The elements of embryology
Sedgwick Adam | Balfour Francis M. (Francis Maitland), 1851-1882 | Foster M. (Michael) Sir, 1836-1907 | Heape Walter, 1855-1929 | Royal College of Physicians of Edinburgh
RSZ.O<b5 THE ELEMENTS OF EMBEYOLOGY. I THE ELEMENTS OF EMBRYOLOGY BY M. FOSTER, M.A., M.D., LL.D., F.R.S., FELLOW OF AND PRELECTOR IN PHYSIOLOGY IN TRINITY COLLEGE, CAMBRIDGE, AND THE LATE FRANCIS M. BALFOUR, M.A., LL.D., F.R.S., FELLOW OF TRINITY COLLEGE, CAMBRIDGE, AND PROFESSOR OF ANIMAL MORPHOLOGY IN THE UNIVERSITY. WALTER HE APE, DEMONSTRATOR IN THE MORPHOLOGICAL LABORATORY OF THE UNIVERSITY OF CAMBRIDGE. Honlon : MACMILLAN AND CO. 1883 [The Right of Translation is reserved.] ©ambrttige : PRINTED BY C. J. CLAY, M.A. & SON, AT THE UNIVERSITY PRESS. PREFACE TO THE SECOND EDITION. When this little work first appeared, it was put for- ward as a Part I, to be followed by other Parts. That plan was however soon abandoned. Nevertheless the volume seemed to have a place of its own ; and my dear lost friend undertook to prepare a second edition, in- tending to add some account of the development of the Mammal with a view of making the work an elementary introduction to vertebrate embryology more particularly suited for medical students. He was occu- pied with the task at the time of his sad death; and indeed a melancholy interest is attached to some of the sheets, by the fact that he had taken them to Switzer- land with him, on that fatal journey. All the first part up to p. 1G0 he had passed for press ; and he had further revised up to about p. 202. The whole of the rest of the volume has been under- VI PREFACE. taken by Mr Adam Sedgwick and Mr Walter Heape. They have attempted to carry out as far as possible what we believe to have been Balfour’s views, and trust that the public will judge leniently of their efforts to perform a difficult task. I have myself been able to do no more than offer general advice from time to time; and though it has not been thought advisable to change the title, the merits as well as the responsi- bilities of the latter part of the work must rest with them. M. FOSTER. Trinity College, Cambridge, March , 1883. TABLE OF CONTENTS. PART I. THE HISTORY OF THE CHICK. CHAPTER I. The Structure of the Hen’s Egg, and the Changes which take PLACE UP TO THE BEGINNING OF INCUBATION . pp. I — 24. The shell and shell-membrane, i — 3. The albumen, 3. The vitelline membrane, 4. The yolk, 5 — 7. Area opaca, 7. Area pellucida, 8. The structure of the blastoderm, 7 — 10. Recapitu- lation, 10. The ovarian ovum, 11 — 15. The descent of the ovum along the oviduct, 15 — 17. Impregnation, 17. Segmentation, 18 — 24. CHAPTER II. Brief Summary of the whole History of Incubation, pp. 25—47. The embryo is formed in the area pellucida, 25. The germinal layers, 25, 26. The extension of the blastoderm over the yolk, 26. The vascular area, 27. The head-fold, 27 — 36. The tail- fold, 37. The lateral folds, 37. The yolk-sac, 37. The alimentary canal, 39. The neural tube, 39, 40. The body-cavity, 41. The somatopleure, 41. The splanchnopleure, 42. The stalk of the yolk-sac, 42, 43. The amnion, 43 — 46. The allantois, 46, 47. Vlll TABLE OF CONTENTS. CHAPTER III. The Changes which take place during the First Day of Incu- bation pp. 48—76. Variations in the progress of development, 48, 49. The embryonic shield, 49. Formation of hypoblast, 51. The germinal wall, 52. The primitive streak, 52 — 54. Formation of primitive streak meso- blast, 54, 55. Hypoblastic mesoblast, 55. Primitive groove, 56, 57. The notochord, 59 — 62. The medullary groove, 62, 63. Amnion, 63. The changes taking place in the three layers, 63 — 66. The germinal wall, 65, 66. The increase of the head-fold, 66. The closure of the medullary canal, 66, 67. The cleavage of the mesoblast ; formation of spanchnopleure and somatopleure, 68. The vertebral and lateral plates, 69. The mesoblastic somites, 70. The sinus rhomboidalis, 71. The neurenteric passage, 71 — 74. Formation of the vascular area, 74) 75* Recapitulation, 75, 76. CHAPTER IV. The Changes which take place during the First Half of the Second Day . . . pp. 77 — 95. Increasing distinctness and prominence of embryo, 77. The first cerebral vesicle, 78, 79. The auditory pits, 81. Increase in number of mesoblastic somites, 81. The fore-gut, 82. The heart, 82 — 89. The vascular system, 89 — 94. Formation of blood-vessels, 92 — 94. The rudiment of the Wolffian duct, 94. Summary, 94, 95. CHAPTER V. The Changes which take place during the Second Half of the Second Day . . . pp. 96—108. Increasing prominence of the embryo; the tail-fold and lateral folds, 96. Continued closure of medullary canal, 96— 9S. The brain, 98—101. The optic vesicles, 98. The second aud third cerebral vesicles, 100. The cerebral hemispheres, 100. First appearance of cranial nerves, 100, 10 1. The notochord, 10 1. The cranial flexure, iot. The auditory vesicle, 101. Increase of curvature of heart, 101, TABLE OF CONTENTS. IX 102. Auricular appendages, 102. Vascular System, 102 — 106. Commencement of circulation, 102. The primitive aortae and first pair of aortic arches, 102, 103. The vitelline vessels and sinus ter- minalis, 103, 104. The course of the circulation, 105. The second and third pairs of aortic arches, 105, 106. The Wolffian duct and first appearance of Wolffian body, 106. The growth of the amnion, 107. The first appearance of the allantois, 107. Summary, 107, 108. CHAPTER VI. The Changes which take place during the Third Day, pp. 109 — 194. The diminution of the albumen, 109. The spreading of the opaque and vascular areas, 109, no. The vascular area, no — 113. The continued folding-in of the embryo, 113. The increase of the amnion, 113. The change in position of the embryo, 113 — 116. The curvature of the body, 116. The cranial flexure, 116, 117. The brain, 117 — 123. Growth of the vesicle of the cerebral hemispheres, 117. The lateral ventricles, 117. The vesicle of the 3rd ventricle or thalamencephalon, 117. The rudiment of the pineal gland, 117, 118. The infundibulum, 119. The stomodaeum, 119. The pituitary body, 119 — 121. Changes in the mid-brain, the corpora bigemina, crura cerebri and iter, 12 1. Changes in the hind-brain, the medulla, cerebellum, 4th ventricle, 121, 122. Changes in the neural canal, 122, 123. The cranial and spinal nerves, 123 — 132. The neural band, 123 — 126. The fifth, seventh, ninth and tenth cranial nerves, 126, 127. Later develop- ment of cranial nerves, 127 — 129. The spinal nerves, 129. The shifting of point of attachment of nerves, 131. Anterior roots, 131. The eye, 137 — 15 6. The first changes in the optic vesicles, 132, 133. The secondary optic vesicle and development of the lens, 134 — 137. The choroidal fissure, 137 — 140. The choroid, sclerotic and cornea, 140, 1 41. The further development of the optic vesicle, 141. The ora serrata, 142. The iris, 142. Pigment epithelium of choroid, 142. The ciliary processes, uvea, ciliary muscle and ligamentum pectinatum, 144. The histological changes in the retina, 144 — 146. Optic nerve, 146, 147. The choroid fissure, 147. The pecten, 148. The histo- logical changes in the lens, 149, 150. The vitreous humour, 150. The cornea, 150 — 153. The aqueous humour, 153. Summary of the development of the eye, 154, 155. The eyelids, 155. The lacrymal glands and duct, 155, 156. The organ of hearing, 156 — 161. Closure X TABLE OF CONTENTS. of the auditory involution, 157. The otic vesicle, 157. The mem- branous labyrinth, 158, 159. The osseous labyrinth, 159, 160. Com- parison of ear with eye, 160, 161. The organ of smell, 161, 162. The olfactory lobes and nerves, 162. The visceral arches and visceral clefts, 162 — 167. Superior maxillary, and fronto-nasal processes, 164, 165. Fate of first visceral cleft, 165, 1 66. The meatus audi- torius externus, 1 66. The tympanic membrane, 166. The Eustachian tube and tympanic cavity, 165, 166. The fenestra ovalis and rotunda, 166. The columella, 166, 167. The vascular system, 167 — 170. The aortic arches, 167. Changes in the heart, 167, 168. The venous system, 169, 170. The meatus venosus, cardinal veins and ductus Cuvieri, 169, 170. The alimentary canal, 171 — 185. Folding in of the splanchnopleure, tail-fold, 171, 172. The mesentery, 172, 173. (Esophagus and stomach, 173. The intestine, 174. The postanal gut, neurenteric canal and proctodaeum, 174 — 176. The lungs, 176 — 178. The liver, 178 — 181. The pancreas, 181. The thyroid body, 181, 182. The spleen, 182. The growth and blood-vessels of the allantois, 182 — 184. The mesollast, 185 — 193. The muscle-plates, 186 — 189. The intermediate cell-mass and Wolffian body, 189 — 193. A typical Wolffian tubule, 193. Change of position of Wolffian duct, 193. Summary, 193, 194. CHAPTER VII. The Changes which take place on the Fourth Day, pp. 195 — 231. Appearance on opening the egg, 195. Growth of amnion, 195, 196. The vitelline duct, 196. Increase of cranial flexure and tail-fold, 196 — 198. The first appearance of the limbs, 198. The growth of the brain, 200. The face, 202. Changes in the nasal pits, 202. The sto- modaeum and mouth, 202, 203. The cranial nerves, 203. Changes in the mesoblastic somites, 204 — 212. The membranous vertebral column, 205. The secondary segmentation of the vertebral column and formation of the permanent vertebrie, 205 — 207. Recapitulation, 207, 208. The changes in the notochord, 208—211. Ossification of vertebrae, 209, 210. The changes in the muscle plates, 211, 212. Wolffian body and duct, 212 — 214. The Mullerian duct, 214 — 218. The kidney and ureter, 218 — 220. The ovaries and testes, 220 — 223. Fate of the embryonic urinogenital organs, 223, 224. Changes in the arterial system, 224 — 226. Changes in the venous system ; veins of the liver, 226 — 229. Changes in the heart ; the ventricular septum, 229, 230. Summary, 230, 231. TABLE OF CONTENTS. XI CHAPTER VIII. The Changes which take place on the Fifth Day, pp. 232 — 274. Appearance on opening the egg, 232. The changes in the limbs, -33» 234- The pectoral and pelvic girdles ; the ribs and sternum, 234, 235. The development of the skull, 235 — 246. The cranium, 235. The parachordals and notochord, 237, 238. The trabeculae, 239 — 241. The sense capsules, 241, 242. Membrane and cartilage hones, 242. Skeleton of visceral arches, 242 — 245. Table of hones, 246. The changes in the face, 246 — 251. The posterior nares, 251. Changes in the spinal cord; its histological differentiation, 251 — 254. The central canal; and the posterior and anterior fissures, 254 — 256. Changes in the heart, 256 — 264. Septum in the bulbus and semilunar valves, 257 — 259. The cardiac valves, 262. The foramen ovale and Eustachian valve, 262 — 264. The pericardial and pleural cavities, 264 — 269. Histological differentiation and the fate of the three primary layers, 269—273. Summary, 273, 274. CHAPTER IX. From the Sixth Day to the End of Incubation, pp. 275 — 303. The appearance of distinct avian characters, 275. The foetal appendages during the 6th and 7th days, 276 — 278. During the 8th, 9th and 10th days, 278. From the nth to the 16th days, 278, 279. From the 16th day onwards, 279, 280. Changes in the general form of the embryo during the 6th and 7th days, 280—282. During the 8th — iothdays, 282. From the nth day onwards, 282. Feathers, 282. Nails, 283. Ossification, 283. Changes in the venous system before and after the commencement of pulmonary respiration, 283 — 289. Changes in the arterial system, the modification of the aortic arches, 289 297. Summary of the chief phases of the circulation, 297 — 303. Hatching, 303. Xll TABLE OF CONTENTS. PART II. THE HISTORY OF THE MAMMA- LIAN EMBRYO. Introduction, pp. 307, 308. CHAPTER X. General Development op the Embryo . . pp. 309 — 341. The ovarian ovum, 309, 310. The egg-membranes, 310. Ma- turation and impregnation, 310 — 312. Segmentation, 312 — 314. The blastodermic vesicle, 314 — 316. The formation of the layers, 316 — 320. The primitive streak and groove, 319, 320. The medullary groove, 320, 321. The mesoblast, 321 — 325. The notochord, 325, 326. The rudiment of the neurenteric canal, 326. Recapitulation, 326. The vascular area, 326. General growth of the embryo, 327 — 334. The human embryo, 335 — 341. Embryos of guinea-pig, etc. with so- called inversion of the layers, 341. CHAPTER XI. Embryonic Membranes and Yolk-Sac . . . pp. 342 — 364. The typical development of the embryonic membranes , 342 — 352. Vascular area of rabbit, 343 — 346. The yolk-sac or umbilical vesicle; amnion, 343. The subzonal membrane, 346. Attachment of blasto- dermic vesicle to uterine walls, 347. The formation of the chorion, 348. Mesoblast and blood-supply of the allantois, 348, 349. The placenta, 349, 350. The fate of the embiyonic membranes, 350 — 352. Deciduate and non-deciduate type of placenta, 352. Comparative history of the mammalian foetal membranes, 352 — 359. Foetal mem- branes of Monotremata and Marsupialia, 352. The discoidal pla- centa, 353, 354. The metadiscoidal placenta, 354 — 358. The zonary placenta, 358, 359. The diffuse and polycotyledonary placenta, 359. Comparative histology of the placenta, 359 — 363. Evolution of the placenta, 364. TABLE OF CONTENTS. xin CHAPTER XII. The Development of the Organs in Mammalia, pp. 365—422. The organs derived from the epiblast, 365 400. Hairs, 365. Glands, 3 66. The hind-brain, 367—370. The mid-brain, 370, 371. General development of fore-brain, 371. Thalamencephalon, 371 —37 6. Pituitary body, 372, 373. Pineal gland, 373—376- Cerebral hemispheres, 376 — 385. The olfactory lobes, 385. Histogenesis of brain, 385—387. The eyes, 387—390. Choroid slit, membrana capsulo-pupillaris and arteria centralis retinae, 389. The auditory organ, 390—397. Accessory auditory structures, 397— 399. The nasal organ, and organ of Jacobson, 399. Cranial and spinal nerves ; sym- pathetic system, 400. Organs derived from the mesoblast, 400 — 417. The vertebral column, 400, 401. The skull, 401. The visceral arches, 402. Man- dibular and hyoid arches ; malleus, incus, and stapes, 403 — 405. Bibs; sternum; pectoral and pelvic girdles, 405. Skeleton of the limbs, 406. Body-cavity ; pericardial, pleural cavities and dia- phragm, 406. The vascular system, 406 — 413. The heart, 406, 407. The ar- terial system, 407 — 409. The venous system, 409 — 413. The supra- renal bodies, 413, 414. The urinogenital organs, 414 — 417. Wolffian duct and body ; kidney ; ovary and testis, 414, 415. Genital cord, 415. Urinogenital sinus and external generative organs, 415 —417. Alimentary canal and its appendages , 417 — 422. Splanchnic mesoblast and mesentery, 419, 420. Stomodaeum, 420, 421. Hard and soft palate, 420, 421. Teeth, 421. Proctodaeum, 422. APPENDIX . . pp. 423 — 471. Incubators, 423 — 425. Hardening reagents, 425 — 428. Staining reagents, 428 — 432. Imbedding, 432 — 434. Cutting sections, 434, 435. Mounting sections, 436. Preservation of embryos as a whole, 436, 437- XIV TABLE OF CONTENTS. Practical directions for obtaining and studying chick embryos , 437 — 460. Examination of a 36 — 48 hours embryo, 437 — 444. Of an embryo of about 48 — 50 hours, 444 — 447. Of an embryo at the end of the 3rd day, 447 — 451. Of an embryo of the 4th day, 451 — 453. Of a blastoderm of 20 hours, 453 — 456. Of an unincubated blasto- derm, 457. Of the process of segmentation, 458. Of the later changes of the embryo, 459. Of the development of blood-vessels, 459, 460. Practical directions for obtaining and studying Mammalian em- bryos, 460 — 471. Animals and breeding, 460, 461. Examination and treatment of segmenting ova, 461 — 464. Of the blastodermic vesicle, 72 — 90 hours, 465. Of the blastodermic vesicle of 7 days, 465, 466. Of an 8 days embryo, 466 — 468. Of an embryo of 8 days 12 hours, 468, 469. Of the foetal membranes of an embryo of 14 days, 469, 470. Note A. Automatic microtome, 471. Note B. New method of mounting sections, 471. PART I. THE HISTORY OF THE CHICK. CHAPTER I. THE STRUCTURE OF THE HEN’S EGG, AND THE CHANGES WHICH TAKE PLACE UP TO THE BEGINNING OF IN- CUBATION. In a hens egg quite newly laid we meet with the following structures. Most external is the shell (Fig. 1, s.), composed of an organic basis, impregnated with calcic salts. It is sufficiently porous to allow of the interchange of gases between its interior and the exter- nal air, and thus the chemical processes of respiration, feeble at first, but gradually increasing in intensity, are carried on during the whole period of incubation. It is formed of two layers, both of which may contain pigment. The inner layer is by far the thickest, and is perforated by vertical canals which open freely on its inner aspect. Superficially these canals appear to be closed by the extremely thin outer layer. They are probably of some importance in facilitating the pene- tration of air through the shell. Lining the shell, is the shell-membrane, which is double, being made up of two layers : an outer thicker F. & B. 1 2 THE HEN’S EGG. [chap. (Fig. 1, s. m), and an inner thinner one (i. s. m.). Both 'of these layers consist of several laminae of felted fibres of various sizes, intermediate in nature between connec- tive and elastic fibres. Fig. l. Diagrammatic Section op an Unincubated Fowl’s Egg (modified from Allen Thomson). bl. blastoderm, w. y. white yolk. This consists of a central flask-shaped mass and a number of layers arranged con- centrically around this. y. y. yellow yolk. v. t. vitelline membrane, x. layer of more fluid albumen immediately surrounding the yolk. w. albumen consisting of alternate denser and more fluid layers, ch. 1. chalaza. a. ch. air- chamber at the broad end of the egg. This chamber is merely a space left between the two layers of the shell-mem- brane. i. s. m. internal layer of shell-membrane, s. m. external layer of shell-membrane, s. shell. I.] THE WHITE OF THE EGG. 3 Over the greater part of the egg the two layers of the shell-membrane remain permanently in close appo- sition ; but at the broad end they tend to separate, and thus to develope between them a space into which air finds its way. This air-chamber, as it is called, is not to be found in perfectly fresh eggs, but makes its appearance in eggs which have been kept for some time, whether incubated or not, and gradually increases in size, as the white of the egg shrinks in bulk from evaporation. Immediately beneath the shell-membrane is the white of the egg or albumen (Fig. 1, w.), which is, chemi- cally speaking, a mixture of various forms of proteid material, with fatty, extractive, and saline bodies. The outer part of the white, especially in eggs which are not perfectly fresh, is more fluid than that nearer the yolk. Its average composition may be taken as 12-0 p. c. proteid matter, 1'5 p. c. fat and extractives, 5 p. c. saline matter, chiefly sodic and potassic chlorides, with phosphates and sulphates, 86*0 p. c. water. The white of the egg when boiled shews in section alter- nate concentric layers of a transparent and of a finely granular opaque material. In the natural condition, the layers corre- sponding to these opaque layers are composed of more fluid albumen, while those corresponding to the transparent layers are less fluid, and consist of networks of fibres, containing fluid in their meshes. The innermost layer, however, immediately surrounding the yolk (Fig. 1, is of the more fluid finely granular kind. In eggs which have been hardened a spiral arrange- ment of the white may be observed, and it is possible to 1—2 THE HEN’S EGG. 4 [chap. tear off laminae in a spiral direction from left to right, from the broad to the narrow end of the egg. Two twisted cords called the chalazce (Fig. 1, ch. 1), composed of coiled membranous layers of denser albu- men, run from the two extremities of the egg to the opposite portions of the yolk. Their inner extremities expand and merge into a layer of denser albumen sur- rounding the fluid layer next the yolk. Their outer extremities are free, and do not quite reach the outer layer of the white. Thus they cannot serve to suspend the yolk, although they may help to keep it in position, by acting as elastic pads. The interior of each chalaza presents the appearance of a succession of opaque white knots ; hence the name chalazse (hailstones). The yolk is enclosed in the vitelline membrane (Fig. 1, v. t.), a transparent somewhat elastic membrane easily thrown into creases and wrinkles. It might almost be called structureless, but under a high power a fine fibrillation is visible, and a transverse section has a dotted or punctuated appearance ; it is probably there- fore composed of fibrils. Its affinities are with elastic connective tissue. The whole space within the vitelline membrane is occupied by the yolk. To the naked eye this appears tolerably uniform throughout, except at one particular point of its surface, at which may be seen, lying imme- diately under the vitelline membrane, a small white disc, about 4 mm. in diameter. This is the blastoderm, or cicatricula. A tolerably typical cicatricula in a fecundated egg will shew an outer white rim of some little breadth, and within that a circular transparent area, in the centre of THE WHITE YOLK. Ij 5 which, again, there is an opacity, varying in appearance, sometimes homogeneous, and sometimes dotted. The disc is always found to be uppermost whatever be the position of the egg, provided there is no restraint to the rotation of the yolk. The explanation of this is to be sought for in the lighter specific gravity of that portion of the yolk which is in the neighbourhood of the disc, and the phenomenon is not in any way due to the action of the chalazae. A section of the yolk of a hard-boiled egg will shew that it is not perfectly uniform throughout, but that there is a portion of it having the form of a flask, with a funnel-shaped neck, which, when the egg is boiled, does not become so solid as the rest of the yolk, but remains more or less fluid. The expanded neck of this flask-shaped space is situated immediately underneath the disc, while its bulbous enlargement is about in the middle of the yolk. We shall return to it directly. The great mass of the yolk is composed of what is known as the yellow yolk (Fig. 1, y. y.). This consists of spheres (Fig. 2, A.) of from 25/x to 100/r,1 in diameter filled with numerous minute highly refractive granules ; these spheres are very delicate and easily destroyed by crushing. When boiled or otherwise hardened in situ , they assume a polyhedral form, from mutual pressure. The granules they contain seem to be of an albuminous nature, as they are insoluble in ether or alcohol. Chemically speaking the yolk is characterized by the presence in large quantities of a proteid matter, having many affinities with globulin, and called vitellin. This exists in peculiar associa- 1 fx = ’001 mm. 6 THE HEN’S EGG. [CHAP. tion with the remarkable body Lecithin. (Compare Hoppe- Seyler, Hdh. Phys. Chem. Anal.) Other fatty bodies, colouring matters, extractives (and, according to Dareste, starch in small quantities), &c. are also present. Miescher (Hoppe-Seyler, Chem. Untersuch. p. 502) states that a considerable quantity of nuclein may be obtained from the yolk, probably from the spherules of the white yolk. Fig. 2. B - A A. Yellow yolk-sphere filled with fine granules. The outline of the sphere has been rendered too bold. B. White yolk-spheres and spherules of various sizes and pre- senting different appearances. (It is very difficult in a woodcut to give a satisfactory representation of these pe- culiar structures.) The yellow yolk, thus forming the great mass of the entire yolk, is clothed externally by a thin layer of a different material, known as the white yolk, which at the edge of the blastoderm passes underneath the disc, and becoming thicker at this spot forms, as it were, a bed on which the blastoderm rests. Immediately under the middle of the blastoderm this bed of white yolk is connected, by a narrow neck, with a central mass of similar material, lying in the middle of the yolk (Fig. 1, w. y.). When boiled, or otherwise hardened, the white yolk does not become so solid as the yellow yolk ; hence the appearances to be seen in sections of the hardened yolk. The upper expanded extremity of this neck of THE YELLOW YOLK. !•] 7 white yolk is generally known as the “ nucleus of Pander.” Concentric to the outer enveloping layer of white yolk there are within the yolk other inner layers of the same substance, which cause sections of the hardened yolk to appear to be composed of alternate concentric thicker laminae of darker (yellow) yolk, and thinner laminae of lighter (white) yolk (Fig. 1, w, y.). The microscopical characters of the white yolk elements are very different from those of the yellow yolk. It is composed of vesicles (Fig. 2, B.) for the most part smaller than those of the yellow yolk (4/x- — 75^), with a highly refractive body, often as small as 1/x, in the interior of each ; and also of larger spheres, each of which contains a number of spherules, similar to the smaller spheres. Another feature of the white yolk, according to His, is that in the region of the blastoderm it contains numerous large vacuoles filled with fluid; they are sufficiently large to be seen with the naked eye, but do not seem to be present in the ripe ovarian ovum. It is now necessary to return to the blastoderm. In this, as we have already said, the naked eye can distin- guish an opaque white rim surrounding a more trans- parent central area, in the middle of which again is a white spot of variable appearance. In an unfecundated cicatricula the white disc is simply marked with a number of irregular clear spaces, there being no proper division into a transparent centre and an opaque rim. The opaque rim is the commencement of what we shall henceforward speak of as the area opaca ; the central transparent portion is in the same way the 8 THE HEN’S EGG. [chap. beginning of the area pellucida. In the part corre- sponding to the area opaca the blastoderm rests imme- diately on the white yolk ; underneath the area pellu- cida is a shallow space containing a nearly clear fluid, to the presence of which the central transparency seems to be due. The white spot in the middle of the area pellucida appears to be the nucleus of Pander shining through. Vertical sections of the blastoderm shew that it is formed of two layers. The upper of these two layers is composed, see Fig. 3, ep, of a single row of cells, with their long axes arranged vertically, adhering together so as to form a distinct membrane, the edge of which rests upon the white yolk. After staining with silver nitrate, this membrane viewed from above shews a mosaic of uniform polygonal cells. Each cell is composed of granular protoplasm filled with highly refractive globules ; and in each an oval nu- cleus may be distinguished. They are of a nearly uniform size (about 9 fi) over the opaque and the pellucid areas. The under layer (Fig. 3, l ), is composed of cells which vary considerably in diameter; but even the smaller cells of this layer are larger than the cells of the upper layer. They are spherical, and so filled with granules and highly refractive globules, that a nucleus can rarely be seen in them : in the larger cells these globules are identical with the smaller white yolk spheres. The cells of this layer do not form a distinct mem- brane like the cells of the upper layer, but lie as a somewhat irregular network of cells between the upper layer and the bed of white yolk on which the blastoderm I.] THE BLASTODERM. rests. The lowest are generally the largest. The layer is thicker at the peri- phery than at the centre : and rests on a bed of white yolk, from which it is in parts separated by a more or less de- veloped cavity, containing probably fluid jmlk matter about to be absorbed. In the bed of white yolk nuclei are present, which are destined to become the nuclei of cells about to join the lower layer of the blastoderm. These nuclei are gene- rally more numerous in the neighbour- hood of the thickened periphery of the blastoderm than elsewhere. Amongst the lower layer cells are to be found Fig. 3. Section of a Blastoderm of a Fowl’s Egg AT THE COMMENCEMENT OF INCUBATION. The thin but complete upper layer ep com- posed of columnar cells rests on the in- complete lower layer l , composed of larger and more granular cells. The lower layer is thicker in some places than in others, and is especially thick at the periphery. The line below the under layer marks the upper surface of the white yolk. The larger so-called formative cells are seen at b, lying on the white yolk. The figure does not take in quite the whole breadth of the blastoderm ; but the reader must under- stand that both to the right hand and the left ep is continued farther than l, so that at the extreme edge it rests directly on the white yolk. 9 10 THE HEN’S EGG. [CHAP. peculiar large spherical bodies, which superficially re- semble the larger cells around them, and have been called formative cells. Their real nature is still very doubtful, and though some are no doubt true cells, others are perhaps only nutritive masses of yolk. The opacity of the peripheral part of the blastoderm is in a large measure due to the collection of the lower layer cells in this region, and the thickening, so caused, appears to be more pronounced for a small arc which subsequently constitutes the hinder border of the area pellucida. Over nearly the whole of the blastoderm the upper layer rests on the under layer. At the circumference however the upper layer stretches for a short distance beyond the under layer, and here consequently rests directly on the white yolk. To recapitulate : — In the normal unincubated hen’s egg we recognize the blastoderm, consisting of a com- plete upper layer of smaller nucleated granular cells and a more or less incomplete under layer of larger cells, filled with larger granules ; in these lower cells nuclei are rarely visible. The thin flat disc so formed rests, at the uppermost part of the entire yolk, on a bed of white yolk, and a peripheral thickening of the lower layer causes the appearance in the blastodermic disc of an area opaca and an area pellucida. The great mass of the entire yolk consists of the so-called yellow yolk composed of granular spheres. The white yolk is composed of smaller spheres of pecu- liar structure, and exists, in small part, as a thin coating around, and as thin concentric laininoe in the substance of the yellow yolk, but chiefly in the THE OVARIAN OVUM. 11 I-J form of a flask-shaped mass in the interior of the yolk, the upper somewhat expanded top of the neck of which forms the bed on which the blastoderm rests. The whole yolk is invested with the vitelline mem- brane, this again with the white ; and the whole is covered with two shell-membranes and a shell. Such an egg has however undergone most important changes while still within the body of the hen; and in order to understand the nature of the structures which have just been described, it will be necessary to trace briefly the history of the egg from the stage when it exists as a so-called ovarian ovum in the ovary of a hen up to the time when it is laid. In birds the left ovary alone is found in the adult ; and is attached by the mesovarium to the dorsal wall of the abdominal cavity, on the left side of the vertebral column. It consists of a mass of vascular stroma in which the ova are imbedded, is covered superficially by a layer of epithelium, continuous with the epithelial lining of the peritoneal cavity. The appearance of the ovary varies greatly according to the age of the indi- vidual. In the mature and sexually active females it is almost wholly formed of pedunculated and highly vascular capsules of various sizes, each containing a more or less developed ovum ; in the young animal however it is much more compact, owing to the absence of advanced ova. If one of the largest capsules of the ovary of a hen which is laying regularly be opened, it will be found to contain a nearly spherical (or more correctly, ellipsoidal with but slightly unequal axes) yellow body enclosed in a delicate membrane. This is the ovarian ovum or egg. 12 THE HEN’S EGG. [chap. Examined with care the ovum, which is tolerably uni- form in appearance, will be found to be marked at one spot (generally facing the stalk of the capsule and form- ing the pole of the shorter axis of the ovum) by a small disc differing in appearance from the rest of the ovum. This disc which is known as the germinal disc or discus Fig. 4. Section through the Germinal Disc of the ripe Ovarian Ovum of a Fowl while yet enclosed in its Capsule. a. Connective-tissue capsule of the ovum. b. follicular epithe- lium, at the surface of which nearest the ovum lies the vitelline membrane, c. granular material of the germinal disc, which becomes converted into the blastoderm. (This is not very well represented in the woodcut. In sections which have been hardened in chromic acid it consists of fine granules.) w. y, white yolk, which passes insensibly into the fine granular material of the disc, x, germinal vesicle enclosed in a distinct membrane, but shrivelled up by the action of the chromic acid, y, space originally completely filled up by the germinal vesicle, before the latter was shrivelled up by the action of the chromic acid. proligerus, consists of a lenticular mass of protoplasm (Fig. 4, c), imbedded in which is a globular or ellipsoidal body (Fig. 4, a?), about 310//, in diameter, called the germinal vesicle. This has a delicate wall, and its con- tents are clear and fluid in the fresh state, but become granular upon the addition of reagents. I-] THE OVARIAN OVUM. 13 The rest of the ovum is known as the yolk. This consists of two elements, the white yolk- and the yellow yolk-spheres, which are distributed respectively very much in the same way as in the laid egg, the yellow yolk forming the main mass of the ovum, and the white yolk being gathered underneath and around the disc (Fig. 4, w. y ), and also forming a flask-shaped mass in the interior. The delicate membrane surrounding the whole is the vitelline membrane. The youngest ova in the ovary of a fowl, in common with those of all other animals, present the characters of a simple cell. Such a cell is dia grammatically repre- sented in Fig. 5. It is seen to consist of a naked protoplasmic body containing in its interior a nucleus — the germinal vesi- cle— which in its turn envelopes a nucleolus — constituting what is known as the germinal spot. Such young ova are enclosed in a capsule of epithelium, named the follicle or follicular mem- brane, and are irregularly scat- tered in the stroma of the ovary. The difference between such an immature ovum and the ripe ovum just described is very great, but throughout its growth the ovum retains the characters of a cell, so that the mature ova- rian ovum, equally with the youngest ovum in the ovary, is a single cell. The most striking changes which takes place in the Fig. 5. Diagram op the Ovum. (From Gegen- baur.) a. Granular proto- plasm. b. Nucleus (ger- minal vesicle), c. Nu- cleolus (germinal spot). 14 THE HEN’S EGG. [chap. course of the maturation of the ovum concern the body of the cell rather than the germinal vesicle. As the body grows in size a number of granules make their appearance in its interior. These granules are formed by the inherent activity of the protoplasm, which is itself nourished, in a large measure at any rate, by the cells of the follicle. The outermost layer of the proto- plasm remains free from these granules. As the ovum grows older the granules become larger, first of all in the centre, and subsequently at the periphery, and take the form of white yolk-spherules. The greater part of them become at a later stage converted into yellow yolk-spheres, while a portion of them, situated in the position of the white yolk of the ripe ovum, retain their original characters. The germinal vesicle, which in the youngest ova is situated centrally or subcentrally, travels in the course of the growth of the ovum towards the periphery, and the protoplasm immediately surrounding it remains relatively free from yolk granules, and so constitutes the germinal disc. In the younger ova there is but a single germinal spot in the germinal vesicle, but as the ova enlarge several accessory germinal spots make then appearance, while in the ripe ovum it seems doubtful whether there is any longer a trace of a germinal spot. The cells of the follicular epithelium are at first arranged in a single row, but at a later stage become two or more rows deep : they undergo however a nearly complete atrophy in the ripe ovum. Around the follicular epithelium there is present a membrana propria, and in the later stages of the growth of the 15 I.] THE OVARIAN OVUM. ovum this is in its turn embraced by a highly vascular connective-tissue capsule. The youngest ova are, as has already been stated, quite naked. In ova of about 1*5 mm. the superficial layer of the ovum becomes converted into a radiately striated membrane called the zona radiata. At a later period a second membrane, placed between the zona radiata and the cells of the follicle, makes its appearance, but its mode of origin is still unknown. As the ovum approaches maturity the zona radiata disappears, and in the ripe ovum the second membrane, which has already been spoken of as the vitelline membrane, alone remains. From what has just been stated it follows that in an egg which has been laid the yolk alone constitutes the true ovum. The white and the shell are in fact accessory structures formed during the passage of the ovum down the oviduct. When the ovarian ovum is ripe and about to be discharged from the ovary, its capsule is clasped by the open infundibulum of the oviduct. The capsule then bursts, and the ovum escapes into the oviduct, its longer axis' corresponding with the long axis of the oviduct, the germinal disc therefore being to one side. In describing the changes which take place in the oviduct, it will be convenient, following the order pre- viously adopted, to treat first of all of the formation of the accessory parts of the egg. These are secreted by the glandular walls of the oviduct. This organ therefore requires some description. It may be said to consist of four parts : — 1st. The dilated infundibulum 16 THE HEN’S EGG. [chap. with an abdominal opening. 2nd. A long tubular portion — the oviduct proper — opening by a narrow neck or isthmus into the 3rd portion, which is much dilated, and has been called the uterus ; the 4th part is some- what narrow, and leads from the uterus into the cloaca. The whole of the mucous membrane lining the oviduct is largely ciliated. The accessory parts of the egg are entirely formed in the 2nd and 3rd portions. The layer of albumen which immediately surrounds the yolk is first de- posited ; the chalazae are next formed. Their spiral character and the less distinctly marked spiral arrange- ment of the whole albumen is brought about by the motion of the egg along the spiral ridges into which the interior of the second or tubular portion of the oviduct is thrown. The spirals of the two chalazae are in different directions. This is probably produced by their peripheral ends remaining fixed while the yolk to which their central ends are attached is caused to rotate by the contractions of the oviduct. During the formation of the chalazae the rest of the albumen is also deposited ; and finally the shell-membrane is formed in the narrow neck of the 2nd portion, by the fibrilla- tion of the most external layer of albumen. The egg passes through the 2nd portion in little more than 3 hours. In the 3rd portion the shell is formed. The mucous membrane of this part is raised into nume- rous flattened folds, like large villi, containing follicu- lar glands. From these a thick white fluid is poured out, which soon forms a kind of covering to the egg, in which the inorganic particles are deposited. In this portion of the oviduct the egg remains from 12 to 18 IMPREGNATION. 17 ’I-] hours, during which time the shell acquires its normal consistency. At the time of laying it is expelled from the uterus by violent muscular contractions, and passes with its narrow end downwards along the remainder of the oviduct, to reach the exterior. Impregnation. This process occurs in the upper portion of the oviduct; the spermatozoa being found actively moving in a fluid which is there contained. We have as yet, as far as the fowl is concerned, no direct observations concerning the changes preceding and following upon impregnation ; nor indeed concern- ing the actual nature of the act of impregnation. In other types however these processes have been followed with considerable care, and the result has been to shew that prior to impregnation a division of the ovum takes place into two very unequal parts. The smaller of these parts is known as the polar body, and plays no further part in the development. In the course of the division of the ovum into these two parts the germinal vesicle also divides, and one part of it enters the polar body, while a portion remains in the larger segment which continues to be called the ovum, and is there known as the female pronucleus. Im- pregnation has been found to consist essentially in the entrance of a single spermatozoon into the ovum, followed by the fusion of the two. The spermatozoon itself is to be regarded as a cell, the head of which corresponds to the nucleus. When the spermatozoon enters the ovum the substance forming its tail becomes mingled with the protoplasm of the latter, but the head enlarges and constitutes a distinct body called the male pronucleus, which travels towards and finally fuses with F. & b. 9 THE HEN’S EGG. 18 [chap. the female pronucleus to constitute the nucleus of the impregnated ovum. Segmentation. There follows upon the impregna- tion a remarkable process known as the segmentation. The process consists essentially in the division of the impregnated ovum by a series of successive segmenta- tions into a number of cells, of which the whole of the cells of the future animal are the direct descendants. In the majority of instances this process results in the division of the whole ovum into cells ; but in cases of ova where there is a large amount of food yolk, only that part of the ovum in which the protoplasm is but slightly loaded with food material, and which we have already described as the germinal disc, becomes so divided. The remainder of the ovum constitutes a food reservoir for the use of the developing embryo and is known as the food yolk. The segmentation in such ova, of which that of the fowl is one of the best known examples, is described as being partial or meroblastic1. In order to understand the process of segmentation in the fowl’s ovum it must be borne in mind that the germinal disc is not sharply separated from the re- mainder of the ovum, but that the two graduate insen- sibly into each other. The segmentation commences in the lower part of the oviduct, shortly before the shell has begun to be formed. Viewed from above, a furrow is seen to make its 1 For a fuller account of the relation between holoblastic and meroblastic segmentation the reader is referred to the treatise on Comparative Embryology by Balfour, Vol. i. chapter iii. Surface Views of the early Stages of the Segmentation in a Fowl’s Egg. (A and C after Coste.) A represents the earliest stage. The first furrow ( b ) has begun to make its appearance in the centre of the germinal disc, whose periphery is marked by the line a. In B, the first furrow is completed nearly across the disc, and a second similar furrow at right angles to the first has appeared. The disc thus becomes divided somewhat irregularly into quadrants by four (half) furrows. In a later stage (C) the meridian furrows b have increased in number, from four, as in B, to nine, and cross furrows have also made their appearance. The disc is thus cut up into small central (c) and larger peripheral (d) segments. Several new cross furrows are seen just beginning, as ex. gr. close to the end of the line of reference d. appearance, running across the germinal disc, though not for the whole breadth, and dividing it into two halves (Fig. 6, A). This primary furrow is succeeded by a second at right angles to itself. The surface thus becomes divided into four segments or quadrants (Fig. 6, B). 2—2 20 THE HEN’S EGG. [CHAP. Tlie second furrow cuts the first somewhat excen- trically. The first four furrows do not extend through the whole thickness of the germinal disc, and the four seg- ments marked out by them are not separated from the disc on their lower aspect. Each of these is again bisected by radiating furrows, and thus the number of segments is increased from four to eight (it may be seven or nine). The central portion of each segment is then, by a cross furrow, cut off from the peripheral portion, giving rise to the appearance of a number of central smaller segments, surrounded by more external elongated segments (Fig. 6, C ). The excentricity in the arrangement of the segments is moreover still preserved, the smaller segments being situated nearer one side of the germinal disc. The excentricity of the segmentation gives to the segmenting germinal disc a bilateral symmetry, but the relation between the axis of symmetry of the segmenting germinal disc and the long axis of the embryo is not known. Division of the segments now proceeds rapidly by means of furrows running in various directions. And it is important to note that the central segments divide more rapidly than the peripheral, and con- sequently become at once smaller and more numerous (Kg- 7)- Meanwhile sections of the hardened blastoderm teach us that segmentation is not confined to the sur- face, but extends through the mass of the blastoderm ; they shew us moreover that division takes place by means of not only vertical, but also horizontal furrows, i. e. furrows parallel to the surface of the disc (Fig. 8). I] SEGMENTATION. 21 Fig. 7. Surface View of the Germinal Disc of a Hen’s Egg DURING THE LATER STAGES OF SEGMENTATION. 