglia are, more or less, covered by the ‘periganglionic glands.’

The ganglia vary much in size; that of the first spinal nerve is the

smallest. The ganglion is not attached to the dorsal root alone but

is prolonged a short distance beyond the point of union of the two

roots; this is best seen in the nerves of the lumbar plexus. Each

ganglion is a yellowish-white rounded or oval body, with an outer thick

(sometimes 0.15 mm. thick) sheath of connective-tissue, connected by

the epineurium of the nerve roots with the _Dura mater_. This sheath is

composed of bundles of parallel, wavy, white, connective-tissue fibres,

which enclose a large number of connective-tissue corpuscles, and here

and there contains brown pigment.

[Footnote 58: The _Gangl. coccygeum_ is in the urostyle.]

[Illustration: Fig. 118.

Ventral view of the brain and spinal cord, to show the points of exit

of the spinal nerves.

_M1–10_ Spinal nerves.

_VG_ Trigeminal ganglion.

_W1–10_ Vertebrae.

_XG_ Ganglion of vagus.

]

By means of longitudinal and transverse sections it may be seen

that the ganglion surrounds the ventral root, but that it forms a

thinner layer on that side, where the dorsal root joins the ventral;

although at this point the two roots and the ganglion are firmly

united together, the thick fibrous sheath of the ganglion prevents any

communication between the nervous elements of the ventral root and

those of the ganglion.

The sheath contains nerve-cells and fibres, and sends in a few fine

processes to support these structures. The fibres of the posterior

root enter and pass out of the ganglion without suffering any changes,

except that the nerve fibres are slightly separated by the presence of

a few nerve-cells, and that a distinct increase in the number of nerve

fibres takes place. The nerve-cells, of which the ganglion is chiefly

composed, and upon the number of which its size directly depends, are

arranged chiefly around the fibres of the dorsal root; such few as lie

between the fibres are smaller than the rest and vary more in their

relative numbers; at times one or two isolated cells are found in the

capsule or even in the adjoining ‘periganglionic gland.’ The cells near

the capsule are somewhat smaller than the deeper cells.[59] Each of

these cells, which are usually pear-shaped, possesses only one process;

the cell membrane is thick, resistant, and possessed of an external

nucleated, endothelial covering, the space around being probably

a pericellular lymph-space; it often contains one to three small

fat-globules; the protoplasm of these has, according to v. Lenhossék, a

concentric fibrillation; the nucleus is round, clear, and distinct, and

relatively larger the smaller the nerve-cell; it is usually placed in

the centre of the cell.

[Footnote 59: Larger cells of deeper layer 80 µ to 90 µ in diameter,

sometimes one or two as large as 0·1 mm. v. Lenhossék.]

[Illustration: Fig. 119.

Ventral view of the spinal ganglia; on the right side they are still

hidden by the ‘periganglionic glands.’ ]

[Illustration: Fig. 120.

Schema of spinal ganglion.

_A_ Ventral root.

_C_ Ramus communicans.

_G_ Ganglion

_O_ Dorsal division.

_P_ Dorsal root.

_R_ Ventral division.

]

In that portion of the cell towards the process is a portion brighter

and less easily stained than the rest of the cell (Polarkernen,

Courvoisier); this apparently possesses one or two nuclei, and is

regarded as a cell by Lenhossék (Polarzellen); the process of each cell

soon acquires a medullated sheath, and after a short course, 0.09 mm.

to 0.35 mm., divides usually about the third constriction (Lenhossék).

These processes all pass peripherally, even those which appear at first

to pass in the opposite direction curve round again; it is chiefly due

to these fibres that the spinal nerve is larger than its two roots.

In addition to the above, smaller, irregular cells from 5 µ to 7 µ in

length are found; Rawitz regards them as young cells, Lenhossék as

cells capable of (entwicklungsfähige) developing into nerve-cells. The

ganglia contain only few blood-vessels.]

The above description is founded upon that of Lenhossék, and has been

confirmed by the translator in every particular, except that only

very indistinct and unsatisfactory indications of the concentric

fibrillation so clearly delineated by Lenhossék were obtained; the

structure of the Gasserian ganglion was found to correspond exactly

with the description here given of the structure of the spinal ganglia.

The nucleus was oftener at the side than at the centre of the cell.

*c.* The ‘*Periganglionic Glands*’ (Periganglionäre Kalkdrüsen, v.

Lenhossék; Kalksäckchen, Ecker, and others; Crystal capsules, Wyman;

Calcareous Sacs or Masses, of other writers), (Fig. 119). These bodies

are found on the ganglia of all the spinal nerves and on the Gasserian

ganglion of the trigeminal nerve; they have recently been carefully

investigated by v. Lenhossék. According to his description there are

usually two to each ganglion, lying on its sides and ventral surface;

each consists of a connective-tissue capsule which sends in a few fine

trabeculae to support a system of glandular tubes: these usually run

parallel to the long axis of the gland, and are about 14 µ to 15 µ in

diameter; each tube is lined with a single layer of somewhat columnar

epithelium, the cells of which have sharp, distinct outlines and oval

nuclei.

A _membrana propria_ was not made out with certainty; the lumen of

the tubes is wide and contains a milky fluid, which gives the whole

structure its characteristic appearance; it has long been known

(Blasius, 1681, mentions the fact, _l.c._, p. 291) that this fluid

contains calcareous matter, which effervesces and dissolves in the

presence of hydrochloric acid; the crystals vary in form but are

chiefly oval (Wyman).

These glands bear no definite proportion to the size of the nerves to

which they are attached (Fig. 118); they already contain calcareous

matter during the tadpole stage before the limbs are developed

(Stannius, Wyman). No ducts have been traced to these glands. This

description, founded upon that of Lenhossék, can be confirmed by the

translator in every particular; his sections, however, would lead him

to believe that the glands are far more vascular than the description

and the drawings of Lenhossék imply.]

*d.* The *Branches* of the complete spinal nerves formed by the union

of the dorsal and ventral roots with the ganglion (Fig. 120) are

usually described as two, a dorsal and a ventral branch, which latter

at once gives off a _Ramus communicans_ to the sympathetic system;

these two branches are given off almost immediately beyond the ganglion.

II. The Individual Nerves.

A. The Dorsal Branches (Fig. 121).

[Illustration: Fig. 121.

Dorsal branches of the spinal nerves. The _Fascia dorsalis_ is

dissected, removed from the left side.

_rc_ Rami cutanei.

_rm_ Rami musculares.

]

The dorsal branches are smaller than the ventral; they pass upwards

between the inner borders of the _Musc. intertransversarii_ and the

articular processes of the vertebrae to reach the under surface of the

_M. longissimus dorsi_; the general course of these nerves is very

similar for all; each nerve gives off--

(_a_) A twig to the _Musc. intertransversarius_ and then divides into

two branches.

(_b_) A _Ramus muscularis_ (Fig. 121 _rm_), which passes outwards to

supply the _M. longissimus dorsi_.

(_c_) A _Ramus cutaneus_ (Fig. 121 _rc_); this nerve continues under

the extensor muscle of the back, running on the arch of the vertebra

towards the middle line; it then pierces the muscle and Fascia dorsalis

to reach the large dorsal lymph-sac (_Sacc. cranio-dorsalis_), through

which it courses to end in the skin. In its course through the sac the

nerve is connected with a small artery and vein by a small amount of

connective-tissue, the whole being enclosed in a sheath of endothelium

continuous with that of the general cavity.

The three anterior cutaneous nerves pierce the _Fascia dorsalis_ near

the spinous processes, the fourth a little to the side, the fifth

perforates the _M. coccygeo-iliacus_ at about the junction of its

anterior and middle thirds; the sixth and seventh perforate the same

muscle, but more posteriorly. There are seven of these _Rami cutanei_,

the first coming from the second spinal nerve, the rest from the

succeeding six spinal nerves.

B. The Ventral Branches.

The ventral branches of the spinal nerves are larger than their

corresponding dorsal branches: each nerve gives off immediately a

_Ramus communicans_ to the sympathetic system (Figs. 117 _SM_, 120

_C_); these _Rami communicantes_ will be described with the sympathetic

system; the remaining portions of the ventral divisions of the original

spinal nerves are usually known as the spinal nerves; they will be

described as such in detail.

*1.* The *hypoglossal nerve*[60] (_N. hypoglossus_, _N. spinalis I_),

(Figs. 114 115, and 117 _M1_), or first spinal nerve, is of small

size; it leaves the vertebral canal between the first and second

vertebrae to lie between the _Musculi intertransversarii_ and the

pharynx, where it has in front the _Arteria vertebralis_, behind the

_Arteria_ and _Nervus brachialis_; passing under the _Levator anguli

scapulae_, it touches the _Aorta ascendens_ and crosses the _Vagus_

and the Carotid-gland, and thus reaches the space between the _Musculi

sterno-hyoidei_ and the _Petro-hyoideus_; then curving forwards sharply

it turns under the _Mylo-hyoideus_ to course between the fibres of the

_Genio-hyoideus_ to the root of the tongue, where it ends. It first

lies to the inner and then to the outer side of the glossopharyngeal

nerve. In its course the nerve gives off the following branches:--

[Footnote 60: [Observers have differed considerably as to the relations

of this nerve: Volkmann describes it as the first nerve of the neck,

and describes a ganglion on its dorsal root; Vogt denies the presence

of this ganglion, and regards it as a true cranial nerve; Stannius

asserts that no ganglion exists, and holds the nerve to represent the

first two spinal nerves; Wyman describes the two roots and regards it

as a spinal nerve; Hoffmann regards this nerve as the second spinal

(N. spinalis II); Fürbringer as the representative of two spinal

nerves; de Watteville describes two roots and names it first spinal

nerve.--TRANS.]]

(_a_) One or two communicating twigs (Figs. 114 and 122) to the second

spinal nerve.

(_b_) Muscular twigs to the _Longissimus dorsi_, _Intertransversarii

capitis_, _Levator anguli scapulae_, and the _Retrahens scapulae_.

(_c_) When near the glossopharyngeal it supplies twigs to the

_Geniohyoideus_, _Sternohyoideus_, and _Omohyoideus_, where it

bifurcates.

(_d_) One of the branches passes inwards to supply the _Hyoglossus_.

(_e_) The other terminal branch passes forwards with the _Ramus

lingualis_ and the glossopharyngeal, between the fibres of the

_Geniohyoideus_, which it partially supplies, and sends twigs to the

neighbouring parts.

(_f_) [Hoffmann describes a communicating branch to the pneumogastric

nerve, which the translator has not been able to discover, and which no

other observer has mentioned.]

*2.* The *second spinal* or *brachial nerve* (_N. brachialis_, _N.

spinalis II_), (Fig. 122) is a large nerve, leaving the vertebral canal

between the second and third vertebrae; it then accompanies the _Art.

axillaris_ along the anterior border of the transverse process of the

third vertebra, over the hinder end of the _Musc. levator scapulae_ and

the anterior end of the _m. transverso-scapularis major_, to the inner

border of the _Musc. subscapularis_, under which it passes into the

arm. In this course it gives off or receives the following branches:--

(_a_) It first receives a branch from the second spinal nerve.

(_b_) It gives off a large branch, the _N. coraco-clavicularis_

(Fig. 122 _Cc_), which first accompanies a branch of the _Art.

axillaris_ on the _Musc. levat. scapulae_, then courses forwards and

outwards over the _Musc. subscapularis_ to pass from above into the

_Foramen ovale_ between the clavicle and coracoid bones. It gives off

two branches:

(1) One immediately beyond its origin, which runs backwards over the

_Musc. subscapularis_ to supply the _Musc. obliq. abdominis internus_

(Fig. 122 _Cc_′).

(2) While in the _Foramen ovale_, the _N. coraco-clavicularis_

bifurcates; the anterior twig runs forwards and outwards to the

_deltoideus_ and also supplies a recurrent filament to the _Musc.

sterno-radialis_; the posterior twig enters the upper surface of the

_Musc. stero-radialis_.

(_c_) At the outer extremity of the _Musc. transverso-scapularis_ the

brachial nerve gives off a posterior branch, which at once bifurcates:

(1) The first branch passes into the under surface of the _Musc.

latissimus dorsi_ (_ld_), giving a twig to the _Infraspinatus_.

(2) The second is the _Ramus cutaneus axillaris_ (_IIc_); it passes on

the under surface and posterior border of the _Musc. latissimus dorsi_

to the skin of axilla and dorsal surface of the upper arm.

(_d_) Several small branches are given off from the upper and anterior

surfaces of the brachial nerve near the _Foramen ovale_; they

accompany an artery to pass forwards and upwards between the _Musc.

subscapularis_ and the _M. transverso-scapularis_ on the one side, and

the long head of the _M. triceps brachialis_ on the other, and are lost

in the _M. deltoideus_ and the _M. infraspinatus_.

[Illustration: Fig. 122.

The brachial plexus.

_Cc_ N. coraco-clavicularis.

_Cc_′ Nerve to musc. obliq. abdom internus.

_d_ Deltoid muscle.

_la_ Musc. lev. ang. scap.

_ld_ Musc. lat. dorsi.

_oi_ Musc. obliq. abdom. intern.

_sc_ Musc. sterno-mastoid.

_ss_ Musc. subscapularis.

_SI_ First spinal nerve.

_SII_ Second spinal nerve.

_SIII_ Third spinal nerve.

_IIc_ Ramus cutaneus axillaris.

_IIIc_ Ramus cutaneus abdominalis.

]

The _N. brachialis_ continues its course by passing between the long

and inner heads of the _M. triceps_, reaches the arm and splits into

two nerves, the _N. ulnaris_ and the _N. radialis_.

[*I.*] The _N. ulnaris_ (Fig. 123 _U_) escapes from cover of the long

head of the _M. triceps_ and runs obliquely over its inner head

towards the outer side of the _Plica cubiti_; in this course it

supplies four branches:

α. The _Ramus subscapularis_ passes dorsally to the _M. subscapularis_.

β. A _Ramus pectoralis_ (Fig. 123 _up_), passes forwards to the _M.

abdomino-pectoralis_ and to the skin of the pectoral region (_R.

cutaneus pectoralis_).

γ. The _N. cutaneus antibrachii superior s. medialis_ (_uc_) passes to

the skin of the inner side of the forearm; this and the two following

nerves are given off near the _Plica cubiti_.

[Illustration: Fig. 123.

Nerves of the central surface of the arm. The hand pronated.

_rc_ Upper cutaneous branch of the N. radialis.

_rc_′ Lower cutaneous branch 4-,f the N radialis

_U_ N. ulnaris.

_uc_ R. cutan. sup. of the N. ulnaris.

_uc_′ R. cutan. inf. of the N ulnaris.

_up_ R. pectoralis of the N. ulnaris.

]

δ. The _N. cutaneus antibrachii inferior s. dorsalis s.

musculo-cutaneus_ (_uc_′) at once supplies branches to the _M. flexor

carpi radialis_[61], and then runs downwards upon the muscle and

bifurcates:--

[Footnote 61: As this muscle increases in size during the breeding

season. it would be interesting to know whether a corresponding change

takes place in the nerve.]

[1] The _R. lateralis_ is distributed, by two twigs, in the _M. flexor

carpi ulnaris_ and in the skin of the second finger, to which it

supplies the _Rami digitales volaris_ and _dorsalis_.

[2] The _R. dorsalis_, after giving a cutaneous twig to the second

finger, is distributed by numerous twigs to the skin of the dorsal

surface of the hand.

The _N. ulnaris_ then passes between the _M. flexor carpi radialis_ and

the tendon of the _M. sterno-radialis_, sinks deeply into the _Plica

cubiti_, where it lies between the _M. flexor carpi radialis_ and

the _Flexor carpi ulnaris_, then between the latter and the _Flexor

anti-brachii medialis_ on the one side and the _Flexor digitorum

communis_ on the other: it supplies branches to all these muscles, and

while still in the forearm divides into two terminal branches:--

α. The _R. ulnaris medialis_ is the smaller; it runs inwards, passes

under the tendon of the _M. extensor carpi ulnaris_ into the palm of

the hand and ends as the _N. volaris digiti V medialis_.

β. The _R. ulnaris lateralis_ passes near the thumb-rudiment, covered

by the _M. abductor pollicis_, deeply into the palm of the hand, and

supplies by bifurcating branches the adjacent sides of the second,

third, fourth, and fifth fingers (_R. volares_); it also gives twigs to

the muscles of the palm.

[II.] The _N. radialis_ (Figs. 123, 124, and 125 _R_) passes immediately

beyond its origin from the _N. brachialis_, between the humerus and the

_M. triceps_, runs along the bone to its outer side; in this course it

gives off:--

[Illustration: Fig. 124.

Nerves of the ventral surface of the arm. The hand supinated, and the

superficial layer (Flex. carpi rad. and uln., Flex. dig. commun.) has

been removed.

_R_ N. radialis.

_u_ N. ulnaris.

_uc_ R. cutan. inf. of the N. ulnaris.

_um_ Bifurcation of the N. ulnaris.

]

α. _Rami musculares_ to the separate parts of the _M. triceps_.

β. The _Ramus cutaneus superior_ (Fig. 125 _rc_) to the skin of the

outer surface of the upper arm and forearm.

The nerve then sinks deeply under the origins of the _M. extensor carpi

ulnaris_ and the _M. extensor digitorum communis_, where it bifurcates

after giving off some small twigs:--

γ. Small muscular twigs to the extensor muscles of the hand and fingers.

[Illustration: Fig. 125.

The N. radialis.

_ec_ M. extensor carpi ulnaris.

_ed_ M. extensor digit. comm.

_R_ N. radialis.

_R1_ R. lateralis.

_R2_ R. medialis.

_rc_ R. cutaneus superior.

_rc_′ R. cutaneus inferior.

]

δ. The _R. radialis medialis_ (_R2_), the smaller terminal branch,

supplies a branch to the _M. extensor digit. comm. longus_ and the skin

over the carpus (_Ramus cutaneus inferior_, _rc_′), and ends on the

muscles of the little finger as the _R. dorsalis digiti V medialis_.

ε. The _R. radialis lateralis_ (_R_), the larger of the two branches,

supplies the extensor muscles of the fingers and gives bifurcating

branches, _Rami digitales dorsales_, to the adjacent sides of the

second, third, fourth, and fifth fingers (the _Ramus dorsalis

lateralis_ of the second finger is supplied by the _N. ulnaris_).

*3.* The *third spinal nerve* (_N. spinalis III_) (Fig. 122 _SIII_)

leaves the vertebral canal by the foramen between the third and fourth

vertebrae, and runs outwards and forwards upon the large transverse

process of the third vertebra to the brachial nerve (_N. spinalis II_).

Its behaviour at this point is subject to considerable variation; at

times it joins the brachial nerve completely, at other times the two

nerves are simply in contact; or lastly, it may send a small twig to

the brachial nerve. Externally to this point the two nerves supply a

number of small branches, which may belong almost entirely to either

one nerve or the other; these nerves again inosculate in a variable

manner, to form an axillary plexus. The more constant branches are:--

(1) One or more twigs to the _Musc. transverso-scapularis major_.

(2) Several branches, usually two larger and one smaller; they course

in the _M. obliq. abdom. internus_ (Fig. 122 _oi_) to the _M. rectus_,

where they give off branches, then pierce the muscle to reach the skin

(_R. cutaneus abdominalis_), (Fig. 122 _IIIc_).

*4.* The *fourth spinal nerve* (_N. spinalis IV_) (Fig. 126 _M4_)

appears between the fourth and fifth vertebrae; it runs on the ventral

surface of the transverse process of the fifth vertebra and upon the

_Musc. intertransversarius_ obliquely outwards and downwards, and

reaches the deeper surface of the _Musc. obliquus internus_, into which

it descends, about opposite the articulation between the eighth and

ninth vertebrae; it then bifurcates into:--

(1) A _Ramus cutaneus abdominalis_, which pierces the muscle and

supplies the skin.

(2) A _Ramus muscularis_, which supplies the broad abdominal muscle and

the _M. rectus abdominis_.

*5.* The *fifth spinal nerve* (_N. spinalis V_) (Fig. 126 _M5_)

emerges through the foramen between the fifth and sixth vertebrae,

runs obliquely downwards and outwards over the transverse processes

of the sixth and seventh vertebrae and the corresponding _M.

intertransversarii_, and at a point nearly opposite the middle of the

urostyle pierces the _M. obliquus internus_.

The rest of its course corresponds exactly with that of the fourth

spinal nerve.

*6.* The *sixth spinal nerve* (_N. spinalis VI_) (Fig. 126 _M6_),

after escaping from the vertebral canal between the sixth and seventh

vertebrae, runs obliquely downwards and outwards on the under

surfaces of the transverse processes of the seventh, eighth, and

ninth vertebrae, then under the iliac bone to a point opposite the

hinder half of the urostyle, where it descends under cover of the _M.

obliquus abdom. internus_. In the rest of its course it repeats the

corresponding courses of the fourth and fifth spinal nerves.

[Illustration: Fig. 126.

Ventral view of the brain and spinal cord, to show the points of exit

of the spinal nerves.

_M1–10_ Spinal nerves.

_VG_ Trigeminal ganglion.

_W1–10_ Vertebrae.

_XG_ Ganglion of the vagus.

]

*7*, *8*, *9*, and *10*. The *seventh*, *eighth*, *ninth*, and *tenth

spinal nerves* (_Nervi spinales VII_, _VIII_, _IX_, and _N. spinalis X

s. N. coccygeus_) (Figs. 126 and 127 _M7_, _M8_, _M9_, and _M10_) are

best described together, as they are intimately associated to form the

sciatic plexus (_Plexus ischiadicus_, _Plexus cruralis_).

Within the vertebral canal the roots of these nerves form the _Cauda

equina_; the seventh nerve leaves the canal between the seventh and

eighth vertebrae, the eighth nerve between the eighth and ninth

vertebrae, the ninth between the sacrum or ninth vertebra and the

urostyle, and the tenth by the _Canalis coccygeus_ in the urostyle;

from these points the four nerves run obliquely on the ventral surface

of the _M. ilio-coccygeus_ to the pelvis, where they form the sciatic

plexus. Although subject to some variation, the usual arrangement is

that the seventh and eighth nerves unite to form a trunk, which then

receives the ninth nerve; the large nerve so formed is the sciatic

nerve: a branch of the tenth usually then joins either the sciatic

plexus or the sciatic nerve; the plexus gives off or receives the

following branches:--

(_a_) Like other spinal nerves, these nerves give off _Rami

communicantes_ (see Sympathetic System).

(_b_) The _M. ilio-hypogastricus_ (Fig. 127 _M7a_). This arises

from the seventh spinal nerve, before its union with the eighth

spinal nerve; it runs obliquely outwards and downwards on the _Musc.

ilio-coccygeus_ and under the border of the _M. obliquus abdom.

internus_, where it bifurcates:--

(1) One branch, the _Ramus cutaneus abdominalis_, pierces the _M.

obliq. abdom. internus_ in the same fashion as the cutaneous branches

of the fourth, fifth, and sixth spinal nerves; it supplies the skin of

the abdomen.

(2) The second or _Ramus muscularis_ supplies the flat abdominal

muscles and the _M. rectus abdominis_.

(_c_) The _M. cruralis_ (Fig. 127 _M7b_) also arises from the seventh

spinal nerve at or just beyond its point of union with the eighth

spinal nerve. It accompanies the _A. cruralis_ on the pelvic wall to

the thigh, where it lies upon the _M. ilio-psoas_ in a triangle between

the _M. rectus fem. anterior_ and the _M. adductor longus_, and divides

into two main branches:--

(1) Muscular twigs to the _M. ilio-psoas_, _M. rectus femoris anticus_,

_MM. adductores longus_ and _brevis_.

(2) The _Ramus cutaneus femoris_, which runs downwards in the hinder

wall of the Lymph-sac (_Saccus iliacus_), and supplies the skin of the

under and outer surfaces of the thigh.

[Illustration: Fig. 127.

The sciatic plexus, from a drawing by Wiedersheim.

_a_ Communicating nerve from the ninth spinal nerve.

_a_′ Nerve to musc. lev. ani.

_b_ Nerve to oviduct.

_c_ Urostyle.

_cc_ Communicating twig from the ninth spinal nerve.

_ci_ M. coccygeoiliacus.

_c.I_ M. coccygeosacralis.

_d_ Nerve to bladder.

_d_′ Twig to the m. iliacus.

_e_ Twig from sciatic plexus to coccygeal plexus.

_i_ M. intertransversarius.

_il_ M. ilio-coccygeus.

_ip_ M. ilio-psoas.

_lev_ M. lev. ani.

_oi_ M. obliq. abdom. internus.

_t_′ Transverse process of sacrum.

_Ao_ Aorta.

_Com_ Point of union of nerves _a_ and _cc_.

_Is_ Ischium.

_M5_ Fifth spinal nerve.

_M6_ Sixth spinal nerve.

_M7_ Seventh spinal nerve.

_M8_ Eighth spinal nerve.

_M9_ Ninth spinal nerve.

_M10_ Tenth spinal nerve.

_M7a_ M. iliohypogastricus.

_M7b_ M. cruralis.

_Ni_ Sciatic nerve.

_Ovd_ Oviduct.

_Rect_′ Rectum.

_S7_, _S8_, _S9_, _S10_ Rami communicantes of seventh,

eighth, ninth, and tenth spinal nerves.

_Ves_ Bladder.

8, 9 Eighth and ninth vertebrae.

]

(_d_) The _Ramus dorsalis_ is a very small nerve; according to Waldeyer

it possesses only twenty nerve-fibres. It arises from the coccygeal

nerve immediately beyond the _Canalis coccygeus_, perforates the _M.

ilio-coccygeus_, and runs on its dorsal surface obliquely over the

lymph-heart, without supplying it; the nerve then pierces the fascia to

supply the skin of the back and thigh.

(_e_) The _Ramus abdominalis_ also arises from the coccygeal nerve; it

is larger than the _R. dorsalis_, and arises at the same point; the

nerve runs on the ventral surface of the _M. ilio-coccygeus_ towards

the lymph-heart, and inosculates with the sympathetic. It is from this

nerve that the branch to the sciatic plexus is usually given off; it

further supplies a varying number of branches, which with branches

derived directly or indirectly from the sciatic nerve, form a plexus

(_Plexus coccygeus_); the two plexuses may together be regarded as a

_Plexus ischio-coccygeus_ or a _Plexus sacro-coccygeus_.

(_f_) In addition to the _Rami communicantes_ given off by the spinal

nerves near the vertebral column, the sciatic plexus supplies a few

twigs, usually two (Fig. 127).

(_g_) Other small twigs pass from the sympathetic system forwards and

outwards to join the sciatic plexus or the sciatic nerve. According to

Waldeyer two of these are very constant.

(_h_) Branches to the oviduct (Fig. 127 _b_).

(_i_) Branches to the rectum (Fig. 127 _cc_).

(_k_) Branches to the bladder (Fig. 127 _d_).

(_l_) Branches to the _M. levator ani_ (Fig. 127 _a_′).

(_m_) A branch to the lymph-heart, which runs along the anterior border

of the _M. lev. ani_, on to its dorsal surface, and then direct to the

lymph-heart.

*Variations* in the *sciatic plexus*. The arrangement of the nerves

in the sciatic plexus is subject to many variations; according to

Wiedersheim most of these variations belong to two chief classes:

either the _N. coccygeus_ inosculates directly with the ninth

spinal nerve by one or several branches, or it joins the ninth

nerve indirectly by uniting with its branches. A case of the latter

arrangement is seen in Fig. 127, and is thus described by Wiedersheim:--

‘After cutting through the pelvic symphysis and drawing to one side

the contained viscera, namely, the hinder end of the oviduct, the

rectum, and the bladder, one sees a row of small twigs (_a_, _b_, _cc_,

_d_, and _e_) arising from the inner, hinder, and anterior surfaces

of the ninth spinal nerve: the first (_a_) runs backwards parallel

with the _N. coccygeus_ over the _M. ilio-coccygeus_ to join this

nerve at the point marked _Com_. The twig _cc_ behaves in like manner

after receiving a twig from _b_. The twig _b_ arises from the inner

surface of the ninth nerve between the two foregoing and close to its

union with the eighth spinal nerve; from this origin twig _b_ passes

almost transversely outwards to the hinder extremity of the oviduct

(“uterus”), and partly to the rectum, crossing in its course twig _a_,

the sciatic nerve, and the urostyle. Its branches form a net-like

plexus with the terminal branches of the last sympathetic ganglion and

with a branch _cc_ from the point of union _Com_.

‘A second branch (_a_′), arising from the point of junction _Com_, is

a continuation of the _N. coccygeus_ (_M10_); it passes vertically

downwards towards the hinder end of the cloaca and at the upper border

of the _Levator ani_, divides into two branches, which are distributed

to the inner and outer surfaces of this muscle and to the cloaca. Other

branches pass dorsally to the lymph-hearts, while a third set pass to

the hindermost part of the bladder.

‘These three sets of nerves, to the _M. coccygeus_, _M. levator

ani_, and to the lymph-hearts, are not supplied entirely by the _N.

coccygeus_, as this is reinforced by one or more branches (_e_) from

the sciatic plexus: this branch (_e_) supplies twigs to the _M.

ilio-coccygeus_, which is also supplied anteriorly from the trunk of

the _N. coccygeus_.

‘The bladder receives a special branch (_d_), which arises from the

sciatic plexus at the junction of the eighth and ninth spinal nerves;

this nerve gives a twig (_d_′) to the _M. iliacus_.’

*I.* The *sciatic nerve* (_N. ischiadicus_) (Fig. 128 _I_) is the

largest nerve of the body; it passes under the _M. coccygeus_, between

the origins of the _M. vastus externus_ and of the _M. pyramidalis_:

lies then between the _M. biceps_ and the _M. pyramidalis_, and later

between the _M. biceps_ and the _M. semimembranosus_; lastly, it

bifurcates under the _M. biceps_ to form the _N. tibialis_ (_II_) and

the _N. peroneus_ (_III_). In this course it gives off:--

(_a_) A twig to the _M. coccygeo-iliacus_, while still in the pelvis.

(_b_) The _N. cutaneus femoris posterior_ (Fig. 128, 2), which passes

between the _M. pyramidalis_ and the _M. vastus externus_, to appear

behind and beneath the former; it accompanies an artery of like name to

supply the skin of the hinder and inner surfaces of the thigh.

(_c_) A little below the foregoing it gives off a collateral branch

(Fig. 128, 3), which passes under the _M. pyramidalis_ and divides to

form--

(1) A branch to the upper third of the _M. semimembranosus_.

(2) A branch to the _Rectus internus minor_. A twig of this branch

(_bb_) pierces the muscle transversely in company with the _Arteria

cutanea_, and passes to the skin of the middle of the inner surface of

the thigh.

(3) Branches to both heads of the _semitendinosus_.

(4) Branches to the _M. adductor magnus_.

(_d_) Branches covered by the _M. pyramidalis_ to the _M. quadrat.

femoris_ and _M. obturator_.

(_e_) A branch (5) forwards to the _M. biceps_.

(_f_) A branch (6) which accompanies the _Art. circumflexa genu

lateralis sup._ forwards to the _M. extensores cruris_, the _M. vastus

externus_, and the _M. rectus anterior_.

[Illustration: Fig. 128.

Distribution of the sciatic nerve.

_I_ The sciatic nerve.

_II_ N. tibialis.

_III_ N. peroneus.

1 Branches to the M. pyramidalis.

2 N. cut. fem. posterior.

3 Large collateral branch.

5 Branch to the M. biceps.

6 Branch accompanying the art. circumfl. genu lateralis sup.

7 Branch of tibial nerve to the M. gastrocnemius.

8 N. suralis.

_ad_‴ M. adductor magnus.

_b_ M. biceps.

_bb_ Twig accompanying the art. cutanea.

_ci_ M. coccygeo-iliacus.

_cp_ N. cut. cruris lateralis.

_ct_ R. cut. cruris posterior.

_g_ M. gastrocnemius.

_gl_ M. glutaeus.

_p_ M. pyramidalis.

_pe_ M. peroneus.

_ra_ M. rectus anterior.

_ri_′ M. rectus internus minor.

_sm_ M. semimembranosus.

_st_ M. semitendinosus.

_ve_ M. vastus externus.

]

*II.* The _Nervus tibialis_ (Figs. 128 _II_, 129 _T_) passes backwards

and inwards to supply the extensors of the foot and the flexors of the

toes. Its branches are:--

(_a_) The _R. cutaneus cruris posterior_ (Figs. 128 and 129 _ct_),

accompanied by an artery and vein, passes to the skin of the calf.

(_b_) A twig to the upper part of the gastrocnemius (Fig. 129 _g_′).

The main nerve, after supplying this twig, bifurcates to form the next

two nerves.

(_c_) The _Nervus suralis_ (Figs. 128 8; 129 _s_) runs downwards on the

inner border of the _M. gastrocnemius_ and gives off:--

(1) A twig (_g_″) to the upper third of the muscle.

[Illustration: Fig. 129.

Nerves of the leg and sole of the foot.

_ab5_ Branch to the M. abduct. digit. V.

_ad5_ Branch to the M. adduct. digit. V.

_ar_ M. rectus anterior.

_b_ M. biceps.

_cs_ Ramus cutaneus medius.

_ct_ Ramus cutaneus cruris posterior.

_dvI_ Ramus digitalis volaris I.

_dvII_ Branch supplying R. digit. volaris I and II.

_dvIII_ Second branch of the N. tibialis.

_F5_ Branch to the M. flex. brev. digit. V.

_flp_ Branches to the M. plantaris and the M. flexor digitorum.

_g_′ Branch to the M. gastrocnemius.

_g_″ Branch to the M. gastrocnemius.

_pc_ N. peroneus.

_s_ N. suralis.

_sm_ M. semimembranosus.

_T_, _t_ Nervus tibialis.

]

(2) The _Ramus cutaneus medius_ (_cs_) is given off below the middle

of the _M. gastrocnemius_; it passes to the skin of the lower third

of the leg. After giving off the latter branch, the _N. suralis_

runs downwards along the inner aspect of the _Tendo achillis_ to the

_Aponeurosis plantaris_ of the sole of the foot as far as the calcar,

and gives off:--

(3) Branches (_flp_) to the _Musc. plantaris_ and the _M. flexor

digitorum_.

(4) A branch to the _M. abductor hallucis_.

(5) The _R. digitalis volaris I_ (_dvI_) supplies the inner side of the

first toe by its terminal twig.

(_d_) The _Nervus tibialis_ (Fig. 129 _t_) passes downwards on the

hinder surface of the tibio-fibula, sinks into the _M. tibialis

posticus_ to appear again at its hinder border; it then runs over the

ankle-joint into the sole of the foot, where it lies midway in the

space between the two _Ossa tarsi_ and between the _M. extensor tarsi_

and the _M. abductor digiti I longus_; the nerve then runs downwards in

the groove of the small _Os tarsi_ and divides to form three branches:--

(1) The first branch (_dvII_) runs to the space between the second and

third fingers, where it bifurcates:--

(α) The _Rami digital. volaris I_ run transversely over the muscles of

the second toe, supply the muscles of the first toe, and bifurcate to

form the _R. digital. volaris I_ and _II_.

(β) The _Ram. digit. volar. II_ divides in the space between the second

and third fingers to supply the _Flex. phalang._ and the adjacent sides

of these toes.

(2) The second branch (_dvIII_) runs over the _M. flex. metatarsi_

of the third toe to the space between the third and fourth toes, and

bifurcates to supply the adjacent sides of these toes and the web

between them.

(3) The third branch at once divides to supply:--

(α) The _M. transv. metatarsi_.

(β) The _MM. lumbricales_ of the fourth toe.

(γ) The _M. abductor digit. V_ (_ab5_), the _M. flexor brevis dig. V_

(_F5_), and the _M. adductor dig. V_ (_ad5_); it then ends as--

(δ) A bifurcating branch forming the _R. digit. volar._ of the fourth

and fifth toes.

*III.* The _Nervus peroneus_ (_N. peroneus communis superior_,

Ecker), (Fig. 130 _pc_) is the second division of the sciatic nerve;

it passes between the outer head of origin of the _M. gastrocnemius_

and the tendon of insertion of the _M. biceps_, it then lies on the

tibio-fibula between the _M. gastrocnemius_ and the _M. peroneus_,

where it is accompanied by the _Vena tibialis postica_, then runs

downwards upon the _M. extensor cruris_ and the _Flexor tarsi ant._,

and passes under the _M. tibial. anticus_ and bifurcates; its branches

are:--

*a.* The _N. cutaneus cruris lateralis_ (Fig. 128 _cp_); which, like

other cutaneous nerves, runs in a common sheath together with an artery

and vein to the skin. It arises close to the sciatic nerve.

*b.* Muscular branches to the _MM. peroneus_, _tibialis anticus_,

_extensor cruris brevis_, and the _flexor tarsi anterior_.

[Illustration: Fig. 130.

Distribution of the _N. peroneus_.

_A_ Inner terminal twig of N. peron. comm. inf.

_B_ Outer terminal twig of N. peron. comm. inf.

_cpl_ N. cutaneus dorsi pedis lateralis.

_pc_ N. peroneus.

_pci_ N. peroneus communis inferior.

_pl_ N. peroneus lateralis.

_pm_ N. peroneus medialis.

]

*c.* The _N. peroneus medialis_ (Fig. 130 _pm_) is the smaller of

the two terminal branches of the _N. peroneus_; it courses with the

_Art. tibialis antica_ under the _MM. flexores tarsi anterior_ and

_posterior_ and supplies the latter.

*d.* The _N. peroneus lateralis_ (Fig. 130 _pl_) is the larger terminal

branch of the _N. peroneus_; it passes downwards between the heads of

the _M. tibialis anticus_ to the tendon of origin of the _Flexor tarsi

posterior_, where it gives off two branches (Nos. 1 and 2); the nerve

then unites with the _N. peroneus lateralis_ to form a common stem, the

_N. peroneus communis inferior_ (Ecker). The _N. peroneus lateralis_

gives off:--

(1) The _N. cutaneus dorsi pedis lateralis_ (_cpl_), which passes to

the skin of the outer side of the dorsum of the foot.

(2) A second branch to the _M. extensor_ of the fourth and fifth toes.

*e.* The _N. peroneus communis inferior_ (Fig. 130 _pci_) runs on

the dorsum of the foot in company with the _Art. dorsalis pedis_,

underneath the _MM. extensores digiti I_ and _II_; it supplies several

branches and then bifurcates. It gives off:--

(1) Branches to the _MM. extensor longus_ and _brevis digiti I_.

(2) _Rami digitales dorsales_ to the adjacent sides of the first and

second toes.

(3) Branches to the _MM. extensores digiti II_.

(4) The inner, terminal branch (Fig. 130 _A_) at once divides:--

(α) The outer branch runs between the third and fourth toes as far as

the commencement of the web, where it bifurcates to form two _Rami

cutanei_, which course along the adjacent sides of these toes as far as

their apices.

(β) The inner branch passes to the extensor muscle, and, in part, to

the adjacent sides of the third and fourth toes.

(5) The outer, terminal branch (Fig. 130 _B_) of the _N. peroneus

communis inferior_ passes to the muscles of the fourth and fifth toes,

and supplies _Rami cutanei dorsales_ to the outer side of the fourth

and inner side of the fifth toes.

Cutaneous branches of the _N. tibialis_ supply the outer side of the

fifth and inner side of the first toe.

G. THE SYMPATHETIC SYSTEM (_Sympathicus_).

_(Re-written by the translator.)_

The *sympathetic cord* or *chain* is a row of nervous ganglia

(vertebral or lateral ganglia), connected by nerve-fibres, and lying

on either side of the vertebral column (Figs. 117 and 131); with the

exception of the last spinal nerve there is usually one sympathetic

ganglion associated with each spinal nerve; in the case of the tenth

spinal nerve there may be only one ganglion or as many as twelve.

The first ganglion (Figs. 111, opposite _ics_; 117 _S_1) is placed on

the hypoglossal nerve just as it emerges from the first intervertebral

foramen; it is large, but smaller than the second; its _Ramus

communicans_ is represented by several fine and very short fibres,

which connect the ganglion with the nerve. This ganglion is connected

with the second by two or three nervous threads, between which passes

the subclavian artery, a true _Annulus Vieussenii_ being thus formed

(de Watteville). The other ganglia are connected by single bands of

fibres. The first ganglion supplies also branches to the axillary

artery and to the cardiac plexus.

The second ganglion (Figs. 117 and 131) is the largest, and is closely

applied to the brachial nerve; as in the case of the first ganglion and

hypoglossal nerve, it is attached to the second spinal nerve without a

distinctly marked _Ramus communicans_.

The third ganglion (_Ganglion cardiacum basale_, Gaskell and Gadow)

is sometimes fused with the second, but is usually close to the third

spinal nerve: it has a short but distinct _Ramus communicans_.

[Illustration: Fig. 131.

Sympathetic cord. From Ecker (Icones physiologicae, Pl. XXIV, Fig. 3).

The heart, lungs, and liver have been removed; the stomach, intestine,

kidneys, and testes drawn to the right side; the left sympathetic cord

is thus pulled to the right side to expose the Rami communicantes.

_S_ Sympathetic cord attached to the ganglion of the vagus.

The numbers refer to the ganglia, which are enumerated from before

backwards.

]

Behind the third ganglion the sympathetic cord is continued backwards

along the corresponding aortic arch, then parallel with and close

to the abdominal aorta (Figs. 117, 127, and 131), receiving _Rami

communicantes_, which are long and well marked, from each of the spinal

nerves; the fourth, fifth, and sixth nerves usually supply each one

_Ramus communicans_, the seventh two, and the eighth and ninth each

two or three: from the tenth nerve it receives three or more, as many

as twelve having been noted. The ganglia are usually more or less

spindle-shaped or flattened and triangular; the hinder part of the cord

usually receives in addition two or three branches from the sciatic

plexus and twigs from the _R. abdominalis_ of the _N. coccygeus_.

The branches and communications of the sympathetic cords are as

follows:--

*a.* Communicating branches between the two cords; these are extremely

numerous and irregular, forming a net-like plexus, which surrounds the

abdominal aorta and other adjacent structures, and gives off numerous

small twigs to the neighbouring vessels and organs.

*b.* Communications with the cranial nerves; these are two branches

(Wiedersheim), (Figs. 111 and 116) which pass from the first ganglion

to the _Ganglion nervi vagi_, where one terminates, the other leaves

the ganglion to pass on to the Gasserian ganglion (Figs. 111 and 116

_VS_): according to Gaskell a _single_ nerve passes from the first

ganglion to the _Ganglion nervi vagi_, whence one portion of its fibres

is continued to the Gasserian ganglion, the remainder accompanying

the pneumogastric nerve without any connection with the ganglion;

he therefore names this nerve the vago-sympathetic. (See Gasserian

ganglion, p. 168.)

*c.* Communications with the spinal nerves; these are:--

(1) The _Rami communicantes_.

(2) Communications between the sympathetic ganglia or their branches

and the spinal nerves or their branches (Fig. 117); by means of these

connections, fibres of the sympathetic system are conducted by the

spinal nerves and their branches to all parts of the body.

*d.* Branches to the heart, which form the following ganglia:--

(1) A relatively large plexus lies on the auricles in the median plane

immediately beneath the division of the _M. hyoglossus_. It supplies a

network of fibres to the auricles and the adjacent large vessels. It is

said to communicate at various points with the pneumogastric nerve.

(2) A smaller ganglion of oval form, supplies twigs to the neighbouring

vessels and a communicating branch to the hypoglossal nerve

(Wiedersheim).

*e.* Branches to the abdominal viscera; these form intricate plexuses

by which the organs are supplied: the one best known is the solar

plexus (Fig. 131); it is formed chiefly from branches derived from the

third, fourth, and fifth ganglia, and supplies the stomach, etc.; other

plexuses for the various viscera are known by corresponding names, such

are the _Plexus hepaticus_, _renalis_, _genitalis_, _haemorrhoidalis_,

and _vesicalis_.

The sympathetic system is characterised by the fact that the branches

form intricate plexuses, which include numerous ganglia and which are

very irregular; it is also characteristic that most of its fibres are

non-medullated. According to the investigations of Gaskell, the fibres

of the sympathetic system arise in mammalia as very fine medullated

fibres from the posterior vesicular (Clarke’s) columns (Mason has

recently described cells in the frog’s spinal cord, which he holds to

be homologous with the cells of these columns); they leave the cord by

both the ventral and dorsal roots of the spinal nerves, and are thus

connected with two sets of ganglia, (1) with the ganglia of the dorsal

roots, and (2) through the _Rami communicantes_ with the sympathetic

ganglia (vertebral or lateral ganglia); these two sets of ganglia

Gaskell terms proximal. By means of the branches from the sympathetic

ganglia (_Rami efferentes_) part of the fibres pass to another set

of ganglia, the solar plexus, etc., which he terms prevertebral or

collateral; from these the fibres pass to be distributed to the various

viscera and blood-vessels, where a fourth set of very small ganglia

(terminal ganglia) is found. The prevertebral and terminal ganglia

are together classed as distal ganglia. Gaskell holds that the fine

medullated fibres from the cord lose their medullary sheath in one or

other of these ganglia according to the function they fulfil.

The inhibitory fibres of the heart and vaso-dilator fibres of the

blood-vessels continue as white fibres along the vago-sympathetic and

spinal nerves to the distal ganglia (Bidder’s ganglion, etc.), where

the medullary sheath disappears: whereas the ‘augmentor’ fibres of

the heart and vaso-constrictor fibres of the blood-vessels lose their

medullary sheath in the proximal ganglia and pass on as non-medullated

fibres. In the same way the nerve-fibres that bring about contraction

of the circular muscle fibres of the hollow viscera lose their

medullary sheaths in the proximal ganglia, while those fibres, the

influence of which negatives the former, become pale fibres in the

distal ganglia.

Waters has demonstrated that in the frog the various spinal nerves

have each a localised physiological action upon the blood-vessels and

muscular walls of various parts of the alimentary canal: he shows that

The third spinal nerve supplies the oesophagus.

The fourth spinal nerve supplies the stomach.

The fifth spinal nerve supplies the upper third of the small intestine.

The sixth spinal nerve supplies the lower two thirds of the small

intestine.

The seventh spinal nerve supplies the large intestine.

The eighth spinal nerve supplies the bladder, this supply being,

however, not so definite as the others given above.

It has long been known that the branches of the spinal ganglia (ganglia

of the posterior roots) are together larger (one-third, Lenhossék),

and contain more fibres than the ventral and dorsal roots together;

this is supposed to be in part due to an acquisition of new fibres

derived from the ganglia. The majority of recent observers hold that

each ganglionic cell has only one process, which, however, soon

bifurcates; whether any of the fibres so formed pass as far as the cord

or beyond its blood-vessels is doubted by most observers, and denied by

Gaskell. These remarks and the description of the cells of the spinal

ganglia (p. 176) hold good for the lateral or vertebral ganglia and the

prevertebral ganglia (solar ganglion, etc.) of the sympathetic system;

the terminal ganglia will be described with the organs in which they

are found.

H. HISTOLOGICAL NOTES ON THE NERVOUS SYSTEM.

(In order to render the foregoing description of the nervous

system more complete, the following notes have been added by the

translator.)

[The histological elements of the nervous system are nerve-cells

and nerve-fibres; of these the nerve-cells have been described with

the parts in which they occur; it may simply be remarked that later

observers have been unable to discover the ‘spiral cells’ described by

Beale, Arnold, and others. The nerve-fibres, as in most other animals,

are of two kinds, medullated and non-medullated.

*1.* Medullated nerve-fibres or white fibres are found in all cranial

and spinal nerves, with the exception of the olfactory nerves, and in

many of the sympathetic nerves (see Sympathetic System); also in the

white matter of the brain and spinal cord; examined microscopically

the fibres are seen to consist of an external sheath or neurilemma, a

medullary sheath, and an axis-cylinder:--

*a.* The neurilemma (Sheath of Schwann, Outer or Primitive Sheath) is a

nucleated endothelial layer covering the nerve-fibre; it is continuous

with the corresponding coat of the nerve-cells, and is uninterrupted

throughout the length of the nerve; at the nodes, however, it dips down

towards the axis-cylinder, the circular groove so formed being filled

with cement substance.

*b.* The medullary sheath (White substance of Schwann); the presence of

this sheath is the chief cause of the whiteness of these nerves; the

thickness of the sheath varies considerably, and towards the ultimate

distribution of the nerve it is entirely lost. At more or less regular

intervals along the course of the nerve-fibre the continuity of the

medullary sheath is broken, and gives the fibres the appearance of

being constricted at these places; such constrictions are known as

nodes of Ranvier; the portion between two such nodes being termed

an internode. Each internode possesses an oval, flattened, granular

nucleus at about its middle and placed between the neurilemma and the

medullary sheath; the nucleus has a nucleolus. In the fresh state the

medullary sheath seems to be fluid; it is of a fatty nature.

Medullary segments are caused by breaks in the continuity of the

medullary sheath, which are seen only in nerves which are no longer in

their normal condition, and are especially well marked after treatment

with osmic acid. The breaks are oblique; hence the conical end of one

segment fits into the funnel-shaped end of the next. How far they

correspond to pre-existent structures is uncertain. The segments in the

frog vary in length from 0.010 to 0.040 mm.

*c.* The axis-cylinder is the essential part of every nerve-fibre;

it shows a longitudinal striation corresponding to the fine fibrils

(primitive fibrillae) of which it consists; these fibrils often exhibit

minute varicosities: at times it has the appearance of being invested

with a very delicate structureless sheath. It is continuous through the

nodes of Ranvier.

*2.* Non-medullated nerve-fibres (Grey or Varicose Fibres; Fibres of

Remak); these occur chiefly in branches and plexuses of the sympathetic

system; they consist of a neurilemma and an axis-cylinder, which agree

exactly with the corresponding elements found in the medullated fibres.

The nerve-fibres, whether medullated or non-medullated, are bound

together by connective-tissue to form nerves. A number of fibres bound

together by connective-tissue to form a slender cord is known as a

funiculus; a small nerve may consist of one such funiculus; the sheath

surrounding it is known as the perineurium, and sends in supporting

processes between the fibres (endoneurium): when several funiculi are

bound together to form a large nerve the common sheath is known as

the epineurium. These sheaths support nerves (_nervi nervorum_) and

vessels (_vasa vasorum_) supplying the nervous elements, and their

intercellular spaces form lymph-canals. The whole nerve is surrounded

by an endothelial coat, which helps to form a lymph-space, which more

or less completely surrounds the nerve.

The ultimate distribution of the nerve-fibres will be included in the

description of the various organs in which they end.]

SECTION IV.

THE VASCULAR SYSTEM.

THE VASCULAR SYSTEM.

LITERATURE.

THE HEART AND BLOOD-VESSELS.

*Aeby*, Ueber den feineren Bau der Blutcapillaren. Centralbl. f. d.

med. Wiss. 1865, p. 209.

*Altmann, R.*, Ueber Corrosion in der Histologie. Centralbl. f. d.

med. Wiss. 1878, p. 245.

*Aubert, H.*, Die Innervation der Kreislaufsorgane. Hermann’s

Handbuch der Physiologie. Leipzig, 1880. Vol. VI, Pt. I, p. 377.

*Beck, K.*, Zur Kenntniss der Herznerven. Arch. f. mik. Anat. 1884.

Vol. XXIV, pp. 11–19.

*v. Bezold*, Untersuchungen über die Innervation des Herzens.

Leipzig, 1863.

*Bidder, F.*, Ueber functionell verschiedene und räumlich getrennte

Nervencentra im Froschherzen. Arch. f. Anat. u. Physiol. 1852,

p. 163.

*Bidder, F.*, Zur näheren Kenntniss des Froschherzens und seiner

Nerven. Arch. f. Anat. u. Physiol. 1866, p. 1.

*Bidder, F.*, Endigungsweise der Herzzweige des N. vagus beim

Frosche. Arch. f. Anat. u. Physiol. 1868, p. 1.

*Bidder, F.*, and *Gregory*, Beiträge zur Physiologie der

Herzbewegung beim Frosche. Dorpat, 1865.

*Boas, J. E. V.*, Ueber den Conus arteriosus und die Arterienbogen

der Amphibien. Morph. Jahrb. 1881. Vol. VII, pp. 271–273.

*Boas, J. E. V.*, Beiträge zur Angiologie der Amphibien. Morph.

Jahrb. 1882. Vol. VIII, pp. 169–187.

*Bobretzky, C.*, Ueber die Entwickelung der Capillargefässe.

Centralbl. f. d. med. Wiss. 1885, p. 769.

*Bremer, L.*, Die Nerven der Capillaren der kleinen Arterien und

Venen. Arch. f. mik. Anat. 1882. Vol. XXI, p. 663.

*Brenner, A.*, Ueber das Verhältniss des N. laryngeus inferior vagi

zu einigen Aortenvarietäten des Menschen und zu dem Aortensystem

der durch Lungen athmenden Wirbelthiere überhaupt. Arch. f. Anat.

u. Physiol. 1883, pp. 373–397.

*Brücke, E.*, Ueber Mechanik des Kreislaufes des Blutes bei Fröschen.

Wiener Sitzungsb. 1851. Vol. VI, p. 61; also p. 114.

*Brücke, E.*, Beiträge zur vergl. Anatomie u. Physiologie des

Gefässsystems der Amphibien. Denkschriften d. Wiener Academie.

1852. Vol. III, p. 335.

*Burdon-Sanderson, J.*, Circulation of the Blood, in Handb. for the

Physiological Laboratory. London, 1873.

*Burow*, De vasis sanguiferis ranarum. Diss. inaug. Regiomontani,

1834.

*Calori, L.*, Sugli organi della circulatione e della respirazione

dei gyrini della Rana esculenta e della Salamandra cristata. Nuov.

Ann. delle Scienz. nat. de Bologna. 1838.

*Champness, F.*, The Septum Atriorum of the Frog and the Rabbit.

Journ. of Anat. and Physiol. 1874, p. 340.

*Chapmann, S. H.*, Beiträge zur Kenntniss des Baues des normalen und

entzündeten Pericardiums der Batrachier. Wiener Med. Jahrbücher.

*Chrzonzczewsky, N.*, Ueber die feinere Structur der Blutcapillaren.

Virchow’s Arch. 1866. Vol. XXXV, p. 169.

*Darwin, F.*, Contributions to the anatomy of the sympathetic ganglia

of the Bladder in their relation to the vascular system. Quart.

Journ. Micros. Sci. 1874. Vol. XIV, p. 109.

*v. Deen*, De differentia et nexu inter nervos vitae animalis et

vitae organicae. Diss. inaugur. Lugduni; also in Arch. f. Anat. u.

Physiol. 1834, p. 477.

*v. Deen*, Over de zijelingsche Takken dar zwervende van den Proteus

anguinis (includes frog-larva). Bijdragen tot de Naturkundige

Wetenschappen. 1834.

*Dogiel, J.*, Die Ganglienzellen des Herzens bei verschiedenen

Thieren und beim Menschen. Arch. f. mik. Anat. 1877. Vol. XIV,

p. 471.

*Dogiel, J.*, Die Nervenzellen und Nerven des Herzventrikels beim

Frosche. Arch. f. mik. Anat. 1882. Vol. XXI, p. 21.

*Dumeril, A. M. C.*, and *Bibron*, *G.*, Erpétologie générale, ou

histoire naturelle complète des reptiles. Paris, 1841.

*Eberth, C. J.*, Ueber den feineren Bau der Blutcapillaren bei den

Wirbelthieren. Centralbl. f. d. med. Wiss. 1865, p. 196.

*Eberth*, Elemente der quergestreiften Muskelfasern, bes. des

Herzens. Virchow’s Arch. 1866. Vol. XXXVII, p. 100.

*Ecker, A.*, Icones physiologicae. Leipzig, 1851–9.

*Engelmann, T. W.*, Der Bulbus Aortae des Froschherzens. Physiol.

Untersuch. in Gemeinsch. mit. J. Hartog und J. J. Verhoff.

Pflüger’s Arch. f. d. ges. Physiol. 1882. Vol. XXIX, pp. 425–468.

*Friedländer*, Ueber die nervösen Centralorgane des Froschherzens.

Unters. a. d. physiol. Labor. in Würzburg. 1867. Vol. II, p. 159.

*Fritsch, G.*, Zur vergleichenden Anatomie der Amphibienherzen. Arch.

f. Anat. u. Physiol. 1869, p. 654.

*Gaskell, W. H.*, On the augmentor (accelerator) nerves of the heart

of cold-blooded animals. Journ. of Physiol. 1884. Vol. V, p. 46.

*Gaskell, W. H.*, and *Gadow*, On the anatomy of the cardiac nerves

in certain cold-blooded animals. Journ. of Physiol. 1884. Vol. V,

p. 362.

*Gastaldi*, Neue Untersuchungen über die Muskulatur des Herzens.

Würzb. Naturf. Zeitschr. 1862. Vol. III, pp. 6–9.

*Gegenbaur, C.*, Grundzüge der vergleichende Anatomie. 2nd edit.,

1870.

*Gerlach, L.*, Ueber die Nervenendigungen in der Muskulatur des

Froschherzens. Virchow’s Arch. 1876. Vol. LXVI, pp. 187–223.

*Golubew, A.*, Beiträge zur Kenntniss des Baues and der

Entwicklungsgeschichte der Capillargefässe des Frosches. Arch. f.

mik. Anat. 1869. Vol. V, pp. 49–89.

*Gompertz, C.*, Ueber Herz und Blutkreislauf bei nackten Amphibien.

Arch. f. Anat. u. Physiol. Phys. Abt. 1884, p. 242.

*Gruby*, Sur le système veineux de la grenouille. Annales des

Sciences nat. 2nd series. Zool. 1842. Vol. XXVII, p. 207.

*Gruby*, Recherches anatomiques sur le système veineux de la

grenouille. Paris, 1842.

*Gscheidlen, R.*, Zur Lehre v. d. Nervenendigungen in den glatten

Muskelfasern. Arch. f. mik. Anat. 1877. Vol. XIV, p. 321.

*Heidenhain*, Disquisitiones de nervis organisque centralibus cordis.

Dissert. inaug. Berol., 1854; and in Arch. f. Anat. u. Physiol.

1858, p. 479.

*His, W.*, Ueber die Endigung der Gefässnerven (in mesentery).

Virchow’s Arch. 1863. Vol. XXVIII, pp. 427–428.

*Hoffmann, C. K.*, Die Lungengefässe der Rana temporaria. Dissert.

Dorpat, 1875.

*Hoffmann, C. K.*, in Bronn’s Klassen und Ordnungen des Thierreichs.

Leipzig und Heidelberg, 1873–1878. Vol. VI, pp. 509–514.

*Hoffmann, T.*, Die Lungen-Lymphgefässe der R. temporaria. Dissert.

Dorpat, 1875.

*Huizinga, D.*, Untersuchungen über die Innervation der Gefässe in

der Schwimmhaut des Frosches. Pflüger’s Arch. f. d. ges. Physiol.

1875. Vol. XI, p. 207.

*Huschke*, Ueber die Carotidendrüse einiger Amphibien. Zeit. f.

Physiol. von Friedmann u. Treviranus. 1831. Vol. IV, p. 113.

*Hyrtl, J.*, Vorläufige Anzeige über gefässlose Herzen. Wiener

Sitzungsb. 1859. Vol. XXXIII, p. 572.

*Hyrtl, J.*, Ueber die sogenannten Herzvenen der Batrachier. Wiener

Sitzungsb. 1864. Vol. L, Pt. II, p. 42.

*Hyrtl, J.*, Ueber das Verhalten der Leberarterie zur Pfortader bei

Amphibien u. Fischen. Wiener Sitzungsb. 1864. Vol. XLIX, Pt. I,

p. 167.

*Jantschitz, J.*, Materialien zur Anatomie der Nerven des

Pericardiums. Rudneff’s Journ. f. normale und patholog. Histologie

u. klinisch. Med. St. Petersburg, 1874, pp. 417–442.

*Klug, F.*, Ueber die Beschleunigungsnerven des Froschherzens.

Centralbl. f. d. med. Wiss. 1881, p. 945.

*Klug, F.*, Ueber die Herznerven des Frosches. Arch. f. Anat. u.

Entwicklungsgesch. 1881, pp. 330–346. Abstract in Centralbl. f. d.

med. Wiss. 1882, p. 259.

*v. Kölliker, A.*, Handbuch der Gewebelehre. 5th edit.

*Küttner*, Beitrag zu den Kreislaufsverhältnissen in der Froschlunge.

Virchow’s Arch. Vol. LXI, p. 21.

*Lahousse, P.*, Die Structur des Nervenplexus in der

Vorhofscheidewand des Froschherzens. Arch. f. Anat. u. Physiol.

1886, pp. 191–196.

*Lambotte, H. A.*, Observations anatomiques et physiologiques sur les

appareils sanguins et respiratoires de Batraciens anourés. Mém.

cour l’Acad. de Bruxelles. 1838.

*Langer, C.*, Ueber das Lymphgefasssystem des Frosches. Wiener

Sitzungsb. 1866. Vol. LIII, Pt. I, p. 395; 1867. Vol. LV, Pt. I,

p. 593; 1868. Vol. LVIII, Pt. I, p. 198.

*Langerhans, P.*, Notiz zur Anatomie des Amphibienherzens. Zeit. f.

wiss. Zool. 1873. Vol. XXIII, p. 457.

*Langerhans, P.*, Zur Histologie des Herzens. Virchow’s Arch. 1873.

Vol. LVIII, pp. 65–83.

*Lavdowsky, M.*, Das Säugadersystem und die Nerven der Cornea. Arch.

f. mik. Anat. 1872. Vol. VIII, p. 538.

*Lavdowsky, N.*, Ueber die Fortsätze der Nervenzellen in den

Herzganglien. Arch. f. mik. Anat. 1887. Vol. XXIX, p. 609.

*Lehmann*, Ueber die Nervenendigungen und das Vorkommen von

mikroskopischen Ganglien in den Gefässwandungen. Zeit. f. wiss.

Zool. 1864. Vol. XIV, pp. 346–352.

*Levschin*, Ueber das Lymph- und Blutgefässsystem. Wiener Sitzungsb.

1870. Vol. LXI, Pt. I, p. 67.

*Leydig*, Anatomisch-histologische Untersuchungen über Fische und

Amphibien. 1853.

*Leydig, F.*, Ueber Organe eines sechsten Sinnes. Nova Acta Acad.

Caes. 1868. Vol. XXXIV, pp. 1–102.

*Leydig*, Lehrbuch der Histologie. 1857.

*Löwit, M.*, Beiträge zur Kenntniss der Innervation des Herzens.

Pflüger’s Arch. f. d. ges. Physiol. Chapters I and II, 1880,

Vol. XXIII, p. 313. Chapters III, IV, and V, 1881, Vol. XXV,

pp. 399–496. Chapter VI, 1882, Vol. XXVIII, p. 312. Chapter VII,

1882, Vol. XXIX, p. 469.

*Löwit, M.*, Ueber die Gegenwart von Ganglienzellen im Bulbus aortae

des Froschherzens. Pflüger’s Arch. f. d. ges. Physiol. 1883.

Vol. XXXI, pp. 88–94.

*Ludwig, C.*, Ueber die Herznerven des Frosches. Arch. f. Anat. u.

Physiol. 1848, p. 139.

*Marshall, A. M.*, The Frog. Manchester and London. 1885, pp. 22–31.

*Marshall, C. F.*, Structure and Distribution of striped and

unstriped muscle. Quart. Journ. Micros. Sci. 1887. Vol. XXV.

*Meckel, J. F.*, System der vergleichenden Anatomic. Halle, 1833.

*Milne-Edwards*, Leçons sur la Physiol. et l’Anat. comparée. Vol. II,

p. 407.

*Namias, M.*, Sui ganglii miocardici della Rana. Osservazione ed

esperienzi in Lo Spallanzani. 1881. Vol. X, pp. 402–419.

*Nicolsky, P.*, Ueber Flimmerendothel beim Frosche (in pericardium).

Centralbl. f. d. med. Wiss. 1880, p. 641.

*Oellacher, J.*, Ueber die erste Entwickelung des Herzens und der

Pericardial- oder Herzhöhle bei Bufo cinereus. Arch. f. mik. Anat.

1871. Vol. VII, p. 157.

*v. Openchowski, T.*, Beitrag zur Kenntniss der Nervenendigungen im

Herzen. Arch. f. mik. Anat. 1883. Vol. XXII, p. 408.

*Owen, R.*, Comparative Anatomy. Vol. I, p. 517.

*Owen, R.*, On the structure of the heart in the perennibranchiate

Batrachia. Trans. Zool. Society, 1834. Vol. I, p. 212.

*Pihlermann, R.*, Untersuchungen über die angeblich präformirten

Verbindungswege zwischen den Blut- und Lymphgefässen des Frosches.

Dissert. Dorpat, 1876.

*Pohl-Pincus*, Ueber die Muskelfasern des Froschherzens. Arch. f.

mik. Anat. 1884. Vol. XXIII, p. 500.

*Prévost* and *Lebert*, Mémoire sur la formation des organes de la

circulation et du sang dans le batraciens. Annales des Sciences

nat. 1844; Vol. I, p 193; and in Compt. rend. 1844; Vol. XVIII,

p. 88; and in Froriep’s Notizen, 1844, Vol. XXX, cols. 337–340.

*Ranvier, L.*, Leçons d’anatomie générale. Paris, 1880, p. 469.

*Ranvier, L.*, Appareils nerveux terminaux des muscles de la vie

organique; cœrs sanguins, cœrs lymphatiques; œsophagus; muscles

lisses. Leçons d’anatomie générale au College de France. Paris,

1880. Vol. VII, p. 350.

*Rawitz, B.*, Ueber den Bau der Spinalganglien. Arch. f. mik. Anat.

Vol. XXI, p. 244.

*Reich, M.*, Einige microscopische Studien mit Silbersalpeterlösung

besonders an Gefässen des Auges und anderen Organen. Wiener

Sitzungsb. 1873. Vol. LXVII, Pt. III, pp. 81–96.

*Rokitansky, C.*, Ueber die Scheidewand der Vorhöfe. Wiener Med.

Jahrb. 1871, p. 340.

*Rosenberger*, De centris motuum cordis disquisitiones

anatomico-physiologicae. Dorpat, 1850.

*Rouget, C.*, Mémoire sur le développement, la structure et

les proprietés physiologigues des capillaires sanguins et

lymphatiques. Arch. de physiol. normale et pathologique. 1873,

pp. 662–664.

*Rusconi, M.*, Observations sur le système veineux de la grenouille.

Annales des Sciences nat. 2nd series. Zool. 1845. Vol. IV, p. 283.

*Sabatier*, Etudes sur le cœur dans la série des vertébrés.

Montpellier, 1873; also Ann. Sci. nat. 1873, Vol. XVIII, art. 4;

also Revue Cours. Scient. 1873, pp. 163–165.

*Schmuziger, F.*, Ein Beitrag zur Auswanderung der Blutkörperchen

aus den Gefässen des Frosches. Arch. f. mik. Anat. 1873. Vol. IX,

pp. 709–711.

*Schöbl, D. J.*, Ueber die Blutgefässe des cerebrospinalen

Nervensystems der Urodelen. Arch. f. mik. Anat. 1882. Vol. XX,

p. 87. Also in Sitzungsb. d. k. b. Gesell. d. Wiss. in Prag. 1878,

p. 25.

*Schöbl, D. J.*, Ueber Wundernetze und divertikelbildende Capillaren

bei nackten Amphibien, etc. Arch. f. mik. Anat. 1885. Vol. XXV,

p. 89.

*Schraeder, M. E. G.*, Ueber das Hemmungscentrum des Froschherzens.

Strassburg, 1886.

*Schutz, C. H.*, Das System der Circulation. Stuttgart and Tübingen,

1836.

*Schweigger-Seidel*, Das Herz, in Stricker’s Handbuch d. Gewebelehre.

*Sokoloff*, Ueber die Saftkanälchen (Lymphkanälchen) in den

quergestreiften Muskeln des Frosches. Rudneff’s Journ. f. normal

u. patholog. Histologie und Klin. Med. St. Petersburg, 1873,

pp. 456–459.

*Stirling, W.*, and *Macdonald*, The minute structure of the palatine

nerves of the Frog and the termination of nerves in blood-vessels

and glands. Journ. of Anat. and Physiol. 1884. Vol. XVII, p. 273.

*Stricker, S.*, Untersuchungen über die capillaren Blutgefässe in der

Nickhaut des Frosches. Wiener Sitzungsb. 1865. Vol. LI, Pt. II,

pp. 16–26.

*Stricker, S.*, Studien über den Bau und das Leben der capillaren

Blutgefässe. Wiener Sitzungsb. 1866. Vol. LII, Pt. II, pp. 379–394.

*Treviranus, G. R.*, Ueber die Karotidendrüse einiger Amphibien.

Tiedemann’s Zeitschr. 1831. Vol. IV, p. 113.

*Treviranus, C. R.*, Beobachtungen aus der Zootomie u. Physiologie,

nach dessen Tode herausgegeben von L. C. Treviranus. Bremen, 1839.

*Verhoeff, J. J. W.*, Histologische en physiologische bijdragen

tot de kennis van den bulbus aortae van het kikvorschhart.

Onderzoekingen gedach in het Phys. Laborat. VII, aflev 2. Utrecht,

1882.

*Vignal*, Recherches sur l’appareil ganglion nerve du cœur des

vertébrés. Laboratoire d’histologie du Collége de France. Travaux

de l’année 1881, p. 186; and in Arch. de Physiol. norm. et path.

1881, pp. 673–694 and 910–934.

*Virchow, H.*, Ueber die Kopfgefässe des Frosches. Sitzungsb.

Würzburg, 1880. Vol. XV, p. xxxiv.

*Virchow, H.*, Ueber die Gefässe im Auge und die Umgebung des Auges

beim Frosche. Zeit. f. wiss. Zool. 1881. Vol. XXXV, p. 247.

*Volkmann*, Nachweise der Nervencentra, von welchen die Bewegung der

Lymph- und Blutgefässherzen ausgeht. Arch. f. Anat. u. Physiol.

1844, p. 419.

*Wagner, R.*, Lehrbuch d. vergleichenden Anatomie. Leipzig, 1834–35.

*Wagner, R.*, Icones Zootomicae. Leipzig, 1841.

*Weismann*, Ueber die Muskulatur des Herzens beim Menschen und in dem

Thierreiche. Arch. f. Anat. u. Physiol. 1861, p. 42.

*Wyman, J.*, On the heart and respiration of the Menobranchus and

Batrachians. Proc. of the Boston Soc. of Nat. History. 1856.

Vol. V, p. 51.

*Zimmermann, W.*, Ueber circumvasale Safträume der Glaskörpergefässe

von Rana esculenta. Arch. f. mik. Anat. 1886. Vol. XXVII, p. 410.

THE LYMPHATIC SYSTEM.

*Bacculo, B.*, Nuove ricerche intorno l’apparecchio ganglionare

intrinsico dei cuori linfatici. Naples, 1885.

*Eckhard, C.*, Zeitsch. f. rat. Med. 1850. Vol. IX.

*Eckhard, C.*, Beiträge zur Anat. u. Physiol. 1855, Vol. I, p. 53;

1858, Vol. II, p. 145; 1863, Vol. III, p. 167.

*Eckhard, C.*, Experimentale Physiologie des Nervensystems. Giessen,

1867, p. 208.

*Goltz*, Centralbl. f. med. Wiss. 1863, pp. 17, 497; 1864, p. 690.

*Heidenhain*, Disquisitiones de nervis cordis cordiumque

lymphaticorum. Berlin, 1854.

*His, W.*, Ueber die Wurzeln der Lymphgefässe in den Häuten des

Körpers und über die Theorien der Lymphbildung. Zeit. f. wiss.

Zool. 1863. Vol. XII, p. 223.

*Hoffmann, T.*, Die Lungen-Lymphgefässe der R. temporaria. Dissert.

Dorpat, 1875.

*Hüter, C.*, Ueber den Kreislauf und die Kreislaufsstörungen in der

Froschlunge. Centralb. f. med. Wiss. 1873, Nos. 5 and 6.

*Jourdain, S.*, Sur les sacs sous-cutanés et les sinus lymphatiques

de la région céphalique dans la Rana temporaria. Compt. rend.

1881. Vol. XCIII, pp. 597–600.

*Jourdain, S.*, Sur le système lymphatique des tétards de

grenouilles. Compt. rend. 1883. Vol. XCVI, pp. 271–273.

*Jourdain, S.*, Recherches sur le système lymphatique de la R.

temporaria. Montpellier, 1883; Extr. de la Revue de Sci. nat.

Montpellier, 1884.

*Kahrhel, G.*, Studien über Innervation der Lymphherzen. Medic.

Jahrb. 1886, pp. 392–420.

*Key* and *Retzius*, Studien in der Anatomie des Nervensystems. Arch.

f. mik. Anat. 1873. Vol. IX, p. 308.

*Klein, E.*, On the lymphatic system of the skin and mucous

membranes. Quart. Journ. Micros. Sci. 1881. Vol. XX, pp. 379–406.

*Langendorff, O.*, and *Boll, F.*, Zur Kenntniss der Lymphherzen.

Arch. f. Anat. u. Physiol. 1883, p. 329.

*Langer, C.*, Ueber das Lymphgefässsystem des Frosches. Wiener

Sitzungsb. 1866, Vol. LIII, Pt. I, p. 395; 1867, Vol. LV, Pt. I,

p. 593; 1868, Vol. LVIII, Pt. I, p. 198.

*Levschin*, Ueber das Lymph- und Blutgefässsystem. Wiener Sitzungsb.

1870. Vol. LXI, Pt. I, p. 67.

*Luchsinger, B.*, Zur Lehre der Innervation der Lymphherzen.

Pflüger’s Arch. f. d. ges. Physiol. 1880. Vol. XXIII, p. 304.

*Meyer, J.*, Systema amphibiorum lymphaticum. Diss. Berol., 1845.

*Müller, J.*, Beiträge zur Anatomie und Naturgeschichte der

Amphibien. Tiedemann’s Zeitschr. 1831. Vol. IV, pp. 190–275.

*Müller, J.*, On the existence of four distinct hearts having

regular pulsations connected with the lymphatic system in certain

amphibious animals. Phil. Trans. 1833. Pt. I, pp. 89; also in

Arch. f. Anat. u. Physiol. 1834, p. 296.

*Müller, J.*, Bemerkungen über die Structur der Lymphherzen und der

Lymphgefässe. Arch. f. Anat. u. Physiol. 1839, p. 176.

*Obersteiner, H.*, Ueber einige Lymphräume im Gehirne. Wiener

Sitzungsb. 1870. Vol. LXI, Pt. I, p. 57.

*Pagliani*, Ueber die Funktion der Herzganglien. Moleschott’s

Untersuchungen. 1876. Vol. XI, p. 358.

*Panizza, B.*, Sopra il sistema linfatico dei retelli. Ricerche

Zootomiche. Pavia, 1833.

*Panizza, B.*, Ueber die Lymphherzen der Amphibien. Arch. f. Anat. u.

Physiol. 1834, p. 300.

*Pihlermann, R.*, Untersuchungen über die angeblich präformirten

Verbindungswege zwischen den Blut- und Lymphgefässen des Frosches.

Dissert. Dorpat, 1876.

*Priestly, J.*, An account of the anatomy and physiology of the

batrachian lymph-hearts. Journ. of Physiol. 1878. Vol. I, p. 1.

*Ranvier, L.*, Appareils nerveux terminaux des muscles de la vie

organique; cœurs sanguins, cœurs lymphatiques; œsophagus; muscles

lisses. Leçons d’Anatomie générale faites au Collége de France.

Paris, 1880. Vol. VII, p. 350.

*v. Recklinghausen*, Die Lymphgefässe und ihre Beziehung zum

Bindegewebe. 1862.

*Remak, R.*, Ueber blutlehre Gefässe (Lymphgefässe) im Schwanze der

Froschlarve. Arch. f. Anat. u. Physiol. 1849, p. 102.

*Robinson, C.*, Ueber die Lymphgefässe der Abdominaleingeweide der

Frösche, sowie deren Lymphbehälter. Froriep’s Notizen, 1846. No.

807, col. 225.

*Rusconi, M.*, Lettre du docteur Rusconi à M. Breschet sur le système

lymphatique de la Salamandra commune. Arch. f. Anat. u. Physiol.

1840, p. 8.

*Rusconi, M.*, Ueber die Lymphgefässe der Amphibien. Arch. f. Anat.

u. Physiol. 1843, p. 241.

*Rusconi, M.*, Einige historische Notizen die Lymphgefässe der

Amphibien betreffend. Arch. f. Anat. u. Physiol. 1843, p. 244.

*Rusconi, M.*, Sur les vaisseaux lymphatiques. Annales des Sci. nat.

Series II. Zool. 1841. Vol. XV, p. 249.

*Rusconi, M.*, Riflessioni sopra il sistema linfatico dei Rettili.

Pavia, 1845.

*Scherkey, M. L.*, Ueber die Feststellung und Bedeutung der Centren

der Lymphherzen im Rückenmark. Dissert. Berlin, 1878.

*Scherkey, M. L.*, Zur Lehre der Innervation der Lymphherzen. Arch.

f. Anat. u. Physiol. 1879, p. 227.

*Schweigger-Seidel, F.*, and *Dogiel, J.*, Ueber die Peritonealhöhle

bei Fröschen und ihren Zusammenhang mit dem Lymphgefässsysteme.

Arbeiten aus d. physiol. Anstalt. Leipzig, 1867, pp. 68–76;

Abstract in Centralbl. f. d. med. Wiss. 1867, p. 287.

*Suslowa, N.*, Beiträge zur Physiologie der Lymphherzen. Centralbl.

f. d. med. Wiss. 1867, p. 832.

*Suslowa*, Beiträge zur Physiol. der Lymphherzen. Dissert. Zürich,

1867; also in Zeitsch. f. rat. Med. 3rd series. 1868. Vol. XXXI,

p. 224.

*Waldeyer, W.*, Anatomische und physiologische Untersuchungen über

die Lymphherzen der Frösche. Zeitsch. f. rat. Med. 3rd series.

1864. Vol. XXI, pp. 103–124; Abstract in Centralbl. f. d. med.

Wiss. 1864, p. 73.

*Waldeyer, W.*, Zur Anatomie und Physiologie der Lymphherzen von Rana

und Emys europea. Studien des physiol. Instituts zu Breslau. 1865,

pp. 71–96; Abstract in Centralbl. f. d. med. Wiss. 1865, p. 321.

*Weber, E. H.*, Ueber die Lymphherzen der Amphibien von Panizza.

Arch. f. Anat. u. Physiol. 1834, pp. 300–304.

*Weber, J.*, Beiträge zur Anatomie und Physiologie. Bonn, 1832.

THE BLOOD AND LYMPH.

*Aly, W.*, Ueber die Vermehrung der rothen Blutkörperchen bei

Amphibien. Dissert. Halle, 1884.

*Arndt, R.*, Beobachtungen an roten Blutkörperchen der Wirbelthiere.

Virchow’s Arch. Vol. LXXVIII, p. 1.

*Beale, L. S.*, Observations upon the nature of the red

blood-corpuscle. Quart. Journ. Micros. Sci. 1864, pp. 32–43.

*Bizzozero, G.*, Ueber die Teilung der roten Blutkörperchen.

Centralbl. f. d. med. Wiss. 1881, p. 129.

*Bizzozero, G.*, Ueber die Teilung der roten Blutkörperchen.

Centralbl. f. d. med. Wiss. 1882, p. 577.

*Bizzozero, G.*, and *Torre, A. A.*, Ueber die Entstehung der roten

Blutkörperchen bei den verschiedenen Wirbelthierklassen. Virchow’s

Arch. 1884. Vol. XCV, pp. 1–25.

*Brücke, E.*, Ueber den Bau der rothen Blutkörperchen. Wiener

Sitzungsb. 1867. Vol. LVI, Pt. II, pp. 79–91.

*Donders* and *Moleschott*, Holländische Beiträge. 1848, p. 361.

*Eberth, C. J.*, Ueber die Vermehrung der rothen Blutkörperchen nach

Untersuchungen von W. Aly. Fortschritte d. Med. 1885. Vol. III,

pp. 1–7.

*Fuchs, E.*, Beitrag zur Kenntniss des Froschblutes und der

Froschlymphe. Virchow’s Arch. 1877. Vol. LXXI, p. 78.

*Gaule, J.*, Beobachtungen der farblosen Elemente des Froschblutes.

Arch. f. Anat. u. Physiol. 1880, pp. 375–392.

*Gulliver, G.*, Measurements of the red blood-corpuscles of

Batrachians. Proc. Zool. Soc. London, 1873, p. 162.

*Hewson, W.*, On the figure and composition of the red Particles

of the Blood, commonly called Globules. Phil. Trans. 1773.

Vol. LXIII, Pt. I, p. 303.

*Home, E.*, On the changes the blood undergoes in the act of

coagulation. Phil. Trans. 1818, p. 172.

*Kollmann, J.*, Bau der rothen Blutkörperchen. Zeitschr. f. wiss.

Zool. 1874. Vol. XXIII, p. 462.

*Kusnezoff, F.*, Ueber blutkörperhaltige Zellen der Milz. Wiener

Sitzungsb. 1873. Vol. LXVII, Pt. III, pp. 58–67.

*Moriggia, A.*, Ueber den Durchtritt der farblosen Blutkörperchen

durch die Blutgefässwandungen, etc. Moleschott’s Untersuchungen.

1876. Vol. XI, p. 470.

*Neumann, E.*, Zur Histologie der rothen Blutkörperchen. Centralbl.

f. d. med. Wiss. 1865, p. 481.

*Owsjannikow, P.*, Zur Histologie der Blutkörperchen. Bullet. de

l’acad. science de Pétersbourg. 1865. Vol. VIII, pp. 561–572.

*Peremeschko*, Ueber die Theilung der rothen Blutkörperchen bei

Amphibien. Centralbl. f. d. med. Wiss. 1879, pp. 673–675.

*Ranvier, L.*, Traité technique d’histologie, pp. 148–224.

*Rollett, A.*, Versuche und Beobachtungen am Blute. Wiener Sitzungsb.

Vol. XLVI, Pt. II, pp. 65–98.

*Stricker*, Beobachtungen über die Entstehung des Zellkernes. Wiener

Sitzungsb. 1878. Vol. LXXVI, Pt. III, p. 7.

*Thoma, R.*, Die Ueberwanderung farbloser Blutkörperchen von dem Blut

in das Lymphgefässsystem. Heidelberg, 1873.

*Welcker, H.*, Grösse, Zahl, Volum, Oberfläche und Farbe der

Blutkörperchen bei Menschen und Thieren. Henle u. Pfeuffer’s

Zeitschr. 1864. Vol. XX, pp. 258–307.

PART 1.

THE HEART AND BLOOD-VESSELS.

I. THE HEART.

(The original arrangement of this section has been modified.)

The heart is situated in the middle line above the central portion of

the shoulder-girdle and _M. sternohyoideus_, and below the oesophagus;

the apex is directed backwards and lies between the lobes of the liver,

the base is directed forwards and lies a short distance behind the

larynx. The heart is surrounded by the pericardium.

A. The *Pericardium* [is a very thin membranous sac, which completely

encloses the heart; it is attached, at some distance, to the large

vessels passing to and from the heart, and has inserted into it some

fibres of the _M. obliquus abdominis internus_ (see page 70). A fold

of the pericardium extends from the dorsal surface to the truncus

arteriosus, and through it courses the vena cardiaca.

[Illustration: Fig. 132.

The heart and blood-vessels, seen from the ventral surface; after Howes.

_AC._ Left anterior caval vein.

_C′._ Carotid arch.

_LA._ Left auricle.

_P′._ Right pulmo-cutaneous arch.

_RA._ Right auricle.

_S′._ Left systemic arch.

_TA._ Truncus arteriosus.

_V._ Ventricle.

]

The pericardium is a connective-tissue membrane, and is lined on

either side with endothelium, which is in part ciliated (Leydig); the

endothelium of the inner surface is continued on to the heart and so

forms a closed lymph-sac. The pericardium is more or less pigmented,

and contains both vessels and nerves; it may contain fat-cells].

B. The *Heart*.

[The heart is a hollow muscular organ, composed of: (1) two auricles,

forming the wider anterior portion; (2) the ventricle, placed

behind the auricles; (3) the sinus venosus, situated dorsally; and

(4) the truncus arteriosus, lying ventrally. Examined while still

living, the auricles are seen to be much darker in colour than the

ventricle, due to the blood being seen through their thinner walls;

between the auricles and the ventricle is a distinct groove, the

auriculo-ventricular groove. The whole organ is more or less pigmented,

especially the ventricle (_R. temporaria_ has much less pigment).]

[Illustration: Fig. 133.

The heart, seen from above, and dissected to show the opening from the

Sinus venosus to the right auricle; after Howes.

_C′._ Right carotid arch.

_IC._ Posterior caval vein.

_LA._ Left auricle.

_P′._ Right pulmo-cutaneous arch.

_PV._ Pulmonary vein.

_RA._ Right auricle.

_S′._ Right systemic arch.

_SV._ Sinus venosus.

_SV′._ Opening of Sinus venosus into the right auricle.

_V._ Right anterior caval vein.

]

*a.* The *Sinus venosus* (_Sinus venarum cavarum_), (Fig. 133 _SV._)

is a thin-walled sac lying on the dorsal surface of the heart, and

very slightly to the right side (Hoffmann). It is somewhat triangular

in form, receiving an anterior caval vein at each anterior angle

(_V._), and the posterior caval vein (_IC._) at the posterior angle.

The ventral wall has a transverse opening (_SV′._), by which it

communicates with the right auricle; the opening possesses two valves,

an anterior and a posterior, and is placed close to the auricular

septum.

*b.* The *auricles* [are not always separated, as the septum which

usually brings about the separation is in some cases incomplete, and in

rare instances appears to be absent (Hoffmann)]; according to Ecker,

the auricular septum is sometimes placed so far to the left side

that it appears to be absent (Fig. 135). The septum is attached to

the walls of the auricles so as to form two distinct cavities, which

communicate at the auriculo-ventricular opening, where the septum has

a free concave border (Fig. 134). The right auricle is much larger

than the left, and receives the systemic venous blood from the sinus

venosus by the opening (Figs. 133 and 134 _SV._) already described.

The left auricle receives the blood from the lungs by the pulmonary

vein (Fig. 133 _PV._), which opens into the auricle near the septum

(Fig. 134 _PV._): [the openings from the sinus venosus and from the

pulmonary vein may be so closely approximated as to be separated by the

septum only. The septum is much thinner than the auricular walls]. Both

auricles open into the ventricle by the auriculo-ventricular opening

(Fig. 134).

[Illustration: Fig. 134.

The frog’s heart seen from the ventral surface, and dissected so as to

show its structure. The ventral walls of the truncus arteriosus and of

the auricles and ventricle have been removed. From a drawing by Mr.

Hurst.

_A._ Auriculo-ventricular aperture and valve.

_B._ Aperture leading from ventricle to truncus arteriosus.

_C._ Left carotid arch.

_LA._ Left auricle.

_P._ Left pulmo-cutaneous arch.

_P′._ Style passed down right pulmo-cutaneous arch into the

truncus arteriosus, _T._

_PV._ Opening of pulmonary vein.

_RA._ Right auricle.

_S._ Left systemic arch.

_SA._ Septum atriorum.

_SV._ Opening of Sinus venosus into right auricle.

_V._ Ventricle.

]

*c.* The *ventricle* (Figs. 132, 134, and 136 _V._), examined while

still living, is seen to be bluntly conical in shape and darker on the

left side than on the right; this is due to the ventricular wall of the

right side, near the truncus arteriosus, being somewhat thinner. The

inner wall is not smooth but possesses muscular ridges (_Trabeculae_).

At its base the ventricle communicates with the auricles by the

auriculo-ventricular opening (Figs. 134 and 136), and with the truncus

arteriosus by a separate opening (Fig. 134 _A._). The ventricle is

lined by a layer of endocardial endothelium.

[Illustration: Fig. 135.

Dissection of a case in which the auricular septum is placed more to

the left than is normal.

_S_ Septum.

_SS_ Left auricle.

_V_ Opening into the Sinus venosus.

]

[Illustration: Fig. 136.

Dissection of the heart from the left side; after Howes.

_A._ Auriculo-ventricular opening.

_AC._ Anterior caval veins.

_IC._ Posterior caval vein.

_LA._ Left auricle.

_PV._ Pulmonary vein.

_RA._ Right auricle.

_SV._ Sinus venosus.

_TA._ Truncus arteriosus.

_V._ Ventricle.

]

The *auriculo-ventricular opening* (Fig. 134 _A._) is a large aperture

guarded by two valves, a dorsal and a ventral: each valve is formed

by a reduplication of the endothelium, and contains a small amount of

connective-tissue; its free margin is bound down by a number (about

twelve) of fibrous cords, the _chordae tendineae_, which are attached

by their posterior extremities to the trabeculae.

*d.* The *truncus arteriosus* (Figs. 132 _TA._, 134 _A._) arises at the

base of the ventricle, on the right side of its ventral surface, then

passes forwards and to the left, across the auricles, to reach their

anterior border near the median line, where it divides to form two

vessels.

Internally it is incompletely divided into two compartments by a spiral

valve (Fig. 134); the valve is attached to the dorsal surface and is

free ventrally: when the vessel is fully dilated (artificially), the

valve extends two-thirds of its diameter. The opening of the vessel

into the ventricle is guarded by two semilunar valves (Fig. 134 _A._),

the free margins of which are bound to the inner surface of the tube by

_chordae tendineae_. The distal extremity has a semi-lunar valve in the

left compartment (Fig. 134), which is attached by a delicate band to

the spiral valve (Ecker).

C. *The minute structure of the Heart.*

[The heart consists chiefly of muscle, but possesses also nerve-fibres,

nerve-cells, endothelium, and connective-tissue; the truncus arteriosus

has, in addition, blood-vessels.

*a.* The *muscular structure* of the heart.

(1) The *arrangement* of the *muscle-fibres* varies in the different

portions of the heart. That of the ventricle will be easily understood

by reference to Fig. 137. A layer of circular fibres (_CM_) extends

round the whole ventricle, and from it more or less radial or

transverse bands (_TM_, also Fig. 138 _TM_) pass inwards; these, in the

apical portion of the ventricle, seem to extend almost transversely

across; towards the base they are directed onwards and forwards towards

the auricles. Of these bands some, near the apex, can be traced

directly to the opposite side, others join the wall more anteriorly,

and still others join bands of muscular fibres (_LM_), which may be

described as longitudinal, and form the _Musculi papillares_. In the

anterior portion of the ventricle the same arrangement holds good, but

is modified to form a central cavity; still more anteriorly, at the

junction of auricles and ventricle, the trabecular structure is much

less apparent, the fibres being closer and mainly circular.

[Illustration: Fig. 137.

I. Transverse section through the junction of the hinder and middle

thirds of the ventricle of _R. temporaria_. II. Transverse section

through junction of middle and anterior thirds of the same heart. Occ.

I, Syst. 1, Leitz. The heart was freely distended[62] with dilute

spirit, and so hardened.--G. H.

_CC_ Central cavity.

_CM_ Circular muscle.

_LM_ Longitudinal muscle.

_TM_ Transverse muscle.

]

[Footnote 62: The base of the ventricle measured, when distended, 8 mm.

by 5 mm.]

In the auricles the muscular fibres are chiefly arranged in circular

rings which cross one another; on the inner surface, however, traces

of an arrangement similar to that in the ventricle may be made out. In

the heart, from which the sections in Fig. 137 were drawn, transverse

bands (cut longitudinally), 0.33 mm. in length, were numerous; their

central ends terminated in bands, the fibres of which were cut

transversely. The auricular septum (Fig. 141 a _M_) also possesses

muscular fibres which cross one another in its substance.

The fibres of the truncus arteriosus resemble those of an ordinary

artery; those of the sinus venosus are arranged in rings, mostly

transverse, but others oblique, and a few longitudinal.]

(2) The *structure* of the *cardiac muscle-fibres* (Figs. 138 and 141

a).

[The cardiac muscle of the frog consists of spindle-shaped cells, which

as Pohl-Pincus pointed out, are of two kinds; the main difference

between the two being in the size of the nucleus, which is much broader

in one than in the other.

[Illustration: Fig. 138.

Portion of a transverse section through the middle of the ventricle of

_R. temporaria_. From same heart as the sections in Fig. 136. Oc. I.

Syst. 7, Leitz.--G. H.

_A_ Rod-shaped nuclei of muscle fibre.

_B_ Oval nuclei of muscle fibre.

_CM_ Circular layer of muscle.

_CT_ Connective-tissue.

_E_ Endothelium of the epicardium.

_TM_ Transverse bands of muscle.

]

α. Cells with a broad nucleus (Fig. 138 _B_): the nucleus is lenticular

or egg-shaped, 4 to 7 µ broad, 8 to 14 or 18 µ long; the ratio of

length to breadth being 1 : 2.

β. Cells with a narrow nucleus (Fig. 138 _A_): the nucleus is

rod-shaped, 2·5 to 3·5 µ broad and 25 to 43 µ in length. The ratio

of length to breadth varying from 1 : 8 to 1 : 16. These cells are

more frequent in the right half of the base of the ventricle than the

former variety. In the trabeculae the second variety is more numerous

(Pohl-Pincus).

The muscle-fibre may be simple (Fig. 141 a _M_) or possess branches

(Fig. 138); the fibres are all more or less spindle-shaped, and

striated transversely and longitudinally, but possess no sarcolemma.

The protoplasmic contents of the cell are finely granular; the nucleus

much more coarsely granular and possessing one or more distinctly

marked nucleoli. The striations are due to the presence of a network,

which has been carefully described by Messrs. B. Melland and C. F.

Marshall[63], and is similar to that of ordinary voluntary muscle (see

histology of muscle).]

[Footnote 63: I have had opportunity of examining Mr. Marshall’s slides

and have convinced myself of the correctness of his observations.]

*b.* The *nerves* of the *heart*.

The nervous supply of the heart is derived from the sympathetic system

and from the cardiac branches of the pneumogastric nerve; the course of

which has already been traced (page 175) to the roots of the anterior

caval veins, where they form a simple plexus by means of a connecting

link (Fig. 139). The plexus possesses nerve-cells and sends off two

nerves into the auricular septum; the one (_d_) lies dorsally and is

shorter and thicker than the other (_v_), which lies ventrally in the

septum. They course backwards to the posterior border of the septum,

and there distribute fine branches to the base of the ventricle and

surrounding parts.

[Illustration: Fig. 139.

Course of the cardiac nerves in the auricular septum.

_d_ Dorsal nerve.

_v_ Ventral nerve.

]

[In the whole of their course in the heart they have numerous

nerve-cells, either imbedded between their fibres or attached to the

nerves and their branches.

The nerve-fibres are both medullated and non-medullated. The

nerve-cells (Figs. 139, 140, 141) are usually oval, and are unipolar

or rarely bipolar; sometimes two cells exist in the same envelope,

forming the ‘twin-cells’ of Dogiel (Fig. 141 b II). According to the

recent observations of Lavdowsky and Dogiel each cell receives one

medullated fibre and gives off a fine non-medullated fibre; the two run

together for a short distance, after which the non-medullated fibre

separates to supply muscle-fibres.

[Illustration: Fig. 140.

Group of nerve-cells on the cardiac nerve, from the auricular septum.

Stained with picrocarmine. Occ. III, Syst. 3, Hartnack; after Dogiel.

_F_ Fibres of the cardiac nerve.

_G_ Nerve-cells.

]

Two large clusters of nerve-cells have received special names.

*Remak’s ganglion* is a large group of nerve-cells in the wall of the

sinus venosus.

*Bidder’s ganglion* is formed, collectively, by groups of cells in the

auriculo-ventricular groove, the chief groups being attached to the two

cardiac nerves.

No nerve-cells have been discovered behind that portion of the

ventricle near the auriculo-ventricular groove, nor has a direct

connection between the cells or their processes with the fibres of the

pneumogastric nerve been made out.

Nerve-fibres have been described as existing in all parts of the

ventricle and being connected with the muscle-fibres (Openchowsky and

others); this cannot, as yet, be accepted as proved.

The truncus arteriosus also possesses nerve-fibres and nerve-cells

(Pagliani, Löwit, and others), but their arrangement seems to be very

irregular. Löwit did not succeed in finding the nerve-cells in all

cases examined; and Engelmann denies their occurrence.]

*c* and *d*. [The *endothelium* and *connective-tissue* of the heart.

The connective-tissue of the heart exists only in very small quantity.

A fine layer on the outer surface of the ventricle, especially towards

the auriculo-ventricular groove, can easily be seen (Fig. 138 _CT_);

the various valves, the auricular septum, and more particularly the

truncus arteriosus, all contain connective-tissue. In the latter part

the spiral valve is wholly formed of this tissue.

[Illustration: Fig. 141 a.

Small group of nerve-cells from the auricular septum. Stained with

picrocarmine and silver nitrate, after Dogiel.

_E_ Endothelium (endocardium).

_G_ Nerve-cells.

_M_ Muscle-fibres.

_N_ Nucleus.

_N_′ Nucleolus.

]

[Illustration: Fig. 141 b.

Isolated nerve-cells from frog’s heart. I. Usual form of nerve-cell.

II. ‘Twin-cell.’

_C_ Capsule.

_N_ Nucleus.

_N_′ Nucleolus.

_P_ Process.

]

The whole of the inner surface is lined with endothelium

(endocardium), which covers the various trabeculae; the external

surface of the heart is likewise covered with an epicardial layer of

endothelium (Fig. 138 _E_).]

[Illustration: Fig. 142.

Arteries and veins of the Truncus arteriosus of Bufo vulgaris, after

Hyrtl.

_a_ Arteria bulbi.

_b_ Truncus arteriosus.

_h_, _i_ Hepatic portal veins.

_t_ Right and left synangium.

_v_ Vena bulbi anterior.

_vi_ Left innominate vein.

_vc_ Vena bulbi posterior (V. cardiaca).

_A_ Right auricle.

_A_′ Left auricle.

_V_ Ventricle.

]

*e.* The blood-supply to the heart.

With the exception of the _truncus arteriosus_, no part of the heart

possesses blood-vessels (Hyrtl). [The _truncus arteriosus_ (Fig. 142

_b_) has a rich anastomosis, which receives its blood by the _arteria

bulbi_ (_a_), a branch of the carotid arch. Two veins connect this

anastomosis with the systemic veins, and with the hepatic portal veins:

the former, _vena bulbi anterior_ (_v_), passes over the auricles to

open into the left innominate vein (_vi_); the latter, the _vena bulbi

posterior_ (_vc_) or _vena cardiaca_, as it is more usually named,

passes from the _truncus arteriosus_ in a fold of pericardium, and then

courses backwards to open into the anterior-abdominal vein.]

II. THE ARTERIES.

(The original arrangement of this section has been modified.)

The _Truncus arteriosus_ divides to form two vessels (Figs. 132, 134);

each of these is divided by two partitions into three compartments,

which after a short course together separate to form three distinct

vessels, the carotid, systemic, and pulmo-cutaneous arches.

*1.* The *carotid arch* (_Canalis carotico-lingualis_, Brücke; _Ductus

caroticus_), (Fig. 143 _I_) is the most anterior of the three arches:

it winds round the side of the oesophagus as far as the carotid gland,

where it divides to form two branches; the right carotid arch gives off

a small branch to the _Truncus arteriosus_, the _Arteria bulbi_ (see

above).

[Illustration: Fig. 143.

Schema of the arterial system of Rana esculenta.

_A_ Dorsal aorta.

_c_ Cutaneous artery.

_ca_ Carotid artery.

_d_ Carotid gland.

_ic_ Iliac arteries.

_ie_ External iliac artery.

_ii_ Internal iliac artery.

_J_ Mesenteric artery.

_l_ Lingual artery.

_o_ Occipital artery.

_ov_ Occipito-vertebral artery.

_P_ Pulmonary artery.

_s_ Subclavian artery.

_T_ Truncus arteriosus.

_ug_ Urinogenital arteries.

_v_ Vertebral artery.

_I_ Carotid arch.

_II_ Systemic arch.

_III_ Pulmo-cutaneus arch.

]

The *carotid gland* (Fig. 144 _c_′) is a dilated portion of the vessel.

The interior contains bands or trabeculae of muscle, which give its

walls a more or less spongy structure; the muscle-fibres are not

transversely striated (Hoffmann).

*a.* The *lingual artery* (_Arteria lingualis_, _Arteria

hyoidea-lingualis_, _Arteria hyoidea_), (Figs. 143 _l_, 144 _L_) arises

by two roots from the inner surface of the carotid gland, and courses

forwards and inwards, supplying branches to the thyroid gland and the

_Musculi geniohyoidei_, to the ventral surface of the hyoid; another

continues forwards on the outer border of the _M. hyoglossus_ to supply

branches to the hyoid and tongue.

*b.* The *carotid artery* (_Arteria carotis communis_), (Figs. 143

_ca_, 144 _c_″) arises by several roots from the outer wall of the

carotid gland; and passes over the _Musculus petrohyoideus I_ to the

anterior end of the oesophagus, round which it courses towards the

vertebral column, and then runs forwards to the base of the skull.

The artery then courses forwards, lying immediately above the mucous

membrane and underneath the transverse arm of the parasphenoid, which

it crosses about midway between its back and external end; [at this

point the artery gives off two palatine arteries, then ascends in the

orbit to the origin of the _M. pterygoideus_, and divides into two

branches, the internal carotid artery and the ophthalmic artery]. The

branches are:--

(1) The pharyngeal artery (_Arteria pharyngea ascendens_), (Fig. 145

_p_) which runs towards the opening of the Eustachian tube,

distributes branches to the pharynx, and anastomoses with the _Arteria

pharyngeo-maxillaris_ of the cutaneous artery, and with the _Ramus

inframaxillaris_ of the occipital artery.

[Illustration: Fig. 144.

Right carotid gland.

_c_ Carotid arch.

_c_′ Carotid gland.

_c_″ Carotid artery.

_L_ Lingual artery.

]

(2) [The posterior palatine artery (_Arteria palatina posterior_,

Virchow) immediately divides into a number of small twigs to supply the

mucous membrane of the hinder and outer parts of the gums.]

(3) The anterior palatine artery (_Arteria palatina anterior_, Virchow;

_Arteria palatina_, Ecker), (Fig. 145 _p_′) runs forwards, between

the _Musc. levator bulbi_ and the mucous membrane, with the _Ramus

palatinus_ of the trigeminal nerve. [On reaching the palatine bone,

it curves outwards to the outer anterior angle of the orbit, where it

gives off a branch backwards along the upper jaw to the hinder portion

of the orbit.] In its course it supplies twigs to the surrounding

tissues, but chiefly to the mucous membrane and Harder’s gland.

[Illustration: Fig 145.

Arterial system of _Rana esculenta_.

_A_ Dorsal aorta.

_Ad_ Right systemic arch.

_As_ Left systemic arch.

_c_ (opposite fore-limb) Carotid artery.

_c_ (in abdomen) Coeliac artery.

_cm_ Cutaneous artery.

_e_ Lingual artery.

_H_ Heart.

_H_′ Harderian gland.

_L_ Liver.

_Lu_ Lung.

_m_ Mesenteric artery.

_M_ Stomach.

_M_′ Spleen.

_o_ Ophthalmic artery.

_p_ Pharyngeal artery.

_p_′ Anterior palatine artery.

_S_ Subclavian artery.

_I_ Carotid arch.

_II_ Systemic arch.

_III_ Pulmo-cutaneous arch.

]

(4) The internal carotid artery (_Art. cerebralis_, Virchow); see

Arteries of Brain, p. 162.

(5) The ophthalmic artery (_Art. ophthalmica_), (Fig. 145 _o_) [is

larger than the internal carotid artery at this point; it crosses the

origins of the _M. rectus externus_ and of the _M. retrahens bulbi_

and reaches the optic nerve; the artery then runs forwards along the

under surface of this nerve and of the eyeball, lying close to the

sclerotic coat as far as a little beyond the equator, where it pierces

the sclerotic coat so obliquely that the choroid coat is only reached

at the ciliary processes. In its course the artery supplies:--

α. Muscular branches, given off at the point where the artery reaches

the eyeball.

β. Two arteries (_AA. ciliares_) to the choroid, given off at the same

point as the foregoing.

γ. Two arteries to the iris.

δ. The _Art. hyaloidea_, the terminal portion of the ophthalmic artery.

(For further description of these vessels, see Eye.)].

*2.* The *systemic arch* (_Ductus aorticus_), (Figs. 132, 143, and 145

_II_) is the middle arch of the three; it arises from the middle canal

of the _Ductus arteriosus_, and winds obliquely round the oesophagus

between the _MM. petrohyoidei I_ and _II_ towards the vertebral

column, which it reaches at about the level of the sixth vertebra. The

right systemic arch is continued as the dorsal aorta, the left arch

communicates with it merely by a small opening, and is then continued

as the coeliaco-mesenteric artery.

Branches of the systemic arch:--

*a.* The *laryngeal artery* (_Arteria laryngea_), (Fig. 146 _lg_)

arises from the inner border of the systemic arch before it reaches the

oesophagus; it passes forwards and inwards to be distributed in the

larynx and pharynx. According to Brücke this artery possesses a valve

at its point of origin.

[Illustration: Fig. 146.

Transverse section at level of the larynx.

_C_ Carotid gland.

_G_ Glottis.

_lg_ Laryngeal artery.

_S_ Subclavian artery.

_I_ Carotid arch.

_II_ Aortic arch.

]

*b.* The *oesophageal arteries* (_Arteriae oesophageae_) pass from the

upper part of the arch to the dorsal surface of the oesophagus.

*c.* The *occipito-vertebral artery* (Ecker), (_Art.

occipito-vertebralis_), (Fig. 147 _o_, _v_) [arises from the systemic

arch immediately in front of the transverse process of the second

vertebra, and ascends immediately in contact with the body of the first

vertebra, which it separates from the _MM. intertransversarii capitis_

and the sympathetic cord]. Immediately under the most anterior portion

of the _M. longissimus dorsi_ the artery divides into two branches, the

vertebral and occipital arteries:--

(1) The *vertebral artery* (_Arteria vertebralis_ or

_supra-vertebralis_) (Fig. 147 _v_) courses backwards lying

on the transverse processes of the vertebrae and on the _MM.

intertransversarii_, under cover of the _M. longissimus dorsi_. In this

course it is close to the oblique processes, and may even be partially

covered by these; it extends beyond the sacrum, and gives off:--

α. The _Rami spinales_, which pass through the intervertebral foramina

to the spinal cord (see Arteries of Spinal Cord, p. 162).

[Illustration: Fig. 147.

Dissection to show the occipito-vertebral and the cutaneous arteries.

The skin of the back has been divided by a median incision and drawn to

either side; the right arm dislocated and removed through the opening

in the skin (at _A_). On the left side the extensor muscles of the back

have been removed.

_c.m._ Cutaneous artery.

_c.p._ Arteria cutanea pectoris.

_o_ Occipital artery.

_s_ Scapular artery.

_v_ Vertebral artery.

]

β. The _Rami dorsales_; these supply the muscles of the back and send

branches to the skin, which accompany the cutaneous branches of the

spinal nerves through the _Saccus lymphaticus cranio-dorsalis_.

γ. The _Rami intercostales_ are distributed in part to the _MM.

intertransversarii_; other twigs pass ventrally to the periganglionic

glands; the longest branches accompany the anterior divisions of the

spinal nerves to the muscles and skin of the belly (Fig. 145).

(2) The *occipital artery* (_Arteria occipitalis_), (Figs. 147 and 148

_o_), the anterior branch of the occipito-vertebral artery, pierces

the _M. longissimus dorsi_; then courses forwards under cover of the

_Fascia dorsalis_ and upon the _M. temporalis_, and divides, behind the

eyeball [at the anterior border of the _M. temporalis_, Virchow], into

two branches:--

α. The _Ramus orbito-nasalis_ (Fig. 148 _on_) accompanies the

ophthalmic nerve along the wall of the cranium, and passes through the

sphenethmoid, with the nasal branch of the ophthalmic nerve, to the

nose, where it divides into two branches supplying the mucous membrane

of the nose. In its course through the orbit the artery supplies

branches to the Harderian gland and neighbouring parts, and anastomoses

with the ascending branch of the _Arteria palatina_.

β. The _Ramus maxillaris communis_ (_Art. temporalis_, Virchow),

(Fig. 148 _m_) runs downwards and outwards along the anterior border of

the _M. temporalis_, and passes under the anterior arm of the squamosal

bone to course backwards on the upper jaw. At this point the _Ramus

maxillaris superior_ (_m_′) is given off. The main artery continues its

backward course to the tympanic membrane, where it divides into its

three terminal branches:--

[Illustration: Fig. 148.

Branches of the occipito-vertebral and cutaneous arteries in the head;

side view. The skin (_H_) has been reflected downwards.

_c.m._ Cutaneous artery.

_l.a._ M. levator scapulae.

_m_ Art. maxillaris communis.

_m_′ Art. maxillaris superior.

_m_″ Art. maxillaris inferior.

_o_ Occipital artery.

_on_ Art. orbito-nasalis.

_p.m._ Art. pharyngo-maxillaris.

_v_ Vertebral artery.

]

[(1) The _Ramus maxillaris superior_ (_AA. maxillares superiores_,

Virchow), usually not a single artery but a number of small twigs,

which supply the space between the eye and the upper jaw.

(2) A small ascending branch to the tympanic membrane (Virchow).

(3) A _Ramus auricularis_ (Virchow), which forms a rich anastomosis

on the hinder wall of the tympanic cavity, and supplies a twig to the

tympanic membrane. This passes from the upper border of the membrane

to the point of attachment of the _Columella auris_ (extrastapedial),

round which it forms a circular anastomosis. The _Ramus auricularis_

also anastomoses with the cutaneous artery.

[Illustration: Fig. 149.

Dissection to show the occipito-vertebral and the cutaneous arteries.

The skin of the back has been divided by a median incision and drawn to

either side; the right arm dislocated and removed through the opening

in the skin (at _A_). On the left side the extensor muscles of the back

have been removed.

_c.m._ Cutaneous artery.

_c.p._ Arteria cutanea pectoris.

_o_ Occipital artery.

_s_ Scapular artery.

_v_ Vertebral artery.

]

(4) A branch passing inwards to the fat-body, and lying immediately in

front of the deltoid muscle (Virchow).

(5) A small branch running on to the inner surface of the angle of the

jaw and then forwards (Virchow).

(6) A larger vessel, the _Ramus maxillaris inferior_ (Fig. 148 _m_″),

which arises from the _Ramus maxillaris superior_ under cover of the

tympanic membrane, and accompanies the _Ramus mandibularis_ of the

trigeminal nerve through the _M. masseter_, or between this muscle and

the _M. temporalis_ to the mandible, and then courses forwards to the

chin. A large cutaneous branch is given off half-way along the floor

of the mouth. The artery gives off many small twigs to supply the

neighbouring structures.]

3. The *pulmo-cutaneous arch* (_Ductus pulmo-cutaneus_), (Figs. 132

143, and 145 _III_) is the hindmost (fifth embryonic) of the three

persistent arches; it passes upwards and forwards on the oesophagus,

and at the level of the carotid gland divides into two branches, the

pulmonary artery and the cutaneous artery.

a. The *pulmonary artery* (_Arteria pulmonalis_) runs backwards to the

root of the lung, along the outer surface of which it is continued in a

sinuous course. It supplies the lung.

b. The *cutaneous artery* (_Arteria cutanea magna_), (Fig. 149

_c.m._) [passes along the deeper surface of the _M. petrohyoideus

III_ until near the prootic bone; in this course it travels forwards,

outwards, and upwards, crosses the outer border of the muscles,

and appears between the _M. levator anguli scapulae_ and the _M.

sternocleidomastoideus_. At this point the artery curves sharply round

to reach the angle of the jaw, the hinder border of the _M. depressor

maxillae inferioris_. The curve so formed and the descending limb are

covered by the _M. depressor maxillae inferioris_. The artery then

passes backwards to form a rich anastomosis in the skin almost as far

as the posterior end of the trunk. It supplies the following branches:--

(1) The _Ramus dorsalis_ (Virchow) arises immediately in front of the

curve of the main artery; it ascends behind the _M. temporalis_, and

passes along the anterior border of the _M. depressor maxillae_ to the

skin, where it curves backwards. It supplies small twigs forwards as

far as the eyelids.

(2) The _Ramus auricularis_ (Virchow) also arises in front of the

curve in the cutaneous artery; sometimes it arises from the _Ramus

dorsalis_. The artery winds around the _M. petrohyoideus III_ near its

insertion, and so reaches the posterior wall of the tympanic cavity,

where it anastomoses with the _R. auricularis_ of the _Ramus maxillaris

communis_.]

(3) The _Arteria pharyngo-maxillaris_ (Fig. 148 _p.m._) is a small

branch running forwards and outwards to supply the mucous membrane of

the larynx, Eustachian tubes, gums, the lower jaw, and the skin under

the mouth. It forms a rich anastomosis with branches of the _Ramus

maxillaris inferior_ of the occipital artery, and with others from the

pharyngeal branch of the carotid artery; a large branch passes forwards

to supply the skin of the throat and floor of the mouth.

(4) The _Arteria cutanea pectoris_ (Fig. 149 _c.p._) is distributed to

the skin of the breast.

*4.* The *subclavian artery* (_Arteria subclavia_), (Fig. 150 _s_). The

subclavian artery arises from the systemic arch immediately behind the

occipito-vertebral artery at the level of the second or third vertebra.

The artery then runs outwards alongside and in front of the brachial

nerve, between the _M. subscapularis_ and the _M. triceps_ to the arm,

where it is known as the brachial artery; in its course it gives off:--

*a.* The _Arteria costo-cervicalis_ (Fig. 150, 1). This artery arises

near the systemic arch, and runs backwards, parallel to the vertebral

column, over the brachial nerve, and across the transverse processes of

the vertebrae. It is distributed more especially to the _M. obliquus

internus_.

*b.* The _Arteria coraco-clavicularis_ (Fig. 150, 2) arises more

externally than the last artery; it accompanies the coraco-clavicular

nerve on the dorsal surface of the ventral portion of the

shoulder-girdle, and passes between the _M. deltoideus_ and _M.

subscapularis_ into the _Foramen ovale_, where it divides to supply

these muscles and the _M. pectoralis_.

[Illustration: Fig. 150.

Subclavian artery of the left side.

_A_ Systemic arch.

_A_′ Dorsal aorta.

_c_ Arteria cutanea medialis superior.

_c_′ Arteria cutanea medialis inferior.

_ls_ M. levator scapulae.

_o.v_ Occipito-vertebral artery.

_p_ Arteria pectoralis.

_R_ Radial artery

_s_ Subclavian artery.

_U_ Ulnar artery.

1 Arteria costo-cervicalis.

2 Arteria coraco-clavicularis.

3 Arteria scapularis posterior.

4 Arteria scapularis superior.

5 Cutaneous artery.

6 Arteria subscapularis.

]

*c.* The _Art. scapularis posterior_ (Fig. 150, 3), a small branch

arising opposite at the tip of the second transverse process and

passing backwards over the _MM. transverso-scapulares_, to which it is

distributed.

[Illustration: Fig. 151.

Arteries of the palmar surface of the hand.

_B_ Ulnar artery.

_ci_ R. cutaneus medialis inferior.

_d_ Digital branches.

_F_ M. flexor digitorum communis.

_II_ Thumb.

_V_ Fifth finger.

]

[Illustration: Fig. 152.

Arteries of the dorsal surface of the hand.

_B_ Ulnar artery.

_cri_ R. cutan. rad. inferior.

_R_ Radial artery.

_II_ Thumb.

_V_ Fifth finger.

]

*d.* The _Art. scapularis superior_ (Fig. 150, 4) accompanies several

nerves between the _M. subscapularis_, _M. transverso-scapularis

major_, and the long head of the _M. triceps_ towards the dorsal

surface, where it is distributed to the _MM. infraspinatus_,

_latissimus dorsi_, etc.

*e.* The _Ramus cutaneus maxillaris_.

*f.* The _Art. subscapularis_ (Fig. 150, 6) is distributed in the

subscapular muscle.

A. The *brachial artery* (_Arteria brachialis_), (Fig. 150) accompanies

the brachial nerve under cover of the long head of the _M. triceps_ and

over its inner head to the _Plica cubiti_, into which it sinks near the

tendon of the _M. sternoradialis_; beyond this point it is continued as

the ulnar artery. In its course the brachial artery gives off:--

*a.* The _Arteria pectoralis_ (Fig. 150 _p_).

*b.* The _Arteria cutanea medialis superior_ (Fig. 150 _c_).

*c.* The *radial artery* (_Arteria radialis_), (Fig. 150 _R_) which

winds round the humerus with the radial nerve between the muscles of

the outer side of the forearm, to which it gives branches: in the palm

of the hand it passes between the _M. extensor carpi ulnaris_ and the

_M. flexor antibrachii_ to the dorsal surface of the hand, where it

anastomoses with the ulnar artery. In its course it supplies:--

(1) Muscular branches to the surrounding muscles.

(2) The _Ram. cut. radialis inferior_, which arises from the radial

artery towards the distal end of the forearm, and passes underneath the

_M. flexor carpi ulnaris_ to the skin.

B. The *ulnar artery* (_Arteria ulnaris_), (Figs. 150 _U_, 151 _B_)

accompanies the ulnar nerve towards the hand, passes between the _M.

flexor digitorum communis_ and the _M. anconeus_ to the palm of the

hand, and then winds round the outer side to the dorsal surface of the

carpus to form its terminal branches. The branches are:--

(1) The _Ram. cutan. med. inferior_ (Fig. 151 _ci_); it arises near the

wrist-joint and supplies (α) twigs to the skin of the inner surface;

(β) a branch downwards, which courses along the inner side of the

second finger to its tip; and (γ) a branch which passes upwards to

anastomose with the _R. cutaneus medialis superior_.

(2) In the palm of the hand a branch (Fig. 151 _d_) is given off,

which divides to form _Arteriae digitales volares_: one for the second

finger, two for the third finger, two for the fourth finger, and one

for the fifth finger.

(3) Given off on the dorsum of the hand (Fig. 152) it supplies branches

corresponding exactly with those of the palmar surface.

*5.* The *dorsal aorta* (_Aorta communis s. abdominalis_) (Fig. 153

_A_). The two systemic arches unite at about the level of the sixth

vertebra to form the dorsal aorta: as already mentioned the union is

very incomplete, the dorsal aorta being practically a continuation of

the right systemic arch, and only communicating with the left by means

of a small opening. The branches are as follows:--

*a.* The *coeliaco-mesenteric artery* (_Arteria intestinalis

communis_), (Fig. 153), the true continuation of the left systemic

arch, at once divides to form the gastric and mesenteric arteries.

(1) The *gastric artery* (_Arteria gastrica s. coeliaca_) (_c_)

immediately divides into two branches:--

α. The _Ramus dexter s. anterior_; this artery gives off the _Arteria

hepatica_ to the liver and gall-bladder and passes to the right

(anterior) surface of the stomach, running along the attachment of

the mesentery and supplying branches forwards to the oesophagus and

backwards towards the intestine.

β. The _Ramus sinister s. posterior_ passes to the left (posterior)

surface and behaves like the _Ram. dexter_.

(2) The *mesenteric artery* (_Arteria mesenterica superior_ and

_inferior_, and the _Arteria splenica_), (Fig. 153 _m_) supplies:--

α. The superior mesenteric artery, which supplies the upper part of

the intestine, and gives off a recurrent branch to the stomach, which

anastomoses with the gastric arteries.

[Illustration: Fig. 153.

Arterial system of _Rana esculenta_.

_A_ Dorsal aorta.

_Ad_ Right systemic arch.

_As_ Left systemic arch.

_c_ (opposite fore-limb) Carotid artery.

_c_ (in abdomen) Coeliac artery.

_cm_ Cutaneous artery.

_e_ Lingual artery.

_H_ Heart.

_H_′ Harderian gland.

_L_ Liver.

_Lu_ Lung.

_m_ Mesenteric artery.

_M_ Stomach.

_M_′ Spleen.

_o_ Ophthalmic artery.

_p_ Pharyngeal artery.

_p_′ Anterior palatine artery.

_S_ Subclavian artery.

_I_ Carotid arch.

_II_ Systemic arch.

_III_ Pulmo-cutaneous arch.

]

β. The splenic artery to the spleen.

γ. The inferior mesenteric artery to the lower part of the intestine.

All the arteries to the intestine course in lymph-spaces.

*b.* The *urinogenital arteries* (_Arteriae urino-genitales_),

(Fig. 154 _ug_) are four to six small arteries arising from the ventral

surface of the aorta between the kidneys: they immediately divide into

right and left branches to supply the kidneys, fat-bodies, reproductive

organs, and their ducts (see kidney).

*c.* The *lumbar arteries* (_Arteriae lumbales_), are small paired

vessels to the neighbouring muscles, and send branches through the

intervertebral foramina to the vertebral canal. Sometimes these

arteries arise from the _Art. spinalis_ or _Art. vertebralis_, etc.;

in the former case they pass outwards from the vertebral canal to the

surrounding structures.

*d.* The *haemorrhoidal artery* (_Art. mesenterica inf. s Art.

haemorrhoidalis superior_) is a small median artery running from the

posterior end of the aorta to the large intestine.

*6.* *Bifurcation of the aorta.* *The iliac arteries* (_Arteriae

iliacae communes_). Opposite the middle of the urostyle the aorta

bifurcates to form the iliac arteries; each of these courses on the

corresponding sciatic plexus to the thigh, beyond which it is continued

as the sciatic artery. The branches of the iliac artery are:--

*a.* The *vesico-epigastric artery* (_Arteria epigastrico-vesicalis_),

(Fig. 155 _ev_); it arises near the bifurcation, passes outwards on the

plexus, and divides into two branches:--

(1) The epigastric artery (_Arteria epigastrica_) (_e_) immediately

gives off a recurrent branch to the iliac bone and the _M.

ilio-coccygeus_; it then continues to the ventral muscles of the

abdominal wall, where it runs forwards on the deeper surface of the _M.

obliquus internus_, and is distributed more especially to this muscle.

[Illustration: Fig. 154.

The urinogenital arteries.

_A_ Dorsal aorta.

_H_ Testis.

_N_ Kidney.

_ug_ Urinogenital arteries.

]

(2) The hypogastric artery (_Arteria vesicalis_) (_v_) winds downwards

around the sciatic plexus to reach the mesentery, along which it

courses to the bladder, where it forms a rich anastomosis.

*b.* The *femoral artery* (_Arteria femoralis s. cruralis_), (Fig. 155

_f_) is the representative of the femoral artery of man, although it

does not perform the same functions. The artery passes under the hinder

free border of the _M. obliquus internus_ to the thigh, where it lies

upon the _M. ilio-psoas_, and sends twigs to the neighbouring muscles,

the skin, and lymph-sac (_Lamina inguinalis_), and then divides into:--

(1) A twig, which runs forwards on the ventral surface of the iliac

bone and is covered by the epigastric artery.

(2) A twig, running forwards and downwards on the _MM. ilio-psoas_,

_vastus internus_, and _sartorius_, to send branches to the hip-joint.

*c.* The *spermatic artery* (_Arteria spermatica_), which arises, in

males, from the iliac artery, and ascends to the _Vesicula seminalis_.

[Illustration: Fig. 155.

Bifurcation of the aorta and the iliac arteries.

_e_ Epigastric artery.

_ev_ Vesico-epigastric artery.

_f_ Femoral artery.

_o_ Internal oblique muscle.

_v_ Vesical artery.

]

*7.* *Arteries of the hinder extremity.*

A. The *sciatic artery* (_Arteria ischiadica_), (Fig. 156 _i_) is

the continuation of the iliac artery; it leaves the pelvis between

the hinder (outer) border of the _M. coccygeo-iliacus_ and the inner

surface of the origin of the _M. ilio-psoas_, then lies between the

former muscle and the origin of the _M. vastus externus_. It appears

on the thigh between the _M. vastus externus_ (in front) and the _M.

pyriformis_ (behind); it is the main arterial trunk for the hinder

extremity. The artery courses backwards, lying upon the sciatic

nerve and between the _M. biceps_ and the _M. semimembranosus_; in

this course it runs in a lymph-space placed in the _Septum femorale

superius_ (see page 259). Arrived at the popliteal space the artery

bifurcates to form the peroneal and tibial arteries; near the pelvis

it gives off two branches (*a*) and (*b*), and in its course along the

thigh two branches (*c*) and (*d*).

*a.* The _Art. haemorrhoidalis inferior_ (_h_) runs backwards and

inwards under the _M. pyriformis_ to be distributed in the _M.

sphincter ani_ and the skin of the anus.

*b.* The _Art. cutanea femoris posterior v. superior_ (_cf_′) runs

with the corresponding nerve and the _M. pyriformis_, and over the _M.

semimembranosus_ to the skin of the inner and hinder surfaces of the

thigh.

*c.* The _Rami musculares_ supply the _M. semimembranosus_, the _M.

biceps_, and the _MM. extensores auris_; the branch to the _M. rectus

inferior major_ pierces the muscle in company with the nerve, and is

continued as:--

*d.* The _Art. cutanea femoris media_ (_cf_″) to the skin of the middle

of the inner surface of the thigh. Several small twigs pierce the _M.

rectus internus minor_, reach the skin, and communicate with this

artery.

[Illustration: Fig. 156.

Arteries of the hinder extremity.

_cc_ Art. cutanea calcanei.

_cf_ Cutaneous branch of the femoral artery.

_cf_′ Art. cutanea fem. post. superior.

_cf_″ Art. cutanea femoris media.

_cgl_ Art cut. genu lateralis superior.

_cgl_′ Art. circumflexa genu lateralis inferior.

_cgm_ Art. circumflexa genu superior medialis.

_cgm_′ Art. circumflexa genu inferior medialis.

_ci_ M. coccygeo-iliacus.

_g_ M. gastrocnemius.

_gl_ M. glutaeus.

_h_ Art. haemorrhoidalis inferior.

_i_ Sciatic artery.

_ml_ Art. malleolaris lateralis.

_mm_ Art. malleolaris medialis.

_p_ Art. peronea.

_p_′ Muscular branch to the M. peroneus.

_P_ M. pyramidalis.

_ra_ M. rectus anterior.

_ri_″ M. rectus internus.

_sm_ M. semimembranosus.

_su_ Art. suralis.

_t_ Tibial artery.

_to_ Aperture in tibio-fibula.

_tr_ Art. tarsea.

_ve_ M. vastus externus.

]

B. The *peroneal artery* (_Art. peronea_), (Fig. 156 _p_) runs

outwards under cover of the tendon of the _M. biceps femoris_, and

accompanies the peroneal nerve downwards to give off:--

*a.* The _Art. circumflexa genu lateralis superior_, which runs

forwards on the outer side of the knee, gives a twig to the joint, and

is continued as:--

*b.* The _Art. cutanea genu lateralis superior_ (_cgl_) to the skin of

the anterior and outer sides of the knee. It anastomoses with branches

of the _Arteriae circumflexae genu mediales_ to form a rich anastomosis

(_Rete articulare genu_): other branches pass upwards to inosculate

with the _Art. cutanea femoris anterior_ (_cf_), and others downwards

to anastomose with the next artery.

*c.* The _Art. circumflexa genu lateralis inferior_ (_cgl_′) runs under

the tendon of origin of the gastrocnemius to the outer side of the

knee, and terminates in the skin as the _Art. cutanea genu lateralis

inferior_. Some of the cutaneous twigs pass upwards to anastomose with

the _Art. cut. genu lat. superior_, others downwards to anastomose with

the _AA. malleolares_.

*d.* A muscular branch (_p_′) for the _M. peroneus_; it is the real

continuation of the peroneal artery, and runs behind the outer tendon

of origin of the gastrocnemius to be distributed in the _M. peroneus_.

C. The *tibial artery* (_Arteria tibialis_), (Fig. 156 _t_), arises at

the bifurcation of the sciatic, and runs inwards, meeting the tibial

nerve between the tendons of origin of the _M. gastrocnemius_, and

deep in the popliteal space under cover of the calf-muscles. It then

pierces the _M. tibialis posticus_, and leaves this to perforate the

tibio-fibula (see p. 50), appearing again on the anterior surface as

the anterior tibial artery. Before meeting the tibial nerve this artery

gives off the following branches:--

*a.* The _Art. circumflexa genu superior medialis_ (_cgm_), which winds

round to the front of the knee-joint, and gives off a cutaneous branch,

the _Art. cutanea genu medialis superior_, which is distributed in the

skin of the inner side of the knee and anastomoses above with the _Art.

cutanea femoris media_, below with the following branches (*b*, *c*,

and *d*).

*b.* The _Art. circumflexa genu inferior medialis_ (_cgm_′) supplies

twigs to the knee-joint, and passes to the skin of the knee and inner

surface of the calf as the _Art. cutanea inferior medialis_.

*c.* Muscular twigs to the _M. gastrocnemius_.

*d.* The _Art. suralis_ (_su_); this accompanies the _N. suralis_

downwards along the calf-muscle, and is distributed in the skin of the

inner side of the heel.

*e.* While passing between the fibres of the _M. tibialis posticus_,

the tibial artery gives off muscular twigs; one larger than the rest is

known as the _Ramus descendens_.

D. The *anterior tibial artery* (_Arteria tibialis antica_), (Fig. 157

_t.a._) is the continuation of the tibial artery after piercing the

tibio-fibula. It runs downwards on the anterior surface of the leg,

then between the two heads of the _M. tibialis anticus_ to the dorsum

of the foot, where it is continued as the _Arteria dorsalis pedis_. It

supplies:--

[Illustration: Fig. 157.

Arteries of the dorsal surface of the foot.

_c.c._ Art. cutanea calcanei.

_c.c.i._ Art. cutanea cruris inferior.

_Ft_′ M. flexor tarsi.

_h_ Art. cutanea hallucis.

_i1_ Art. interossea dorsalis I.

_i2_ Art. interossea dorsalis II.

_i3_ Art. interossea dorsalis III.

_i4_ Art. interossea dorsalis IV.

_ml_ Art. malleolaris lateralis.

_m.m._ Art. malleolaris medialis.

_t.a._ Anterior tibial artery.

_ta_* Bifurcation of the dorsalis pedis artery.

_tr_ Art. tarsea.

]

*a.* _Rami musculares_ to the _M. peroneus_, _M. extensor cruris_, and

to the _M. flexor tarsi anterior_.

*b.* The _Art. cutanea cruris inferior_ (_c.c.i._) passes between the

two heads of the _M. tibialis anticus_ to the skin of the anterior and

outer sides of the leg.

*c.* The _Art. cutanea calcanei_ (_c.c._) is given off at the

ankle-joint; it runs outwards on the lower end of the tibio-fibula,

giving branches to the joint, then backwards between the tibio-fibula

and the _M. peroneus_ to be distributed to the skin of the heel. Above

it anastomoses with the _A. circumflexa genu inf. lateralis_, and below

with the malleolar arteries.

*d.* The _Art. malleolaris lateralis_ (_ml_) arises a little beyond

the last artery; it runs outwards under the origin of the _M. flexor

tarsi_, gives branches to the joint and muscles, and terminates in the

skin of the outer border and dorsal surface of the foot.

*e.* The _Art. malleolaris medialis_ (_m.m._) arises at about the

same level as the _Art. malleolaris lat._ from the inner border

of the anterior tibial artery. It supplies branches to the joint,

gives numerous branches to the skin, which anastomose with the _Art.

suralis_, and passes to the inner border of the foot, where it ends by

distributing branches to the _M. plantaris_, the extensor aponeurosis,

and, as the _Art. cutanea plantaris_, to the skin of the sole of the

foot.

E. The *dorsal artery* of the foot (_Art. dorsalis pedis_), (Fig. 157

_t.a._) runs under the _M. flexor tarsi posterior_ to the point _ta_*,

where it is superficial; the artery then bifurcates; the branches are:--

[Illustration: Fig. 158.

Arteries of the sole of the foot.

_i_ Art. interossea.

_pf_ Rami perforantes.

]

*a.* The _Art. tarsea_ (_tr_) which arises from the dorsal artery of

the foot while under cover of the _M. flexor tarsi posterior_; it

passes to the _M. extensor digiti V_ and to the skin of the dorsum and

outer border of the foot. One branch courses along the outer border of

the fifth toe to its tip; another passes upwards to anastomose with the

neighbouring vessels.

*b.* The inner branch, formed by the bifurcation of the dorsal artery

of the foot, at once divides to form:--

(1) The _Art. cutanea hallucis_ (_h_) to the supplemental great toe.

(2) The _Art. interossea dorsalis I_ (_i^1_), which at once bifurcates

to form two _Rami digitales_ for the first and second toes; these

inosculate with the _Art. cutanea plantaris_, and send twigs to the

skin of the inner border of the foot.

*c.* The outer branch formed by the bifurcation of the dorsal artery

divides to form the _Arteriae interossei dorsales II_, _III_, and _IV_

(_i^2_, _i^3_, _i^4_); they are distributed in a rich anastomosis to

the web.

*d.* The _Arteria interossea_ (Fig. 158 _i_) arises from the dorsal

artery of the foot at the tarsus, and perforates the membrane between

the astragalus and calcaneum to reach the sole of the foot. It forms a

rich subcutaneous plexus on the calcar, and anastomoses with the _Rami

perforantes_ (Fig. 158 _pf_), which pass above the upper borders of the

_Musculi interossei_ to reach the plantar surface, where they give off

small _Rami interossei volares_ to anastomose with the _AA. interosseae

dorsales_.

III. THE VEINS.

(The arrangement of this section has been modified.)

The blood, distributed to the body by means of the arteries, is

returned to the heart by the veins, which conduct the blood directly or

indirectly to the auricles. The pulmonary vein opens directly into the

left ventricle, the three _Venae cavae_ open into the _Sinus venosus_.

A. *Vein opening into the left auricle.*

The *pulmonary vein* (_Vena pulmonalis communis_), (Fig. 159 _vp_).

The blood returning from each lung is gathered into a vessel lying

along the inner side of the root of the lung. That of the right side

is somewhat longer than the left; the two vessels (_Venae pulmonales

dextra et sinistra_) run above the corresponding anterior caval veins

to form a common trunk, the pulmonary vein, which opens into the left

auricle (see p. 215).

B. *Veins opening into the* _Sinus venosus_.

The veins opening into the _Sinus venosus_ are the three caval veins;

of these the two anterior open into the anterior angles of the _Sinus

venosus_, the posterior into the posterior angle.

*1.* The *Anterior Caval Veins* (_Vena cava superior_), (Figs. 133 _V_,

159 _C a_) are a pair of large veins opening into the _Sinus venosus_

and returning to it blood from the fore-limbs and sides of the head.

Each vein is formed by the union of a small vein, the external jugular,

with two large veins, the innominate and the subclavian veins.

*a.* The *external jugular vein* (_Vena jugularis externa_), (Fig. 159

_je_) is formed, at the outer border of the hyoid bone, by the union of

two veins:--

(1) The lingual vein (_Vena lingualis_) (Fig. 161 _l_). This originates

at the tip of the tongue and passes in a very tortuous course along the

ventral surface of the hyoid; it receives numerous branches from the

tongue and hyoid muscles.

(2) The mandibular vein (_Vena maxillaris inferior_), (Fig. 161 _m_)

runs along the insertion of the _M. submaxillaris_, and turns inwards

at its hinder border to join the lingual vein.

*b.* The *innominate vein* (_Vena anonyma_), (Fig. 160 _A_) is formed

by the union of the internal jugular and the subscapular veins. By

tracing the innominate vein upwards between the suspensorium and the

shoulder-girdle, the internal jugular vein will be seen to receive

the vertebral vein at the outer border of the _M. levator scapulae_.

The left innominate vein receives the _Vena bulbi anterior_ from the

_Truncus arteriosus_ (p. 222).

[Illustration: Fig. 159.

Schema of the veins of _Rana esculenta_.

_a_ Subclavian vein.

_ab_ Anterior abdominal vein.

_Ad_ Right auricle.

_As_ Left auricle.

_Ca_ Anterior vena cava.

_Cp_ Posterior vena cava.

_cm_ Cutaneous vein.

_d_ Vena dorso-lumbalis.

_D_ Intestine.

_f_ Femoral vein.

_i_ Sciatic vein.

_ic_ Vena communicans iliaca.

_je_ External jugular vein formed by lingual and maxillary

veins _l_ and _m_.

_ji_ Internal jugular vein.

_L_ Hepatic veins.

_N_ Kidney.

_o_ Oviducal veins.

_P_ Hepatic portal vein.

_raI_ Renal portal vein.

_raII_ Secondary renal portal veins.

_re_ Renal veins.

_s_ Subscapular vein.

_SC_ Sinus venosus.

_va_ Innominate vein.

_vc_ Vena bulbi posterior (cardiac vein).

_vp_ Pulmonary veins.

]

(1) The *internal jugular vein* (_Vena jugularis_), (Fig. 160 _J_)

can be traced from the border of the _M. levator scapulae_ upwards

and forwards to the hinder part of the prootic bone: it then passes

forwards under the lateral process of the prootic, in a groove on the

anterior surface of the same bone, towards the hinder angle of the

orbit, where it receives the veins from the cranial cavity (see vessels

of brain, p. 165), and also a branch from the orbit.

[Illustration: Fig. 160.

Distribution of the internal jugular vein and the anterior portion of

the cutaneous vein.

_A_ Innominate vein.

_c.m._ Cutaneous vein.

_f_ Nasal vein.

_h_ Harderian gland.

_H_ Skin, reflected downwards.

_J_ Internal jugular vein.

_L_ Anterior lymph-heart.

_l_ M. levator scapulae.

_O_ M. obliquus internus.

_o_ Vena orbitalis anterior.

_o_′ Vena orbitalis posterior.

_P_ Lateral process of the prootic bone.

_S_ Subscapular vein.

_S_′ Shoulder-girdle.

_T_ Tympanic membrane.

_V_ Vertebral vein.

_2_ Transverse process of the second vertebra.

_3_ Transverse process of the third vertebra.

_4_ Transverse process of the fourth vertebra.

]

α. [The _Vena orbitalis media_ (Virchow); it arises from the anterior

internal angle of the orbit, from the Harderian gland, the nasal

cavity, or from other veins on the anterior wall of the orbit. The

vessel lies close to the cranium, immediately underneath the _Arteria

nasalis_, and sinks under the _M. rectus superior_ to join the internal

jugular vein. In its course it receives the _Vena bulbi superior_ (see

eye), and near its termination communicates with the _Vena orbitalis

posterior_.]

β. The _Vena vertebralis_ (Fig. 160 _V_) arises posteriorly and runs

forwards over the transverse processes of the vertebrae, the _Musculi

intertransversarii_, and the _M. levat. scapulae_, to join the internal

jugular vein. In its course it receives branches (_Venae spinales_)

from the vertebral canal through the intervertebral foramina, and

cutaneous branches which accompany the corresponding arteries and

nerves through the dorsal lymphatic sac. The vein pulsates in

consequence of its connection with the anterior lymph-heart (Müller).

(2) The *subscapular vein* (_Vena subscapularis_), (Fig. 160 _S_). This

vein arises in the transverse abdominal muscles and the muscles of the

shoulder girdle; it unites with the internal jugular vein under the

shoulder-girdle.

*c.* The *subclavian vein* (_Vena subclavia_), (Fig. 159) passes

backwards from the anterior caval vein on the dorsal surface of the

coracoid bone and the clavicle, and then under cover of the _Portio

sternalis_ of the _M. pectoralis major_, to the internal border of the

_Portio abdominalis_ of the _M. pectoralis major_, where it is formed

by the union of the cutaneous vein and the brachial vein.

[Illustration: Fig. 161.

The anterior caval vein and its branches.

_a_ Innominate vein.

_c.m._ Cutaneous vein.

_c.s._ Anterior cava vein.

_i_ Internal jugular vein.

_i_′ External jugular vein.

_l_ Lingual vein.

_m_ Mandibular vein.

_s_ Brachial vein.

_s.c._ Subscapular vein.

]

(1) The cutaneous vein (_Vena cutanea magna_, Ecker; _Vena

musculo-cutanea_, Gruby), (Figs. 160, 161, and 162 _c.m._). This vein

arises by small branches in the head; these are:--

α. The _Vena nasalis_ (Virchow); it arises by an upper and a lower

branch from around the nostril, and courses backwards along the

superior maxillary bone to about the middle of the orbit, where it

joins the facial vein.

β. The _Vena orbitalis anterior_ (Virchow), (_Vena ophthalmica

anterior_, Ecker), (Fig. 160 _o_) commences at the Harderian gland from

a rich anastomosis, receives branches from the floor and anterior wall

of the orbit, and joins the facial vein with the _Vena nasalis_.

[Illustration: Fig. 162.

Course of the cutaneous vein as seen from the side.

_c.m._ Cutaneous vein.

_d_ M. depressor maxillae.

_i_ M. infraspinatus.

]

γ. The _Vena orbitalis posterior_ (Virchow), (_Vena ophthalmica

posterior_, Ecker), (Fig. 160 _o_′) arises on the posterior wall of

the orbit on the _M. pterygoideus_; it passes under the _Processus

zygomaticus_ to join the facial vein, and in its course receives the

_Vena ophthalmica_ (see eye), and communicates with the _Vena orbitalis

medialis_ and the internal jugular vein.

δ. The _Vena facialis_ (Fig. 160) is formed about the middle of the

orbit by the union of the _Vena nasalis_ and the _Vena orbitalis

anterior_; it courses backwards towards the angle of the jaw. On

reaching the _M. depressor maxillae_, about 4 mm. in front of the angle

of the jaw, it passes on to the skin. It receives in its course the

_Venae palpebrales_ (Fig. 162) and the _Vena orbitalis posterior_.

The musculo-cutaneous vein is the direct continuation of the _Vena

facialis_ after this has left the upper jaw; it immediately receives

one or more twigs (_Venae tympanicae_), (Fig. 162) from the tympanic

membrane and the tympanic cavity.

The vein then courses backwards in the outer wall of the lateral

lymph-sac as far as the middle of the trunk, where it passes, by

means of the _Septum abdominale_, to the outer border of the _Portio

abdominalis_ of the _M. pectoralis_; here the vein curves round to pass

forwards between the fibres of this muscle to its inner border where it

terminates by joining the brachial vein. In the whole of this lengthy

course the vein receives numerous branches from the neighbouring parts.

[Illustration: Fig. 163.

Veins in the region of the kidney.

_a_ Renal portal vein.

_a_′ Dorso-lumbar veins.

_abd_ Anterior abdominal vein.

_d_ Intestine.

_f_ Femoral vein.

_i_ Sciatic vein.

_N_ Kidney.

_o_ Right ovary, drawn to the left.

_o_′ Oviduct.

_ov_ Oviducal veins.

_p_ Mesenteric veins.

_r_ Renal veins.

_V_ Bladder.

_v.c.i._ Posterior vena cava.

_Vv_ Vesical veins.

]

(2) The *brachial vein* (_Vena brachialis_) is formed in the _Plica

cubiti_ by the union of two veins.

α. The *radial vein* (_Vena radialis_) arises on the dorsum of the hand

from a venous arch, the distal convexity of which receives the _Venae

digitales dorsales_: the vein courses superficially along the outer

side of the forearm to the elbow.

β. The *ulnar vein* (_Vena ulnaris_) arises from the veins in the

palm of the hand, and courses towards the elbow, lying deeply between

the muscles of the inner and outer surfaces of the forearm. In

the elbow the vein becomes superficial near the tendon of the _M.

sterno-cleidoradialis_, and joins the radial vein to form the brachial

vein.

The cutaneous veins of the fore-limb follow courses corresponding with

those of the arteries.

*2.* The *Posterior Caval Vein* (_Vena cava inferior_), (Fig. 159

_Cp_, 163 _v.c.i._). The posterior caval vein is a median vein

which, commencing between the kidneys, runs forwards to open into the

posterior end of the _Sinus venosus_. It receives the hepatic, renal,

and ovarian or spermatic veins.

*a.* The hepatic veins (_Venae hepaticae_), (Fig. 164 _v.h._) are

two in number, one from either side of the liver; they open into the

posterior caval vein near the _Sinus venosus_.

*b.* The *renal veins* (_Venae renales_ or _Venae revehentes_),

(Fig. 163 _r_) are four or five small vessels on each side; they pass

from the inner border of the kidney inwards and forwards, to form,

by their union, the posterior caval vein. The most anterior receive

branches from the fat-bodies (see kidney).

*c.* The *spermatic* or *ovarian veins* are four or five in number on

each side; they open into the posterior caval vein between the renal

veins.

*3.* The *Portal Systems*.

[There are, in the frog, two portal systems; the renal and the

hepatic. Each of these receives vessels which are derived, directly or

indirectly, from the veins of the hinder extremity.

The femoral vein on entering the pelvis divides to form two branches;

an iliac vein placed dorsally, and a pelvic vein placed ventrally; the

two pelvic veins converge to the middle line of ventral wall of the

abdomen, where they unite to form the anterior abdominal vein.]

*a.* The *renal portal vein* (_Vena renalis advehens princeps_),

(Fig. 163 _a_). This vein is formed by the union of the sciatic and the

iliac veins; it courses forwards to the outer border of the kidney, and

is distributed, by numerous branches, to that organ, chiefly on its

dorsal surface. It receives the following veins:--

(1) The *iliac vein* (_Vena iliaca externa_); this is the dorsal branch

formed by the bifurcation of the femoral vein.

(2) The *sciatic vein* (_Vena ischiadica_), (Figs. 163 and 165 _i_)

arises at the popliteal space, courses upwards in company with the

sciatic nerve between the _M. semitendinosus_ and the _M. biceps_, then

passes, still with the nerve, between the _M. vastus externus_ and the

_M. pyramidalis_ into the pelvis. It continues forwards to join the

iliac vein. At the root of the thigh this vein communicates with the

femoral vein by the _Ramus communicans iliacus_.

(3) The *dorso-lumbar vein* (_Vena dorso-lumbalis_), (Fig. 163 _a_′)

is a vein (or more usually several) arising by twigs in the dorsal and

outer walls of the abdomen; it joins the renal portal vein opposite the

kidney.

(4) The *oviducal veins* (Fig. 163 _ov_) are irregular veins from the

oviducts, which open either into the dorso-lumbar vein, or into the

renal portal vein.

*b.* The *hepatic portal system*.

The liver receives blood from two sources: (1) from the hind-limbs by

means of the anterior abdominal vein, and (2) from the alimentary canal

by the hepatic portal vein.

[Illustration: Fig. 164.

Veins of the liver.

_AA_ Auricles.

_aa_ Right and left branches of anterior abdominal vein.

_b_ Ramus descendens.

_c_ Vena bulbi (Vena cardiaca).

_D_ Intestine.

_G_ Gall-bladder.

_L.l._ Left lobe of liver.

_L.r._ Right lobe of liver.

_P_ Hepatic portal vein.

_P_′ Intestinal veins.

_P_″ Gastric veins.

_S_ Sinus venosus.

_V_ Ventricle.

_Va_ Anterior abdominal vein.

_V.c.i._ Posterior vena cava.

_v.h._ Hepatic veins.

]

The *anterior abdominal vein* (_Vena abdominalis_), (Figs. 159 _ab_,

164 _Va_) is formed, a little in front of the symphysis pubis, by

the union of the two pelvic veins. It courses forwards along the

middle line of the ventral body-wall until opposite the liver, where

it ascends and divides into three branches, of which two enter the

right and left lobes of the liver respectively. The third, the _Ramus

descendens_, joins the hepatic portal vein. The anterior abdominal vein

receives the following veins:--

α. The vesical vein (_Vena vesicalis_), (Fig. 163 _Vv_); a median vein

arising in the groove between the lobes of the bladder from lateral

branches, which extend outwards on the bladder. This vein communicates

above with other vesical twigs which open into the haemorrhoidal vein.

β. Parietal veins, which join the anterior abdominal vein in its course

along the ventral body-wall.

γ. A cardiac vein (_Vena cardiaca_), (Figs. 142, 159 _vc_, 164 _c_),

which joins the anterior abdominal vein just before its division (see

p. 222).

The *hepatic portal vein* (_Vena portarum_), (Figs. 159, 164 _P_) is

formed by the union of the gastric, intestinal, splenic, and one of

the haemorrhoidal veins. The large vein so constituted passes forwards

towards the liver, receives the _Ramus descendens_ from the anterior

abdominal vein, and then passes into the left lobe of the liver. Its

branches are the following:--

(1) The *gastric veins*. The stomach has three veins:--

α. [The _Vena coronaria ventriculi_, which receives blood from the

hinder end of the oesophagus and anterior portion of the stomach.

β and γ. Two other veins which form an _Arcus coronarius_ on the

concave surface of the stomach; the hinder receiving in addition the

small _Venae pancreaticae_ and the _Vena duodenalis_; the trunk so

formed receives the _Vena coronaria ventriculi_ (Hoffmann).

(2) The *intestinal veins*; the veins of that portion of the alimentary

canal between the middle of the duodenum and the anterior end of the

large intestine open into a common vein which joins the hepatic portal

vein.

(3) The *splenic vein* joins either the hepatic portal vein or one of

the intestinal veins.

(4) The *haemorrhoidal veins* are three in number; the anterior opens

into the splenic vein, the remaining two into the portal veins through

the agency of the intestinal veins.]

*4.* *The Veins of the Hinder Extremity.*

The hinder extremity has two venous trunks, the sciatic vein and the

femoral vein.

*a.* The *femoral vein* (_Vena femoralis_), (Fig. 165 _f_) is the

larger venous trunk of the hinder extremity. It arises in the popliteal

space as a direct continuation of the posterior tibial vein, courses

upwards in company with the sciatic artery and nerve, then leaves

these to wind forwards and outwards, and appears between the origins

of the _M. vastus externus_ and of the _M. rectus anterior_. The vein

here gives off the _Ramus communicans iliacus_ to the sciatic vein,

and turns forwards and downwards around the _M. glutaeus maximus_ to

reach the floor of the pelvis, near the femoral artery. The vein then

bifurcates to form the pelvic and iliac veins.

[Illustration: Fig. 165.

Veins of the hinder extremity, half schematic.

_c.i._ Vena communicans iliaca.

_f_ Femoral vein.

_i_ Sciatic vein.

_i_′ Vena cutanea cruris medialis.

_r.a._ M. rectus anterior.

_s.m._ M. semimembranosus.

_v.e._ M. vastus externus.

]

(1) The _Ramus communicans iliacus_ (Fig. 165 _c.i._) forms a

communication with the sciatic at the base of the thigh.

(2) The posterior tibial vein (_Vena tibialis postica_), of which the

femoral is a continuation, arises on the dorsal surface of the toes and

foot. It runs superficially on the _M. extensor longus digiti primi_

over the ankle-joint, courses upwards and winds round the outer surface

of the tibio-fibula to the flexor surface. It receives, in its course,

muscular branches, and at the knee a large cutaneous branch.

*b.* The *sciatic vein*: this vein has already been described (see

p. 247), but the vein from which it arises requires description.

The *anterior tibial* vein (_Vena tibialis antica_) arises on the

dorsum of the foot by twigs from the toes; it courses under the _M.

extensor longus digiti primi_, then upwards on the anterior surface

of the leg, and enters the canal in the tibio-fibula to appear on its

hinder surface. The vein then receives twigs from the _M. peroneus_,

the _M. tibialis posticus_, the _M. gastrocnemius_, and a large

cutaneous branch.

α. The _Vena cutanea cruris medialis_ (Fig. 165 _i_′) receives branches

from both above and below, and enters the popliteal space to join the

anterior tibial vein.

β. The anastomosis on the dorsum of the foot, from which the anterior

tibial vein arises, receives perforating twigs from the plantar surface.

The anatomy of the frog · 第 1 / 8 章