|
|
Line 1,267: |
Line 1,267: |
|
| |
|
| (490) J. K. Thacker. "Ventral fins of Ganoids." Trans, of the Connecticut | | (490) J. K. Thacker. "Ventral fins of Ganoids." Trans, of the Connecticut |
| Acad., Vol. iv. 1877. | | Acad., Vol. iv. 1877. |
| | |
| | |
| | |
| ==CHAPTER XXI. THE BODY CAVITY, THE VASCULAR SYSTEM, AND THE VASCULAR GLANDS==
| |
| | |
| The Body cavity.
| |
| | |
| IN the Ccelenterata no body cavity as distinct from the
| |
| alimentary cavity is present ; but in the remaining Invertebrata
| |
| the body cavity may (i) take the form of a wide space separating
| |
| the wall of the gut from the body wall, or (2) may be present in
| |
| a more or less reduced form as a number of serous spaces, or
| |
| (3) only be represented by irregular channels between the
| |
| muscular and connective-tissue cells filling up the interior of the
| |
| body. The body cavity, in whatever form it presents itself, is
| |
| probably filled with fluid, and the fluid in it may contain special
| |
| cellular elements. A well developed body cavity may coexist
| |
| with an independent system of serous spaces, as in the Vertebrata and the Echinodermata ; the perihaemal section of the
| |
| body cavity of the latter probably representing the system of
| |
| serous spaces.
| |
| | |
| In several of the types with a well developed body cavity it
| |
| has been established that this cavity originates in the embryo
| |
| from a pair of alimentary diverticula, and the cavities resulting
| |
| from the formation of these diverticula may remain distinct, the
| |
| adjacent walls of the two cavities fusing to form a dorsal and a
| |
| ventral mesentery.
| |
| | |
| It is fairly certain that some groups, e.g. the Tracheata, with
| |
| imperfectly developed body cavities are descended from ancestors
| |
| which were provided with well developed body cavities, but how
| |
| far this is universally the case cannot as yet be definitely
| |
| decided, and for additional information on this subject the
| |
| | |
| | |
| | |
| 624 CIIORDATA.
| |
| | |
| | |
| | |
| reader is referred to pp. 355 360 and to the literature there
| |
| referred to.
| |
| | |
| In the Chaetopoda and the Tracheata the body cavity arises
| |
| as a series of paired compartments in the somites of mesoblast
| |
| (fig. 350) which have at first a very restricted extension on the
| |
| ventral side of the body, but eventually extend dorsalwards and
| |
| vcntralwards till each cavity is a half circle investing the
| |
| alimentary tract ; on the dorsal side the walls separating the two
| |
| | |
| | |
| | |
| | |
| FIG. 350. LONGITUDINAL SECTION THROUGH AN EMBRYO OF AGELINA
| |
| LABYRINTHICA.
| |
| | |
| The section is taken slightly to one side of the middle line so as to shew the relation of the mesoblastic somites to the limbs. In the interior are seen the yolk
| |
| segments and their nuclei.
| |
| | |
| i 16. the segments ; pr.l. procephalic lobe ; do. dorsal integument.
| |
| | |
| half cavities usually remain as the dorsal mesentery, while
| |
| ventrally they are in most instances absorbed. The transverse
| |
| walls, separating the successive compartments of the body
| |
| cavity, generally become more or less perforated.
| |
| | |
| Chordata. In the Chordata the primitive body cavity is
| |
| cither directly formed from a pair of alimentary diverticula
| |
| (Cephalochorda) (fig. 3) or as a pair of spaces in the mesoblastic
| |
| plates of the two sides of the body (fig. 20).
| |
| | |
| As already explained (pp. 294 300) the walls of the dorsal
| |
| sections of the primitive body cavity soon become separated
| |
| from those of the ventral, and becoming segmented constitute
| |
| the muscle plates, while the cavity within them becomes
| |
| | |
| | |
| | |
| I
| |
| | |
| | |
| | |
| THE BODY CAVITY.
| |
| | |
| | |
| | |
| 625
| |
| | |
| | |
| | |
| the
| |
| | |
| | |
| | |
| obliterated : they are dealt with in a separate chapter. The
| |
| ventral part of the primitive cavity alone constitutes the
| |
| permanent body cavity.
| |
| | |
| The primitive body cavity in the lower Vertebrata is at first
| |
| continued forwards into the region of the head, but on the
| |
| formation of the visceral clefts the cephalic section of the body
| |
| cavity becomes divided into a series of separate compartments.
| |
| Subsequently these sections of the body cavity become obliterated ; and, since their walls give rise to muscles, they may
| |
| probably be looked upon as equivalent to the dorsal sections of
| |
| the body cavity in the trunk, and will be treated of in connection
| |
| with the muscular system.
| |
| | |
| As a result of its mode of origin the body cavity in
| |
| trunk is at first divided into two
| |
| lateral halves ; and part of the mesoblast lining it soon becomes distinguished as a special layer of epithelium, known as the peritoneal epithelium, of which the part bounding the
| |
| outer wall forms the somatic layer,
| |
| and that bounding the inner wall the
| |
| splanchnic layer. Between the two
| |
| splanchnic layers is placed the gut.
| |
| On the ventral side, in the region of
| |
| the permanent gut, the two halves
| |
| of the body cavity soon coalesce,
| |
| the septum between them becoming
| |
| absorbed, and the splanchnic layers
| |
| of epithelium of the two sides uniting
| |
| at the ventral side of the gut, and
| |
| the somatic layers at the median
| |
| ventral line of the body wall (fig.
| |
| | |
| | |
| | |
| | |
| In the lower Vertebrata the body
| |
| cavity is originally present even in
| |
| the post-anal region of the trunk, but
| |
| usually atrophies early, frequently
| |
| before the two halves coalesce.
| |
| | |
| On the dorsal side of the gut the
| |
| B. III.
| |
| | |
| | |
| | |
| FIG. 351. SECTION THROUGH
| |
| THE TRUNK OF A SCYLLIUM EMBRYO SLIGHTLY YOUNGER THAN
| |
| | |
| 28 F.
| |
| | |
| sp.c. spinal canal ; W. white
| |
| matter of spinal cord ; pr. posterior nerve-roots ; cA. notochord ;
| |
| x. sub-notochordal rod ; ao. aorta ;
| |
| nip. muscle-plate ; nip 1 , inner layer
| |
| of muscle-plate already converted
| |
| into muscles; Vr. rudiment of
| |
| vertebral body ; si. segmental
| |
| tube ; sd. segmental duct ; sp.v.
| |
| spiral valve ; v. subintestinal vein ;
| |
| p.o. primitive generative cells.
| |
| | |
| 40
| |
| | |
| | |
| | |
| 626 ABDOMINAL PORES.
| |
| | |
| | |
| | |
| two halves of the body cavity never coalesce, but eventually the
| |
| splanchnic layers of epithelium of the two sides, together with a
| |
| thin layer of interposed mesoblast, form a delicate membrane,
| |
| known as the mesentery, which suspends the gut from the dorsal
| |
| wall of the body (figs. 119 and 351). On the dorsal side the
| |
| epithelium lining of the body cavity is usually more columnar
| |
| than elsewhere (fig. 351), and its cells partly form a covering for
| |
| the generative organs, and partly give rise to the primitive
| |
| germinal cells. This part of the epithelium is often known as
| |
| the germinal epithelium.
| |
| | |
| Over the greater part of the body cavity the lining epithelium becomes in the adult intimately united with a layer of the
| |
| subjacent connective tissue, and constitutes with it a special
| |
| lining membrane for the body cavity, known as the peritoneal
| |
| membrane.
| |
| | |
| Abdominal pores. In the Cyclostomata, the majority of the Elasmobranchii, the Ganoidei, a few Teleostei, the Dipnoi, and some Sauropsida
| |
| (Chelonia and Crocodilia) the body cavity is in communication with the
| |
| exterior by a pair of pores, known as abdominal pores, the external
| |
| openings of which are usually situated in the cloaca 1 .
| |
| | |
| The ontogeny of these pores has as yet been but very slightly investigated.
| |
| In the Lamprey they are formed as apertures leading from the body cavity
| |
| into the excretory section of the primitive cloaca. This section would
| |
| appear from Scott's (No. 87) observations to be derived from part of the
| |
| hypoblastic cloacal section of the alimentary tract.
| |
| | |
| In all other cases they are formed in a region which appears to belong
| |
| to the epiblastic region of the cloaca ; and from my observations on Elasmobranchs it may be certainly concluded that they are formed there
| |
| in this group. They may appear as perforations (i) at the apices of
| |
| papilliform prolongations of the body cavity, or (2) at the ends of cloacal
| |
| pits directed from the exterior towards the body cavity, or (3) as simple
| |
| slit-like openings.
| |
| | |
| Considering the difference in development between the abdominal pores
| |
| of most types, and those of the Cyclostomata, it is open to doubt whether
| |
| these two types of pores are strictly homologous.
| |
| | |
| In the Cyclostomata they serve for the passage outwards of the generative products, and they also have this function in some of the few Teleostei
| |
| in which they are found ; and Gegenbaur and Bridge hold that the primitive
| |
| mode of exit of the generative products, prior to the development of the
| |
| Miillerian ducts, was probably by means of these pores. I have elsewhere
| |
| | |
| 1 For a full account of these structures the reader is referred to T. W. Bridge,
| |
| "Pori Abdominales of Vertebrata. " Journal of Anat. and Physiol. , Vol. XIV., 1879.
| |
| | |
| | |
| | |
| THE BODY CAVITY.
| |
| | |
| | |
| | |
| 627
| |
| | |
| | |
| | |
| | |
| suggested that the abdominal pores are perhaps remnants of the openings
| |
| of segmental tubes ; there does not however appear to be any definite
| |
| evidence in favour of this view, and it is more probable that they may have
| |
| arisen as simple perforations of the body wall.
| |
| | |
| Pericardial cavity, pleural cavities, and diaphragm.
| |
| | |
| In all Vertebrata the heart is at first
| |
| placed in the body cavity (fig. 353 A),
| |
| but the part of the body cavity containing it afterwards becomes separated as
| |
| a distinct cavity known as the pericardial cavity. In Elasmobranchii, Acipenser, etc. a passage is however left
| |
| between the pericardial cavity and the
| |
| body cavity ; and in the Lamprey a
| |
| separation between the two cavities does
| |
| not occur during the Ammoccete stage.
| |
| In Elasmobranchii the pericardial
| |
| cavity becomes established as a distinct
| |
| space in front of the body cavity in the
| |
| following way. When the two ductus
| |
| Cuvieri, leading transversely from the
| |
| sinus venosus to the cardinal veins, become developed, a horizontal septum,
| |
| shewn on the right side in fig. 352, is
| |
| formed to support them, stretching
| |
| across from the splanchnic to the somatic side of the body cavity, and
| |
| dividing the body cavity (fig. 352) in
| |
| this part into (i) a dorsal section formed
| |
| of a right and left division constituting
| |
| the true body cavity (pp), and (2) a
| |
| ventral part the pericardial cavity (pc).
| |
| The septum is at first of a very small
| |
| longitudinal extent, so that both in
| |
| front and behind it (fig. 352 on the left
| |
| side) the dorsal and ventral sections of the body cavity are in
| |
| free communication. The septum soon however becomes prolonged, and ceasing to be quite horizontal, is directed obliquely
| |
| upwards and forwards till it meets the dorsal wall of the body
| |
| | |
| 40 2
| |
| | |
| | |
| | |
| -ht
| |
| | |
| | |
| | |
| FIG. 352. SECTION
| |
| THROUGH THE TRUNK OF A
| |
| SCYLLIUM EMBRYO SLIGHTLY YOUNGER THAN 28 F.
| |
| | |
| The figure shews the separation of the body cavity from
| |
| the pericardial cavity by a
| |
| horizontal septum in which
| |
| runs the ductus Cuvieri ; on
| |
| the left side is seen the narrow
| |
| passage which remains connecting the two cavities.
| |
| | |
| sp.c. spinal canal ; w. white
| |
| matter of spinal cord ; pr.
| |
| commissure connecting the
| |
| posterior nerve-roots ; ch. notochord ; x. sub-notochordal
| |
| rod ; ao. aorta ; sv. sinus venosus ; cav. cardinal vein ; ht.
| |
| heart ; pp. body cavity ; pc.
| |
| pericardial cavity ; as. solid
| |
| oesophagus ; /. liver ; nip. muscle-plate.
| |
| | |
| | |
| | |
| 628 THE PERICARDIAL CAVITY.
| |
| | |
| Anteriorly all communication is thus early shut off between the
| |
| body cavity and the pericardial cavity, but the two cavities still
| |
| open freely into each other behind.
| |
| | |
| The front part of the body cavity, lying dorsal to the pericardial cavity, becomes gradually narrowed, and is wholly
| |
| obliterated long before the close of embryonic life, so that in
| |
| adult Elasmobranch Fishes there is no section of the body cavity
| |
| dorsal to the pericardial cavity. The septum dividing the body
| |
| cavity from the pericardial cavity is prolonged backwards, till it
| |
| meets the ventral wall of the body at the point where the liver
| |
| is attached by its ventral mesentery (falciform ligament). In
| |
| this way the pericardial cavity becomes completely shut off from
| |
| the body cavity, except, it would seem, for the narrow communications found in the adult. The origin of these communications
| |
| has not however been satisfactorily worked out.
| |
| | |
| The septum between the pericardial cavity and the body
| |
| cavity is attached on its dorsal aspect to the liver. It is at first
| |
| nearly horizontal, but gradually assumes a more vertical position,
| |
| and then, owing to the obliteration of the primitive anterior
| |
| part of the body cavity, appears to mark the front boundary of
| |
| the body cavity. The above description of the mode of formation of the pericardial cavity, and the explanation of its relations
| |
| to the body cavity, probably holds true for Fishes generally.
| |
| | |
| In the higher types the earlier changes are precisely the
| |
| same as those in Elasmobranch Fishes. The heart is at first
| |
| placed within the body cavity attached to the ventral wall of
| |
| the gut by a mesocardium (fig. 353 A). A horizontal septum is
| |
| then formed, in which the ductus Cuvieri are placed, dividing
| |
| the body cavity for a short distance into a dorsal (/./) and
| |
| ventral (p.c) section (fig. 353 B). In Birds and Mammals, and
| |
| probably also in Reptilia, the ventral and dorsal parts of the
| |
| body cavity are at first in free communication both in front of
| |
| and behind this septum. This is shewn for the Chick in
| |
| fig- 353 A an d B, which are sections of the same chick, A being
| |
| a little in front of B. The septum is soon continued forwards
| |
| so as completely to separate the ventral pericardial and the
| |
| dorsal body cavity in front, the pericardial cavity extending at
| |
| this period considerably further forwards than the body cavity.
| |
| | |
| Since the horizontal septum, by its mode of origin, is
| |
| | |
| | |
| | |
| THE BODY CAVITY.
| |
| | |
| | |
| | |
| 629
| |
| | |
| | |
| | |
| necessarily attached to the ventral side of the gut, the dorsal
| |
| part of the primitive body space is divided into two halves by a
| |
| median vertical septum formed of the gut and its mesentery
| |
| (fig- 353 B). Posteriorly the horizontal septum grows in a
| |
| slightly ventral direction along the under surface of the liver
| |
| (fig- 354)j till it meets the abdominal wall of the body at the
| |
| insertion of the falciform ligament, and thus completely shuts
| |
| off the pericardial cavity from the body cavity. The horizontal
| |
| septum forms, as is obvious from the above description, the
| |
| dorsal wall of the pericardial cavity 1 .
| |
| | |
| A. B.
| |
| | |
| | |
| | |
| | |
| | |
| FIG. 353. TRANSVERSE SECTIONS THROUGH A CHICK EMBRYO WITH TWENTYONE MESOBLASTIC SOMITES TO SHEW THE FORMATION OF THE PERICARDIAI,
| |
| CAVITY, A. BEING THE ANTERIOR SECTION.
| |
| | |
| p.p. body cavity; p.c. pericardial cavity; al. alimentary cavity ; au. auricle; v. ventricle; s.v. sinus venosus; d.c. ductus Cuvieri ; ao. aorta; nip. muscle-plate; me.
| |
| medullary cord.
| |
| | |
| With the complete separation of the pericardial cavity from
| |
| the body cavity, the first period in the development of these
| |
| parts is completed, and the relations of the body cavity to the
| |
| | |
| 1 Kolliker's account of this septum, which he calls the mesocardium laterale (No.
| |
| 298, p. 295), would seem to imply that in Mammals it is completed posteriorly even
| |
| before the formation of the liver. I doubt whether this takes place quite so early as
| |
| he implies, but have not yet determined its exact period by my own observations.
| |
| | |
| | |
| | |
| 630
| |
| | |
| | |
| | |
| THE PERICARDIAL CAVITY.
| |
| | |
| | |
| | |
| pericardial cavity become precisely those found in the embryos
| |
| of Elasmobranchii. The later changes are however very different. Whereas in Fishes the right and left sections of the body
| |
| cavity dorsal to the pericardial cavity soon atrophy, in the
| |
| higher types, in correlation with the relatively backward situation of the heart, they rapidly become larger, and receive the
| |
| lungs which soon sprout out from the throat.
| |
| | |
| The diverticula which form the lungs grow out into the
| |
| splanchnic mesoblast, in front of
| |
| the body cavity ; but as they
| |
| grow, they extend into the two
| |
| anterior compartments of the body
| |
| cavity, each attached by its mesentery to the mesentery of the
| |
| gut (fig. 354, lg). They soon moreover extend beyond the region of
| |
| the pericardium into the undivided
| |
| body cavity behind. This holds
| |
| not only for the embryos of the
| |
| Amphibia and Sauropsida, but
| |
| also for those of Mammalia.
| |
| | |
| To understand the further
| |
| | |
| rrianfrps in rhp nerirardial ravitv FlG> 354- SECTION THROUGH
| |
| | |
| pencaraiai cavity THECARDIACREGION OF AN EMBRYO
| |
| | |
| it is necessary to bear in mind its OF LACERTA MURALIS OF 9 MM. TO
| |
| | |
| , ,. ,, ,. . . SHEW THE MODE OF FORMATION OF
| |
| | |
| relations to the adjoining parts. THE PERICARDIAL CAVITY.
| |
| | |
| | |
| | |
| | |
| '-/it
| |
| | |
| | |
| | |
| It lies at this period completely
| |
| ventral to the two anterior pro
| |
| | |
| | |
| ht. heart ; pc. pericardial cavity ;
| |
| al. alimentary tract; lg. lung; /.
| |
| liver ; pp. body cavity ; md. open
| |
| longations of the body Cavity COn- end of Mullerian duct ; wd. Wolffian
| |
| . . duct; vc. vena cava inferior; ao.
| |
| | |
| taming the lungs (fig. 354). Its aorta; ch. notochord; me. medullary
| |
| | |
| dorsal wall is attached to the gut, cord>
| |
| | |
| and is continuous with the mesentery of the gut passing to the
| |
| dorsal abdominal wall, forming the posterior mediastinum of
| |
| human anatomy.
| |
| | |
| The changes which next ensue consist essentially in the
| |
| enlargement of the sections of the body cavity dorsal to the
| |
| pericardial cavity. This enlargement takes place partly by the
| |
| elongation of the posterior mediastinum, but still more by the
| |
| two divisions of the body cavity which contain the lungs
| |
| extending themselves ventrally round the outside of the peri
| |
| | |
| | |
| THE BODY CAVITY.
| |
| | |
| | |
| | |
| 631
| |
| | |
| | |
| | |
| cardial cavity. This process is illustrated by fig. 355, taken
| |
| from an embryo Rabbit. The two dorsal sections of the body
| |
| cavity (pl.p] finally extend so as completely to envelope the
| |
| pericardial cavity (pc\ remaining however separated from each
| |
| other below by a lamina extending from the ventral wall of the
| |
| pericardial cavity to the body wall, which forms the anterior
| |
| mediastinum of human anatomy.
| |
| | |
| By these changes the pericardial cavity is converted into a
| |
| closed bag, completely surrounded at its sides by the two lateral
| |
| halves of the body cavity, which were primitively placed
| |
| | |
| | |
| | |
| SJ3. C.
| |
| | |
| | |
| | |
| | |
| FIG. 355. SECTION THROUGH AN ADVANCED EMBRYO OF A RABBIT TO SHEW
| |
| HOW THE PERICARDIAL CAVITY BECOMES SURROUNDED BY THE PLEURAL
| |
| CAVITIES.
| |
| | |
| ht. heart; pc. pericardial cavity; //./ pleural cavity; Ig. lung; al. alimentary
| |
| tract; ao. dorsal aorta; ch. notochord; rp. rib; st. sternum; sp.c. spinal cord.
| |
| | |
| dorsally to it. These two sections of the body cavity, which in
| |
| Amphibia and Sauropsida remain in free communication with
| |
| the undivided peritoneal cavity behind, may, from the fact of
| |
| their containing the lungs, be called the pleural cavities.
| |
| | |
| In Mammalia a further change takes place, in that, by the
| |
| formation of a vertical partition across the body cavity, known
| |
| as the diaphragm, the pleural cavities, containing the lungs,
| |
| | |
| | |
| | |
| 632 THE VASCULAR SYSTEM.
| |
| | |
| become isolated from the remainder of the body or peritoneal
| |
| cavity. As shewn by their development the so-called pleurae or
| |
| pleural sacks are simply the peritoneal linings of the anterior
| |
| divisions of the body cavity, shut off from the remainder of
| |
| the body cavity by the diaphragm.
| |
| | |
| The exact mode of formation of the diaphragm is not fully
| |
| made out ; the account of it recently given by Cadiat (No. 491)
| |
| not being in my opinion completely satisfactory.
| |
| | |
| BIBLIOGRAPHY.
| |
| | |
| (491) M. Cadiat. "Du developpement de la partie cephalothoracique de 1'embryon, de la formation du diaphragme, des pleures, du pericarde, du pharynx et de
| |
| 1'cesophage." Journal de F Anatomic et de la Physiologic, Vol. xiv. 1878.
| |
| | |
| | |
| | |
| Vascular System.
| |
| | |
| The actual observations bearing on the origin of the vascular
| |
| system, using the term to include the lymphatic system, are
| |
| very scanty. It seems probable, mainly it must be admitted on
| |
| d priori grounds, that vascular and lymphatic systems have
| |
| originated from the conversion of indefinite spaces, primitively
| |
| situated in the general connective tissue, into definite channels.
| |
| It is quite certain that vascular systems have arisen independently in many types ; a very striking case of the kind being
| |
| the development in certain parasitic Copepoda of a closed
| |
| system of vessels with a red non-corpusculated blood (E. van
| |
| Beneden, Heider), not found in any other Crustacea. Parts of
| |
| vascular systems appear to have arisen in some cases by a
| |
| canalization of cells.
| |
| | |
| The blood systems may either be closed or communicate
| |
| with the body cavity. In cases where the primitive body cavity
| |
| is atrophied or partially broken up into separate compartments
| |
| (Insecta, Mollusca, Discophora, etc.) a free communication
| |
| between the vascular system and the body cavity is usually
| |
| present ; but in these cases the communication is no doubt
| |
| secondary. On the whole it would seem probable that the
| |
| vascular system has in most instances arisen independently of
| |
| the body cavity, at least in types where the body cavity is
| |
| | |
| | |
| | |
| THE VASCULAR SYSTEM. 633
| |
| | |
| present in a well-developed condition. As pointed out by the
| |
| Hertwigs, a vascular system is always absent where there is not
| |
| a considerable development of connective tissue.
| |
| | |
| As to the ontogeny of the vascular channels there is still much to be
| |
| made out both in Vertebrates and Invertebrates.
| |
| | |
| The smaller channels often rise by a canalization of cells. This process
| |
| has been satisfactorily studied by Lankester in the Leech 1 , and may easily
| |
| be observed in the blastoderm of the Chick or in the epiploon of a newlyborn Rabbit (Schafer, Ranvier). In either case the vessels arise from a network of cells, the superficial protoplasm and part of the nuclei giving rise
| |
| to the walls, and the blood-corpuscles being derived either from nucleated
| |
| masses set free within the vessels (the Chick) or from blood-corpuscles
| |
| directly differentiated in the axes of the cells (Mammals).
| |
| | |
| Larger vessels would seem to be formed from solid cords of cells, the
| |
| central cells becoming converted into the corpuscles, and the peripheral cells
| |
| constituting the walls. This mode of formation has been observed by
| |
| myself in the case of the Spider's heart, and by other observers in other
| |
| Invertebrata. In the Vertebrata a more or less similar mode of formation
| |
| appears to hold good for the larger vessels, but further investigations are
| |
| still required on this subject. Gotte finds that in the Frog the larger vessels
| |
| are formed as longitudinal spaces, and that the walls are derived from the
| |
| indifferent cells bounding these spaces, which become flattened and united
| |
| into a continuous layer.
| |
| | |
| The early formation of vessels in the Vertebrata takes place in the
| |
| splanchnic mesoblast ; but this appears due to the fact that the circulation
| |
| is at first mainly confined to the vitelline region, which is covered by
| |
| splanchnic mesoblast.
| |
| | |
| The Heart.
| |
| | |
| The heart is essentially formed as a tubular cavity in the
| |
| splanchnic mesoblast, on the ventral side of the throat, immediately behind the region of the visceral clefts. The walls of this
| |
| cavity are formed of two layers, an outer thicker layer, which has
| |
| at first only the form of a half tube, being incomplete on its
| |
| dorsal side; and an inner lamina formed of delicate flattened
| |
| cells. The latter is the epithelioid lining of the heart, and the
| |
| cavity it contains the true cavity of the heart. The outer layer
| |
| gives rise to the muscular wall and peritoneal covering of the
| |
| heart. Though at first it has only the form of a half tube (fig.
| |
| | |
| 1 "Connective and vasifactive tissues of the Leech." Quart. J. of Micr. Science,
| |
| Vol. XX. 1880.
| |
| | |
| | |
| | |
| 634
| |
| | |
| | |
| | |
| THE HEART.
| |
| | |
| | |
| | |
| 356), it soon becomes folded in on the dorsal side so as to form
| |
| for the heart a complete muscular wall. Its two sides, after thus
| |
| meeting to complete the tube of
| |
| the heart, remain at first continuous
| |
| with the splanchnic mesoblast surrounding the throat, and form a provisional mesentery the mesocardium which attaches the heart to
| |
| the ventral wall of the throat. The
| |
| superficial stratum of the wall of
| |
| the heart differentiates itself as the
| |
| peritoneal covering. The inner epithelioid tube takes its origin at the
| |
| time when the general cavity of the
| |
| heart is being formed by the separation of the splanchnicmesoblastfrom
| |
| the hypoblast. During this process
| |
| (fig. 357) a layer of mesoblast remains close to the hypoblast, but connected with the main mass
| |
| | |
| | |
| | |
| | |
| FIG. 356. SECTION THROUGH
| |
| THE DEVELOPING HEART OF AN
| |
| EMBRYO OF AN ELASMOBRANCH
| |
| (Pristiurus).
| |
| | |
| al. alimentary tract ; sp. splanchnic mesoblast ; so. somatic mesoblast ; ht. heart.
| |
| | |
| | |
| | |
| | |
| FIG. 357. TRANSVERSE SECTION THROUGH THE POSTERIOR PART OF THE
| |
| HEAD OF AN EMBRYO CHICK OF THIRTY HOURS.
| |
| | |
| hb. hind-brain; vg. vagus nerve; ep. epiblast; ch. notochorcl ; x. thickening of
| |
| hypoblast (possibly a rudiment of the sub-notochordal rod) ; al. throat; ht. heart;
| |
| //. body cavity; so. somatic mesoblast; sf. splanchnic mesoblast; Ay. hypoblast.
| |
| | |
| | |
| | |
| THE VASCULAR SYSTEM.
| |
| | |
| | |
| | |
| 635
| |
| | |
| | |
| | |
| of the mesoblast by protoplasmic processes. A second layer
| |
| next becomes split from the splanchnic mesoblast, connected
| |
| with the first layer by the above-mentioned protoplasmic
| |
| processes. These two layers form together the epithelioid lining
| |
| of the heart ; between them is the cavity of the heart, which soon
| |
| loses the protoplasmic trabeculae which at first traverse it. The
| |
| cavity of the heart may thus be described as being formed by a
| |
| hollowing out of the splanchnic mesoblast, and resembles in its
| |
| mode of origin that of other large vascular trunks.
| |
| | |
| The above description applies only to the development of
| |
| the heart in those types in which it is formed at a period after
| |
| the throat has become a closed tube (Elasmobranchii, Amphibia,
| |
| Cyclostomata, Ganoids (?)). In a number of other cases, in
| |
| which the heart is formed before the conversion of the throat
| |
| into a closed tube, of which the most notable is that of Mammals
| |
| (Hensen, Gotte, Kolliker), the heart arises as two independent
| |
| | |
| A.
| |
| | |
| | |
| | |
| | |
| B.
| |
| | |
| | |
| | |
| mes fir
| |
| | |
| | |
| | |
| | |
| FIG. 358. TRANSVERSE SECTION THROUGH THE HEAD OF A RABBIT OF THE
| |
| | |
| SAME AGE AS FIG. 144 B. (From Kolliker.)
| |
| B is a more highly magnified representation of part of A.
| |
| | |
| rf. medullary groove; mp. medullary plate; riv. medullary fold; h. epiblast ;
| |
| dd. hypoblast; dd' . notochordal thickening of hypoblast; sp. undivided mesoblast;
| |
| ^.somatic mesoblast; dfp. splanchnic mesoblast; ph. pericardial section of body
| |
| cavity; ahh. muscular wall of heart; ihh. epithelioid layer of heart; vies, lateral
| |
| undivided mesoblast ; sw. part of the hypoblast which will form the ventral wall of
| |
| the pharynx.
| |
| | |
| | |
| | |
| 636
| |
| | |
| | |
| | |
| THE HEART.
| |
| | |
| | |
| | |
| tubes (fig. 358), which eventually coalesce into an unpaired
| |
| structure.
| |
| | |
| In Mammals the two tubes out of which the heart is formed appear at
| |
| the sides of the cephalic plates, opposite the region of the mid- and hindbrain (fig. 358). They arise at a time when the lateral folds which form
| |
| the ventral wall of the throat are only just becoming visible. Each half of
| |
| the heart originates in the same way as the whole heart in Elasmobranchii,
| |
| etc. ; and the layer of the splanchnic mesoblast, which forms the muscular
| |
| wall for each part (ahh), has at first the form of a half tube open below to
| |
| the hypoblast.
| |
| | |
| On the formation of the lateral folds of the splanchnic walls, the two
| |
| halves of the heart become carried inwards and downwards, and eventually
| |
| | |
| | |
| | |
| | |
| FlG. 359. TWO DIAGRAMMATIC SECTIONS THROUGH THE REGION OF THE
| |
| HIND-BRAIN OF AN EMBRYO CHICK OF ABOUT 36 HOURS ILLUSTRATING THE
| |
| FORMATION OF THE HEART.
| |
| | |
| fib. hind-brain ; nc. notochord ; E. epiblast ; so. somatopleure ; sp. splanchnopleure ; d. alimentary tract ; hy. hypoblast ; hs. heart ; of. vitelline veins.
| |
| | |
| | |
| | |
| THE VASCULAR SYSTEM.
| |
| | |
| | |
| | |
| 637
| |
| | |
| | |
| | |
| meet on the ventral side of the throat. For a short time they here remain
| |
| distinct, but soon coalesce into a single tube.
| |
| | |
| In Birds, as in Mammals, the heart makes its appearance as two tubes,
| |
| but arises at a period when the formation of the throat is very much more
| |
| advanced than in the case of Mammals. The heart arises immediately
| |
| behind the point up to which the ventral wall of the throat is established
| |
| and thus has at first a A -shaped form. At the apex of the A , which forms
| |
| the anterior end of the heart, the two halves are in contact (fig. 357),
| |
| though they have not coalesced; while behind they diverge to be continued
| |
| as the vitelline veins. As the folding in of the throat is continued backwards the two limbs of the heart are brought together and soon coalesce
| |
| from before backwards into a single structure. Fig. 359 A and B shews the
| |
| heart during this process. The two halves have coalesced anteriorly (A)
| |
| but are still widely separated behind (B). In Teleostei the heart is formed
| |
| as in Birds and Mammals by the coalescence of two tubes, and it arises
| |
| before the formation of the throat.
| |
| | |
| The fact that the heart arises in so many instances as a
| |
| double tube might lead to the supposition that the ancestral
| |
| Vertebrate had two tubes in the place of the present unpaired
| |
| heart.
| |
| | |
| The following considerations appear to me to prove that this
| |
| conclusion cannot be accepted. If the folding in of the splanchnopleure to form the throat were deferred relatively to the
| |
| formation of the heart, it is clear that a modification in the
| |
| development of the heart would occur, in that the two halves of
| |
| the heart would necessarily be formed widely apart, and only
| |
| eventually united on the folding in of the wall of the throat. It
| |
| is therefore possible to explain the double formation of the heart
| |
| without having recourse to the above hypothesis of an ancestral
| |
| Vertebrate with two hearts. If the explanation just suggested
| |
| is the true one the heart should only be formed as two tubes
| |
| when it arises prior to the formation of the throat, and as a single
| |
| tube when formed after the formation of the throat. Since this
| |
| is invariably found to be so, it may be safely concluded that the
| |
| formation of the heart as two cavities is a secondary mode of
| |
| development, which has been brought about by variations in the
| |
| period of the closing in of the wall of the throat.
| |
| | |
| The heart arises continuously with the sinus venosus, which in
| |
| the Amniotic Vertebrata is directly continued into the vitelline
| |
| veins. Though at first it ends blindly in front, it is very soon
| |
| connected with the foremost aortic arches.
| |
| | |
| | |
| | |
| 638 THE HEART.
| |
| | |
| | |
| | |
| The simple tubular heart, connected as above described, grows
| |
| more rapidly than the chamber in which it is contained, and is
| |
| soon doubled upon itself, acquiring in this way an S-shaped
| |
| curvature, the posterior portion being placed dorsally, and the
| |
| anterior ventrally. A constriction soon appears between the
| |
| dorsal and ventral portions.
| |
| | |
| The dorsal section becomes partially divided off behind from
| |
| the sinus venosus, and constitutes the relatively thin-walled
| |
| auricular section of the heart; while the ventral portion, after
| |
| becoming distinct anteriorly from a portion continued forwards
| |
| from it to the origin of the branchial arteries, which may be called
| |
| the truncus arteriosus, acquires very thick spongy muscular
| |
| walls, and becomes the ventricular division of the heart.
| |
| | |
| The further changes in the heart are but slight in the case of the Pisces.
| |
| A pair of simple membranous valves becomes established at the auriculoventricular orifice, and further changes take place in the truncus arteriosus.
| |
| This part becomes divided in Elasmobranchii, Ganoidei, and Dipnoi into a
| |
| posterior section, called the conus arteriosus, provided with a series of
| |
| transverse rows of valves, and an anterior section, called the bulb us
| |
| arteriosus, not provided with valves, and leading into the branchial
| |
| arteries. In most Teleostei (except Butirinus and a few other forms) the
| |
| conus arteriosus is all but obliterated, and the anterior row of its valves
| |
| alone preserved ; and the bulbus is very much enlarged 1 .
| |
| | |
| In the Dipnoi important changes in the heart are effected, as compared
| |
| with other Fishes, by the development of true lungs. Both the auricular
| |
| and ventricular chamber may be imperfectly divided into two, and in the
| |
| conus a partial longitudinal septum is developed in connection with a
| |
| longitudinal row of valves 2 .
| |
| | |
| In Amphibia the heart is in many respects similar to that of the Dipnoi.
| |
| Its curvature is rather that of a screw than of a simple S. The truncus
| |
| arteriosus lies to the left, and is continued into the ventricle which lies
| |
| ventrally and more to the right, and this again into the dorsally placed
| |
| auricular section.
| |
| | |
| After the heart has reached the piscine stage, the auricular section
| |
| (Bombinator) becomes prolonged into a right and left auricular appendage^
| |
| A septum next grows from the roof of the auricular portion of the heart
| |
| | |
| 1 Vide Gegenbaur, "Zur vergleich. Anat. d. Herzens." Jenaische Zeit., Vol. n.
| |
| 1866, and for recent important observations, J. E. V. Boas, "Ueb. Herz u. Arterienbogenbei Ceratodenu. Protopterus," and " Ueber d. Conus arter. b. Butirinus, etc.,"
| |
| Morphol. Jahrb., Vol. VI. 1880.
| |
| | |
| 2 Boas holds that the longitudinal septum is formed by the coalescence of a row of
| |
| longitudinal valves, but this is opposed to Lankester's statements, "On the hearts of
| |
| Ceratodus, Protopterus and Chimaera, etc. Zool. Trans. Vol. x. 1879.
| |
| | |
| | |
| | |
| THE VASCULAR SYSTEM. 639
| |
| | |
| | |
| | |
| obliquely backwards and towards the left, and divides it in two chambers ;
| |
| the right one of which remains continuous with the sinus venosus, while
| |
| the left one is completely shut off from the sinus, though it soon enters
| |
| into communication with the newly established pulmonary veins. The
| |
| truncus arteriosus 1 is divided into a posterior conus arteriosus (pylangium)
| |
| and an anterior bulbus (synangium). The former is provided with a
| |
| proximal row of valves at its ventricular end, and a distal row at its anterior
| |
| end near the bulbus. It is also provided with a longitudinal septum, which
| |
| is no doubt homologous with the septum in the conus arteriosus of the
| |
| Dipnoi. The bulbus is well developed in many Urodela, but hardly exists
| |
| in the Anura.
| |
| | |
| In the Amniota further changes take place in the heart,
| |
| resulting in the abortion of the distal rows of valves of the conus
| |
| arteriosus 2 , and in the splitting up of the whole truncus arteriosus
| |
| into three vessels in Reptilia, and two in Birds and Mammals,
| |
| each opening into the ventricular section of the heart, and
| |
| provided with a special set of valves at its commencement. In
| |
| Birds and Mammals the ventricle becomes moreover completely
| |
| divided into two chambers, each communicating with one of the
| |
| divisions of the primitive truncus, known in the higher types
| |
| as the systemic and pulmonary aortae. The character of the
| |
| development of the heart in the Amniota will be best understood
| |
| from a description of what takes place in the Chick.
| |
| | |
| In Birds the originally straight heart (fig. 109) soon becomes doubled up
| |
| upon itself. The ventricular portion becomes placed on the ventral and
| |
| right side, while the auricular section is dorsal and to the left. The two
| |
| parts are separated from each other by a slight constriction known as the
| |
| canalis auricularis. Anteriorly the ventricular cavity is continued into the
| |
| truncus, and the venous or auricular portion of the heart is similarly connected behind with the sinus venosus. The auricular appendages grow out
| |
| from the auricle at a very early period. The general appearance of the
| |
| heart, as seen from the ventral side on the fourth day, is shewn in fig. 360.
| |
| Although the external divisions of the heart are well marked even before
| |
| this stage, it is not till the end of the third day that the internal partitions
| |
| become apparent ; and, contrary to what might have been anticipated from
| |
| the evolution of these parts in the lower types, the ventricular septum is the
| |
| first to be established.
| |
| | |
| 1 For a good description of the adult heart vide Huxley, Article "Amphibia," in
| |
| the Encyclopedia Britannic a.
| |
| | |
| 2 It is just possible that the reverse may be true, vide note on p. 640. If however,
| |
| as is most probable, the statement in the text is correct, the valves at the mouth of
| |
| the ventricle in Teleostei are not homologous with those of the Amniota ; the former
| |
| being the distal rov/ of the valves of the conus, the latter the proximal.
| |
| | |
| | |
| | |
| | |
| 640 THE HEART OF AVES.
| |
| | |
| It commences on the third day as a crescentic ridge or fold springing
| |
| from the convex or ventral side of the rounded ventricular portion of the
| |
| heart, and on the fourth day grows rapidly across the ventricular cavity
| |
| towards the concave or dorsal side. It thus forms an incomplete longitudinal partition, extending from the canalis auricularis to the commencement
| |
| of the truncus arteriosus, and dividing the twisted ventricular tube into
| |
| two somewhat curved canals, one more
| |
| to the left and above, the other to
| |
| | |
| the right and below. These commu- A ^) ) CA
| |
| | |
| nicate with each other, above the free
| |
| edge of the partition, along its whole
| |
| length.
| |
| | |
| Externally the ventricular portion
| |
| as yet shews no division into two parts.
| |
| | |
| By the fifth day the venous end of
| |
| the heart, though still lying somewhat
| |
| to the left and above, is placed as far FIG. 360. HEART OF A CHICK ON
| |
| | |
| forwards as the arterial end, the whole THE FOURTH DAY OF INCUBATION
| |
| | |
| VIEWED FROM THE VENTRAL SURFACE.
| |
| | |
| organ appearing to be drawn together.
| |
| | |
| The ventricular septum is complete. L ?.- lef t a , uricular appendage; C.A.
| |
| | |
| , e .. , . , , canahs auricularis ; v. ventricle ; b. trun
| |
| The apex of the ventricles becomes cus arteriosus.
| |
| | |
| more and more pointed. In the auricular portion a small longitudinal fold appears as the rudiment of the
| |
| auricular septum, while in the canalis auricularis, which is now at its greatest
| |
| length, there is also to be seen a commencement of the valvular structures
| |
| tending to separate the cavity of the auricles from those of the ventricles.
| |
| | |
| About the io6th hour, a septum begins to make its appearance in the
| |
| truncus arteriosus in the form of a longitudinal fold, which according to
| |
| Tonge (No. 495) starts at the end of the truncus furthest removed from the
| |
| heart. It takes origin from the wall of the truncus between the fourth and
| |
| fifth pairs of arches, and grows downwards in such a manner as to divide the
| |
| truncus into two channels, one of which leads from the heart to the third and
| |
| fourth pairs of arches, and the other to the fifth pair. Its course downwards
| |
| is not straight but spiral, and thus the two channels into which it divides
| |
| the truncus arteriosus wind spirally the one round the other.
| |
| | |
| At the time when the septum is first formed, the opening of the truncus
| |
| arteriosus into the ventricles is narrow or slit-like, apparently in order to
| |
| prevent the flow of the blood back into the heart. Soon after the appearance
| |
| of the septum, however, semilunar valves (Tonge, No. 495) are developed
| |
| from the wall of that portion of the truncus which lies between the free edge
| |
| of the septum and the cavity of the ventricles 1 .
| |
| | |
| 1 If Tonge is correct in his statement that the semilunar valves develop at some
| |
| distance from the mouth of the ventricle, it would seem possible that the portion of
| |
| the truncus between them and the ventricle ought to be regarded as the embryonic
| |
| conus arteriosus, and that the distal row of valves of the conus (and not the proximal
| |
| as suggested above, p. 639) has been preserved in the higher types.
| |
| | |
| | |
| | |
| THE VASCULAR SYSTEM.
| |
| | |
| | |
| | |
| 641
| |
| | |
| | |
| | |
| The ventral and the dorsal pairs of valves are the first to appear : the
| |
| former as two small solid prominences separated from each other by a
| |
| narrow groove ; the latter as a single ridge, in the centre of which is a
| |
| prominence indicating the point where the ridge will subsequently become
| |
| divided into two. The outer valves appear opposite each other, at a
| |
| considerably later period.
| |
| | |
| As the septum grows downwards towards the heart, it finally reaches
| |
| the position of these valves. One of its edges then passes between the two
| |
| ventral valves, and the other unites with the prominence on the dorsal
| |
| valve-ridge. At the same time the growth of all the parts causes the valves
| |
| to appear to approach the heart, and thus to be placed quite at the top
| |
| of the ventricular cavities. The free edge of the septum of the truncus now
| |
| | |
| A. B.
| |
| | |
| | |
| | |
| | |
| | |
| FlG. 361. TWO VIEWS OF THE HEART OF A CHICK UPON THE FIFTH DAY
| |
| | |
| OF INCUBATION.
| |
| | |
| A. from the ventral, B. from the dorsal side.
| |
| | |
| La. left auricular appendage; r.a. right auricular appendage ; r.v. right ventricle;
| |
| l.v. left ventricle; b. truncus arteriosus.
| |
| | |
| fuses with the ventricular septum, and thus the division of the truncus into
| |
| two separate channels, each provided with three valves, and each communicating with a separate side of the heart, is complete ; the position of
| |
| the valves not being very different from that in the adult heart.
| |
| | |
| That division of the truncus which opens into the fifth pair of arches is
| |
| the one which communicates with the right ventricle, while that which
| |
| opens into the third and fourth pairs communicates with the left ventricle.
| |
| The former becomes the pulmonary artery, the latter the commencement of
| |
| the systemic aorta.
| |
| | |
| The external constriction actually dividing the truncus into two vessels
| |
| does not begin to appear till the septum has extended some way back
| |
| towards the heart.
| |
| | |
| The semilunar valves become pocketed at a period considerably later
| |
| than their first formation (from the H7th to the,i65th hour) in the order of
| |
| their appearance.
| |
| | |
| At the end of the sixth day, and even on the fifth day (figs. 361 and 362),
| |
| the appearance of the heart itself, without reference to the vessels which
| |
| come from it, is not very dissimilar from that of the adult. The original
| |
| | |
| | |
| | |
| B. III.
| |
| | |
| | |
| | |
| 4 1
| |
| | |
| | |
| | |
| 642
| |
| | |
| | |
| | |
| THE HEART OF MAMMALIA.
| |
| | |
| | |
| | |
| r.a
| |
| | |
| | |
| | |
| | |
| l.v
| |
| | |
| | |
| | |
| FIG. 362. HEART OF A
| |
| CHICK UPON THE SIXTH DAY
| |
| OF INCUBATION, FROM THE
| |
| VENTRAL SURFACE.
| |
| | |
| La. left auricular appendage ;
| |
| r,a. right auricular appendage ;
| |
| r.v. right ventricle ; l.v. left ventricle ; b. truncus arteriosus.
| |
| | |
| | |
| | |
| protuberance to the right now forms the apex of the ventricles, and the
| |
| two auricular appendages are placed at the anterior extremity of the heart.
| |
| The most noticeable difference (in the ventral
| |
| view) is the still externally undivided condition of the truncus arteriosus.
| |
| | |
| The subsequent changes which the heart
| |
| undergoes are concerned more with its internal structure than with its external shape.
| |
| Indeed, during the next three days, viz. the
| |
| eighth, ninth, and tenth, the external form of
| |
| the heart remains nearly unaltered.
| |
| | |
| In the auricular portion, however, the
| |
| septum which commenced on the fifth day
| |
| becomes now more conspicuous. It is placed
| |
| vertically, and arises from the ventral wall ;
| |
| commencing at the canalis auricularis and
| |
| proceeding towards the opening into the
| |
| sinus venosus.
| |
| | |
| This latter structure gradually becomes
| |
| reduced so as to become a special appendage
| |
| of the right auricle. The inferior vena cava
| |
| | |
| enters the sinus obliquely from the right, so that its blood has a tendency to
| |
| flow towards the left auricle of the heart, which is at this time the larger of
| |
| the two.
| |
| | |
| The valves between the ventricles and auricles are now well developed,
| |
| and it is about this time that the division of the truncus arteriosus into the
| |
| aorta and pulmonary artery becomes visible from the exterior.
| |
| | |
| By the eleventh to the thirteenth day the right auricle has become as
| |
| large as the left, and the auricular septum much more complete, though
| |
| there is still a small opening, the foramen ovale, by which the two cavities
| |
| communicate with each other.
| |
| | |
| The most important feature in which the development of the Reptilian
| |
| heart differs from that of Birds is the division of the truncus into three
| |
| vessels, instead of two. The three vessels remain bound up in a common
| |
| sheath, and appear externally as a single trunk. The vessel not represented
| |
| in Birds is that which is continued into the left aortic arch.
| |
| | |
| In Mammals the early stages in the development of the heart present no
| |
| important points of difference from those of Aves. The septa in the truncus,
| |
| in the ventricular, and in the auricular cavities are formed, so far as
| |
| is known, in the same way and at the same relative periods in both groups.
| |
| In the embryo Man, the Rabbit, and other Mammals the division of
| |
| the ventricles is made apparent externally by a deep cleft, which, though
| |
| evanescent in these forms, is permanent in the Dugong.
| |
| | |
| The attachment of the auriculo-ventricular valves to the wall of the
| |
| ventricle, and the similar attachment of the left auriculo-ventricular valves
| |
| in Birds, have been especially studied by Gegenbaur and Bernays (No. 492),
| |
| | |
| | |
| | |
| ARTERIAL SYSTEM. 643
| |
| | |
| | |
| | |
| and deserve to be noticed. In the primitive state the ventricular walls
| |
| have throughout a spongy character ; and the auriculo-ventricular valves are
| |
| simple membranous projections like the auriculo-ventricular valves of Fishes.
| |
| Soon however the spongy muscular tissue of both the ventricular and
| |
| auricular walls, which at first pass uninterruptedly the one into the other,
| |
| grows into the bases of the valves, which thus become in the main muscular
| |
| projections of the walls of the heart. As the wall of the ventricle thickens,
| |
| the muscular trabeculas, connected at one end with the valves, remain at the
| |
| other end united with the ventricular wall, and form special bands passing
| |
| between the two. The valves on the other hand lose their muscular
| |
| attachment to the auricular walls. This is the condition permanent in
| |
| Ornithorhynchus. In higher Mammalia the ends of the muscular bands
| |
| inserted into the valves become fibrous, from the development of intermuscular connective tissue, and the atrophy of the muscular elements.
| |
| The fibrous parts now form the chordae tendinea?, and the muscular the
| |
| musculi papillares.
| |
| | |
| The sinus venosus in Mammals becomes completely merged into the
| |
| right auricle, and the systemic division of the truncus arteriosus is apparently not homologous with that in Birds.
| |
| | |
| In the embryos of all the Craniata the heart is situated very
| |
| far forwards in the region of the head. This position is retained
| |
| in Pisces. In Amphibia the heart is moved further back, while
| |
| in all the Amniota it gradually shifts its position first of all into
| |
| the region of the neck and finally passes completely within the
| |
| thoracic cavity. The steps in the change of position may be
| |
| gathered from figs. 109, in, and 118.
| |
| | |
| BIBLIOGRAPHY of the Heart.
| |
| | |
| (492) A. C. Bernays. " Entwicklungsgeschichte d. Atrioventricularklappen."
| |
| Morphol. Jahrbuch,^o\. II. 1876.
| |
| | |
| (493) E. Gasser. " Ueber d. Entstehung d. Herzens beim Hiihn." Archiv f.
| |
| mikr. Anat., Vol. xiv.
| |
| | |
| (494) A. Thomson. "On the development of the vascular system of the foetus
| |
| of Vertebrated Animals." Edinb. New Phil. Journal, Vol. ix. 1830 and 1831.
| |
| | |
| (495) M. Tonge. "Observations on the development of the semilunar valves
| |
| of the aorta and pulmonary artery of the heart of the Chick." Phil. Trans. CLIX.
| |
| 1869.
| |
| | |
| Vide also Von Baer (291), Rathke (300), Hensen (182), Kolliker (298), Gotte (296),
| |
| and Balfour (292).
| |
| | |
| Arterial System.
| |
| | |
| In the embryos of Vertebrata the arterial system consists of
| |
| a forward continuation of the truncus arteriosus, on the ventral
| |
| | |
| 41 2
| |
| | |
| | |
| | |
| 644
| |
| | |
| | |
| | |
| ARTERIES OF PISCES.
| |
| | |
| | |
| | |
| side of the throat (figs. 363, abr, and 364, a), which, with a few
| |
| exceptions to be noticed below, divides into as many branches on
| |
| each side as there are visceral arches. These branches, after
| |
| traversing the visceral arches, unite on the dorsal side of the
| |
| throat into a common trunk on each side. This trunk (figs. 363
| |
| and 364) after giving off one (or more) vessels to the head (c and
| |
| c] turns backv/ards, and bends in towards the middle line, close
| |
| to its fellow, immediately below the notochord (figs. 21 and 116)
| |
| and runs backwards in this situation towards the end of the tail.
| |
| The two parallel trunks below the notochord fuse very early into
| |
| a single trunk, the dorsal aorta (figs. 363, ad, and 364, a"}.
| |
| | |
| | |
| | |
| | |
| ttbr v "a,
| |
| | |
| FIG. 363. DIAGRAMMATIC VIEW OF THE HEAD OF AN EMBRYO TELEOSTEAN,
| |
| WITH THE PRIMITIVE VASCULAR TRUNKS. (From Gegenbaur.)
| |
| | |
| a. auricle ; v. ventricle ; abr. branchial artery ; c'. carotid ; ad. dorsal aorta ;
| |
| s. branchial clefts; sv. sinus venosus; dc. ductus Cuvieri; n. nasal pit
| |
| | |
| There is given off from each collecting trunk from the visceral
| |
| arches, or from the commencement of the dorsal aorta, a subclavian
| |
| artery to each of the anterior limbs ; from near the anterior end
| |
| of the dorsal aorta a vitelline artery (or before the dorsal aortae
| |
| have united a pair of arteries fig. 125, R of A and L of A) to the
| |
| yolk-sack, which subsequently becomes the main visceral artery 1 ;
| |
| and from the dorsal aorta opposite the hind limbs one (or two)
| |
| arteries on each side the iliac arteries to the hind limbs ; from
| |
| these arteries the allantoic arteries are given off in the higher
| |
| types, which remain as the hypogastric arteries after the
| |
| disappearance of the allantois.
| |
| | |
| The primitive arrangement of the arterial trunks is with a
| |
| few modifications retained in Fishes. With the development of
| |
| the gills the vessels to the arches become divided into two parts
| |
| connected by a capillary system in the gill folds, viz. into the
| |
| | |
| 1 In Mammalia the superior inesenteric artery arises from the vitelline artery,
| |
| which may probably be regarded as a primitive crclinco-mescnteric artery.
| |
| | |
| | |
| | |
| ARTERIAL SYSTEM.
| |
| | |
| | |
| | |
| branchial arteries bringing the blood to the gills from the truncus
| |
| arteriosus, and the branchial veins transporting it to the dorsal
| |
| aorta. The branchial vessels to those arches which do not bear
| |
| gills, either wholly or partially atrophy; thus in Elasmobranchii
| |
| the mandibular trunk, which is fully developed in the embryo
| |
| (fig. 193, \av}, atrophies, except for a small remnant bringing
| |
| blood to the rudimentary gill of the spiracle from the branchial
| |
| vein of the hyoid arch. In Ganoids the mandibular artery
| |
| atrophies, but the hyoid is usually preserved. In Teleostei both
| |
| mandibular 1 and hyoid arteries are absent in the adult, except
| |
| that there is usually left a rudiment of the hyoid, supplying the
| |
| pseudobranch, which is similar to the rudiment of the mandibular
| |
| artery in Elasmobranchii. In Dipnoi the mandibular artery
| |
| atrophies, but the hyoid is sometimes preserved (Protopterus),
| |
| and sometimes lost.
| |
| | |
| In Fishes provided with a well developed air-bladder this
| |
| organ receives arteries, which arise sometimes from the dorsal
| |
| aorta, sometimes from the caeliac arteries, and sometimes from
| |
| the dorsal section of the last (fourth) branchial trunk. The
| |
| latter origin is found in Polypterus and Amia, and seems to have
| |
| been inherited by the Dipnoi where the air-bladder forms a true
| |
| lung.
| |
| | |
| The pulmonary artery of all the air-breathing Vertebrata is derived from the pulmonary artery of the
| |
| Dipnoi.
| |
| | |
| In all the types above Fishes considerable changes are
| |
| effected in the primitive arrangement of the arteries in the
| |
| visceral arches.
| |
| | |
| In Amphibia the piscine condition is most nearly retained 2 .
| |
| The mandibular artery is never developed, and the hyoid artery
| |
| is imperfect, being only connected with the cephalic vessels and
| |
| never directly joining the dorsal aorta. It is moreover developed
| |
| later than the arteries of the true branchial arches behind. The
| |
| subclavian arteries spring from the common trunks which unite
| |
| to form the dorsal aorta.
| |
| | |
| In the Urodela there are developed, in addition to the hyoid,
| |
| | |
| 1 The mandibular artery is stated by Gotte never to be developed in Teleostei, but
| |
| is distinctly figured in Lereboullet (No. 71).
| |
| | |
| 2 In my account of the Amphibia, Gotte (No. 296) has been followed.
| |
| | |
| | |
| | |
| 646 ARTERIES OF THE AMNIOTA.
| |
| | |
| four branchial arteries. The three foremost of these at first
| |
| supply gills, and in the Perennibranchiate forms continue to do
| |
| so through life. The fourth does not supply a gill, and very
| |
| early gives off, as in the Dipnoi, a pulmonary branch.
| |
| | |
| The hyoid artery soon sends forward a lingual artery from its
| |
| ventral end, and is at first continued to the carotid which grows
| |
| forward from the dorsal part of the first branchial vessel.
| |
| | |
| In the Caducibranchiata, where the gills atrophy, the following
| |
| changes take place. The remnant of the hyoid is continued
| |
| entirely into the lingual artery. The first branchial is mainly
| |
| continued into the carotid and other cephalic branches, but a
| |
| narrow remnant of the trunk, which originally connected it with
| |
| the dorsal aorta, remains, forming what is known as a ductus
| |
| Botalli. A rete mirabile on its course is the remnant of the
| |
| original gill.
| |
| | |
| The second and third branchial arches are continued as
| |
| simple trunks into the dorsal aorta, and the blood from the fourth
| |
| arch mainly passes to the lungs, but a narrow ductus Botalli still
| |
| connects this arch with the dorsal aorta.
| |
| | |
| In the Anura the same number of arches is present in the
| |
| embryo as in the Urodela, all four branchial arteries supplying
| |
| branchiae, but the arrangement of the two posterior trunks is
| |
| different from that in the Urodela. The third arch becomes at a
| |
| very early period continued into a pulmonary vessel, a relativelynarrow branch connecting it with the second arch. The fourth
| |
| arch joins the pulmonary branch of the third. At the metamorphosis the hyoid artery loses its connection with the carotid, and
| |
| the only part of it which persists is the root of the lingual artery.
| |
| The first branchial artery ceases to join the dorsal aorta, and
| |
| forms the root of the carotid : the so-called carotid gland placed
| |
| on its course is the remnant of the gill supplied by it before the
| |
| metamorphosis.
| |
| | |
| The second artery forms a root of the dorsal aorta. The
| |
| third, as in all the Amniota, now supplies the lungs, and also
| |
| sends off a cutaneous branch. The fourth disappears. The
| |
| connection of the pulmonary artery with both the third and
| |
| fourth branchial arches in the embryo appears to me clearly to
| |
| indicate that this artery was primitively derived from the fonrtli
| |
| arc/i as in the Urodela, and that its permanent connection
| |
| | |
| | |
| | |
| ARTERIAL SYSTEM.
| |
| | |
| | |
| | |
| 647
| |
| | |
| | |
| | |
| with the third arch in the Anura and in all the Amniota is
| |
| secondary.
| |
| | |
| In the Amniota the metamorphosis of the arteries is in all
| |
| cases very similar. Five arches, viz. the mandibular, hyoid, and
| |
| three branchial arches are always developed (fig. 364), but, owing
| |
| to the absence of branchiae,
| |
| never function as branchial arteries. Of these the main parts of
| |
| the first two, connecting the truncus arteriosus with the collecting
| |
| trunk into which the arterial
| |
| arches fall, always disappear, usually before the complete development of the arteries in the posterior arches.
| |
| | |
| The anterior part of the collecting trunk into which these
| |
| vessels fall is not obliterated
| |
| when they disappear, but is on
| |
| the contrary continued forwards
| |
| as a vessel supplying the brain,
| |
| homologous with that found in
| |
| Fishes. It constitutes the internal
| |
| carotid. Similarly the anterior
| |
| part of the trunk from which the mandibular and hyoid arteries
| |
| sprang is continued forwards as a small vessel 1 , which at first
| |
| passes to the oral region and constitutes in Reptiles the lingual
| |
| artery, homologous with the lingual artery of the Amphibia ; but
| |
| in Birds and Mammals becomes more important, and is then
| |
| known as the external carotid (fig. 125). By these changes the
| |
| roots of the external and internal carotids spring respectively
| |
| from the ventral and dorsal ends of the primitive third artery,
| |
| i.e. the artery of the first branchial arch (fig. 365, c and c'} ; and
| |
| thus this arterial arch persists in all types as the common carotid,
| |
| | |
| | |
| | |
| | |
| FIG. 364. DIAGRAM OF THE ARRANGEMENT OF THE ARTERIAL
| |
| ARCHES IN AN EMBRYO OF ONE OF THE
| |
| | |
| AMNIOTA. (From Gegenbaur ; after
| |
| RATHKE.)
| |
| | |
| a. ventral aorta; a", dorsal aorta;
| |
| ' 2 > 3> 4> 5- arterial arches ; c. carotid
| |
| artery.
| |
| | |
| | |
| | |
| 1 His (No. 232) describes in Man two ventral continuations of the truncus arteriosus, one derived from the mandibular artery, forming the external maxillary artery,
| |
| and one from the hyoid artery, forming the lingual artery. The vessel from which
| |
| they spring is the external carotid. These observations of His will very probably be
| |
| found to hold true for other types.
| |
| | |
| | |
| | |
| 6 4 8
| |
| | |
| | |
| | |
| ARTERIAL ARCHES OF THE AMNIOTA.
| |
| | |
| | |
| | |
| and the basal part of the internal carotid. The trunk connecting
| |
| the third arterial arch with the system of the dorsal aorta persists
| |
| in some Reptiles (Lacertilia, fig. 366 A) as a ductus Botalli, but
| |
| is lost in the remaining Reptiles and in Birds and Mammals (fig.
| |
| 366 B, C, D). It disappears earliest in Mammals (fig. 365 C),
| |
| later in Birds (fig. 365 B), and still later in the majority of
| |
| Reptiles.
| |
| | |
| The fourth arch always continues to give rise, as in the Anura,
| |
| to the system of the dorsal aorta.
| |
| | |
| In all Reptiles it persists on both sides (fig. 366 A and B),
| |
| but with the division of the truncus arteriosus into three vessels
| |
| | |
| | |
| | |
| | |
| ad
| |
| | |
| | |
| | |
| FIG. 365. DEVELOPMENT OF THE GREAT ARTERIAL TRUNKS IN THE EMBRYOS
| |
| OF A. A LIZARD ; B. THE COMMON FOWL; C. THE PIG. (From Gegenbaur; after
| |
| Rathke.)
| |
| | |
| The first two arches have disappeared in all three. In A and B the last three are
| |
| still complete, but in C the last two are alone complete.
| |
| | |
| /. pulmonary artery springing from the fifth arch, but still connected with the
| |
| system of the dorsal aorta by a ductus Botalli; c. external carotid; <'. internal
| |
| carotid; ad. dorsal aorta; a. auricle; v. ventricle; n. nasal pit; m, rudiment of
| |
| fore-limb.
| |
| | |
| one of these, i.e. that opening furthest to the left side of the
| |
| ventricle (e and d), is continuous with the right fourth arch, and
| |
| also with the common carotid arteries (c) ; while a second
| |
| springing from the right side of the ventricle is continuous with
| |
| the left fourth arch (Ji and f). The right and left divisions of the
| |
| fourth arch meet however on the dorsal side of the oesophagus to
| |
| give origin to the dorsal aorta (g).
| |
| | |
| In Birds (fig. 366 C) the left fourth arch (h) loses its connection with the dorsal aorta, though the ventral part remains as
| |
| | |
| | |
| | |
| ARTERIAL SYSTEM.
| |
| | |
| | |
| | |
| 649
| |
| | |
| | |
| | |
| the root of the left subclavian. The truncus arteriosus is moreover only divided into two parts, one of which is continuous
| |
| with all the systemic arteries. Thus it comes about that in
| |
| Birds the right fourth arch (e) alone gives rise to the dorsal
| |
| aorta.
| |
| | |
| In Mammals (fig. 366 D) the truncus arteriosus is only
| |
| divided into two, but the left fourth arch (>), instead of the right,
| |
| is that continuous with the dorsal aorta, and the right fourth
| |
| arch (/) is only continued into the right vertebral and right
| |
| subclavian arteries.
| |
| | |
| The fifth arch always gives origin to the pulmonary artery
| |
| (fig. 365, /) and is continuous with one of the divisions of the
| |
| truncus arteriosus. In Lizards (fig. 366 A, i), Chelonians and
| |
| Birds (fig. 366 C, i] and probably in Crocodilia, the right and
| |
| left pulmonary arteries spring respectively from the right and
| |
| left fifth arches, and during the greater part of embryonic life
| |
| the parts of the fifth arches between the origins of the pulmonary
| |
| arteries and the system of the dorsal aorta are preserved as
| |
| ductus Botalli. These ductus Botalli persist for life in the
| |
| Chelonia. In Ophidia (fig. 366 B, Ji) and Mammalia (fig.
| |
| 366 D, m) only one of the fifth arches gives origin to the two
| |
| pulmonary arteries, viz. that on the right side in Ophidia, and
| |
| the left in Mammalia.
| |
| | |
| The ductus Botalli of the fifth arch (known in Man as the
| |
| ductus arteriosus) of the side on which the pulmonary arteries
| |
| are formed, may remain (e.g. in Man) as a solid cord connecting
| |
| the common stem of the pulmonary aorta with the systemic
| |
| aorta.
| |
| | |
| The main history of the arterial arches in the Amniota has
| |
| been sufficiently dealt with, and the diagram, fig. 366, copied
| |
| from Rathke, shews at a glance the character of the metamorphosis these arches undergo in the different types. It merely
| |
| remains for me to say a few words about the subclavian and
| |
| vertebral arteries.
| |
| | |
| The subclavian arteries in Fishes usually spring from the
| |
| trunks connecting the branchial veins with the dorsal aorta.
| |
| This origin, which is also found in Amphibia, is typically found
| |
| in the embryos of the Amniota. In the Lizards this origin
| |
| persists through life, but both subclavians spring from the right
| |
| | |
| | |
| | |
| 650
| |
| | |
| | |
| | |
| ARTERIAL ARCHES OF THE AMNIOTA.
| |
| | |
| | |
| | |
| side. In most other types the origin of the subclavians is
| |
| carried upwards, so that they usually spring from a trunk
| |
| common to them and the carotids (arteria anonyma) (Birds and
| |
| some Mammals); or the left one, as in Man and some other
| |
| Mammals, arises from the systemic aorta just beyond the
| |
| carotids. Various further modifications in the origin of the
| |
| subclavians of the same general nature are found in Mammalia,
| |
| A 13
| |
| | |
| | |
| | |
| | |
| | |
| FIG. 366. DIAGRAMS ILLUSTRATING THE METAMORPHOSIS OF THE ARTERIAL
| |
| | |
| ARCHES IN A LlZARD A, A SNAKE B, A BlRD C AND A MAMMAL D. (From
| |
| Mivart ; after Rathke.)
| |
| | |
| A. a. internal carotid; b. external carotid ; c. common carotid; d. ductus Botalli
| |
| between the third and fourth arches ; e. right aortic trunk ; /. subclavian ; g. dorsal
| |
| aorta; h. left aortic trunk; i. pulmonary artery; k. rudiment of ductus Botalli
| |
| between the pulmonary artery and the system of the dorsal aorta.
| |
| | |
| B. a. internal carotid; b. external carotid; c. common carotid; d. right aortic
| |
| trunk; e. vertebral artery;/, left aortic trunk of dorsal aorta; h. pulmonary artery ;
| |
| i. ductus Botalli of pulmonary artery.
| |
| | |
| C. a. internal carotid ; b. external carotid ; c. common carotid ; d. systemic
| |
| aorta; e. fourth arch of right side (root of dorsal aorta);/, right subclavian; g. dorsal
| |
| aorta; h, left subclavian (fourth arch of left side); i. pulmonary artery; k. and /.
| |
| right and left ductus Botalli of pulmonary arteries.
| |
| | |
| D. a. internal carotid; b. external carotid; c. common carotid; d. systemic aorta;
| |
| c. fourth arch of left side (root of dorsal aorta);/ dorsal aorta; g. left vertebral
| |
| artery; h. left subclavian artery; i. right subclavian (fourth arch of right side); k.
| |
| right vertebral; /. continuation of right subclavian; in. pulmonary artery; n. ductus
| |
| Botalli of pulmonary artery.
| |
| | |
| | |
| | |
| THE VENOUS SYSTEM.
| |
| | |
| | |
| | |
| 6 5 I
| |
| | |
| | |
| | |
| but they need not be specified in detail. The vertebral arteries
| |
| usually arise in close connection with the subclavians, but in
| |
| Birds they arise from the common carotids.
| |
| | |
| BIBLIOGRAPHY of the Arterial System.
| |
| | |
| (496) H. Rathke. " Ueb. d. Entwick. d. Arterien vv. bei d. Saugethiere von
| |
| d. Bogen d. Aorta ausgehen." Miiller's Archiv, 1843.
| |
| | |
| (-197) H. Rathke. " Untersuchungen lib. d. Aortenwurzeln d. Saurier."
| |
| Denkschriften d. k. Akad. Wien, Vol. XIII. 1857.
| |
| | |
| Vide also His (No. 232) and general works on Vertebrate Embryology.
| |
| | |
| TJie Venous System,.
| |
| | |
| The venous system, as it is found in the embryos of Fishes,
| |
| consists in its earliest condition of a single large trunk, which
| |
| traverses the splanchnic mesoblast investing the part of the
| |
| alimentary tract behind the heart. This trunk is directly continuous in front with the heart, and underlies the alimentary
| |
| canal through both its praeanal and postanal sections. It is
| |
| shewn in section in fig. 367, v, and may be called the subintestinal vein. This vein has been found in the embryos of
| |
| Teleostei, Ganoidei, Elasmobranchii and Cyclostomata, and runs
| |
| parallel to the dorsal aorta above, into which it is sometimes
| |
| continued behind (Teleostei, Ganoidei, etc.).
| |
| | |
| In Elasmobranch embryos the subintestinal vein terminates,
| |
| as may be gathered from sections (fig. 368, v.cau), shortly before
| |
| the end of the tail. The same series of sections also shews that
| |
| at the cloaca, where the gut enlarges and comes in contact with
| |
| the skin, this vein bifurcates, the two branches uniting into a
| |
| single vein both in front of and behind the cloaca.
| |
| | |
| In most Fishes the anterior part of this vein atrophies, the
| |
| caudal section alone remaining, but the anterior section of it
| |
| persists in the fold of the intestine in Petromyzon, and also
| |
| remains in the spiral valve of some Elasmobranchii. In
| |
| Amphioxus, moreover, it forms, as in the embryos of higher
| |
| types, the main venous trunk, though even here it is usually
| |
| broken up into two or three parallel vessels.
| |
| | |
| It no doubt represents one of the primitive longitudinal trunks of the
| |
| vermiform ancestors of the Chordata. The heart and the branchial artery
| |
| constitute a specially modified anterior continuation of this vein. The
| |
| | |
| | |
| | |
| 652
| |
| | |
| | |
| | |
| THE SUBINTESTINAL VEIN.
| |
| | |
| | |
| | |
| -p.o
| |
| | |
| | |
| | |
| rp.r.
| |
| | |
| | |
| | |
| dilated portal sinus of Myxine is probably also part of it ; and if this is
| |
| really rhythmically contractile 1 the fact would be interesting as shewing that
| |
| this quality, which is now localised in the heart, was once probably common
| |
| to the subintestinal vessel for its whole length.
| |
| | |
| On the development of the cardinal veins (to be described
| |
| below) considerable changes are
| |
| effected in the subintestinal vein.
| |
| Its postanal section, which is known
| |
| in the adult as the caudal vein,
| |
| unites with the cardinal veins. On
| |
| this junction being effected retrogressive changes take place in the
| |
| praeanal section of the original subintestinal vessel. It breaks up in
| |
| front into a number of smaller
| |
| vessels, the most important of which
| |
| is a special vein, which lies in the
| |
| fold of the spiral valve, and which is
| |
| more conspicuous in some Elasmobranchii than in Scyllium, in which
| |
| the development of the vessel has
| |
| been mainly studied. The lesser of
| |
| the two branches connecting it
| |
| round the cloaca with the caudal
| |
| vein first vanishes, and then the
| |
| larger ; and the two posterior cardinals are left as the sole forward
| |
| continuations of the caudal vein.
| |
| The latter then becomes prolonged
| |
| forwards, so that the two cardinals
| |
| open into it some little distance in
| |
| front of the hind end of the kidneys.
| |
| By these changes, and by the disappearance of the postanal section of the gut, the caudal vein is
| |
| made to appear as a supraintestinal and not, as it really is, a
| |
| subintestinal vessel.
| |
| | |
| From the subintestinal vein there is given off a branch which
| |
| supplies the yolk-sack. This leaves the subintestinal vein close
| |
| | |
| 1 J. Miiller holds that this sack is not rhythmically contractile.
| |
| | |
| | |
| | |
| | |
| FIG. 367. SECTION THROUGH
| |
| THE TRUNK OF A SCYLLIUM
| |
| EMBRYO SLIGHTLY YOUNGER
| |
| | |
| THAN 28 F.
| |
| | |
| sp.c. spinal canal; W. white
| |
| matter of spinal cord ; pr. posterior nerve-roots; ch. notochord ;
| |
| x. sub-notochordal rod ; ao. aorta ;
| |
| mp. muscle plate; ;;//'. inner layer
| |
| of muscle-plate already converted
| |
| into muscles; Vr. rudiment of
| |
| vertebral body; st. segmental
| |
| tube ; sd. segmental duct ; sp.v.
| |
| spiral valve; v. subintestinal vein ;
| |
| p.o. primitive generative cells.
| |
| | |
| | |
| | |
| THE VENOUS SYSTEM.
| |
| | |
| | |
| | |
| 653
| |
| | |
| | |
| | |
| to the liver. The liver, on its development, embraces the
| |
| subintestinal vein, which then breaks up into a capillary system
| |
| in the liver, the main part of its blood coming at this period
| |
| from the yolk-sack.
| |
| | |
| The portal system is thus established from the subintestinal
| |
| vein ; but is eventually joined by the various visceral, and sometimes by the genital, veins as they become successively developed.
| |
| | |
| The blood from the liver is brought back to the sinus venosus by veins known as the hepatic veins, which, like the hepatic
| |
| capillary system, are derivatives of the subintestinal vessel.
| |
| | |
| There join the portal system in Myxinoids and many
| |
| Teleostei a number of veins from the anterior abdominal walls,
| |
| representing a commencement of the anterior abdominal or
| |
| epigastric vein of higher types 1 .
| |
| | |
| In the higher Vertebrates the original subintestinal vessel never attains a
| |
| full development, even in the embryo. It is represented by (i) the ductus
| |
| | |
| | |
| | |
| | |
| FIG. 368. FOUR SECTIONS THROUGH THE POSTANAL PART OF THE TAIL
| |
| OF AN EMBRYO OF THE SAME AGE AS FIG. 28 F.
| |
| | |
| A. is the posterior section.
| |
| | |
| nc. neural canal; al. post-anal gut; alv. caudal vesicle of post-anal gut; x.
| |
| subnotochordal rod; mp. muscle-plate; c/i. notochord; cl.al. cloaca; ao. aorta;
| |
| v.cait. caudal vein.
| |
| | |
| 1 Stannius, Vergleich. Anat., p. 251.
| |
| | |
| | |
| | |
| 654
| |
| | |
| | |
| | |
| THE CARDINAL VEINS.
| |
| | |
| | |
| | |
| venosus, which, like the true subintestinal vein, gives origin (in the Amniota)
| |
| to the vitelline veins to the yolk-sack, and (2) by the caudal vein. Whether
| |
| the partial atrophy of the subintestinal vessel was primitively caused by the
| |
| development of the cardinal veins, or for some other reason, it is at any rate
| |
| a fact that in all existing Fishes the cardinal veins form the main venous
| |
| channels of the trunk.
| |
| | |
| Their later development than the subintestinal vessel as well as their
| |
| absence in Amphioxus, probably indicate that they became evolved, at any
| |
| rate in their present form, within the Vertebrate phylum.
| |
| | |
| The embryonic condition of the venous system, with a single
| |
| large subintestinal vein is, as has been stated, always modified
| |
| by the development of a paired system of vessels, known as the
| |
| cardinal veins, which bring to the heart the greater part of the
| |
| blood from the trunk.
| |
| | |
| The cardinal veins appear in Fishes as four paired longitudinal trunks (figs. 363 and 369), two anterior (/) and two
| |
| posterior (c). They unite into two transverse trunks on either
| |
| side, known as the ductus Cuvieri (dc), which fall into the sinus
| |
| venosus, passing from the body wall to the sinus by a lateral
| |
| mesentery of the heart already spoken of (p. 627, fig. 352). The
| |
| anterior pair, known as the anterior cardinal or jugular veins,
| |
| bring to the heart the blood from the head and neck. They
| |
| are placed one on each side above the level
| |
| of the branchial arches (fig. 299, a.cv). The
| |
| posterior cardinal veins lie immediately dorsal to the mesonephros (Wolfifian body), and
| |
| are mainly supplied by the blood from this
| |
| organ and from the walls of the body (fig.
| |
| 275, c.a.v). In many forms (Cyclostomata,
| |
| Elasmobranchii and many Teleostei) they
| |
| unite posteriorly with the caudal veins in
| |
| the manner already described, and in a large
| |
| number of instances the connecting branch
| |
| between the two systems, in its passage through
| |
| the mesonephros, breaks up into a capillary
| |
| network, and so gives rise to a renal portal
| |
| system.
| |
| | |
| The vein from the anterior pair of fins
| |
| (subclavian) usually unites with the anterior
| |
| jugular vein.
| |
| | |
| | |
| | |
| | |
| j
| |
| | |
| | |
| | |
| FIG. 369. DIAGRAM OF THE PAIRED VENOUS SYSTEM
| |
| | |
| OF A FISH. (From
| |
| Gegenbaur. )
| |
| | |
| j. jugular vein
| |
| (anterior cardinal
| |
| vein) ; c. posterior
| |
| cardinal vein; //. hepatic veins ; sv. sinus
| |
| venosus ; dc. ductus
| |
| Cuvieri.
| |
| | |
| | |
| | |
| THE VENOUS SYSTEM. 655
| |
| | |
| The venous system of the Amphibia and Amniota always
| |
| differs from that of Fishes in the presence of a new vessel, the
| |
| vena cava inferior, which replaces the posterior cardinal veins;
| |
| the latter only being present, in their piscine form, during
| |
| embryonic life. It further differs from that of all Fishes, except
| |
| the Dipnoi, in the presence of pulmonary veins bringing back
| |
| the blood directly from the lungs.
| |
| | |
| In the embryos of all the higher forms the general characters
| |
| of the venous system are at first the same as in Fishes, but with
| |
| the development of the vena cava inferior the front sections of
| |
| the posterior cardinal veins atrophy, and the ductus Cuvieri,
| |
| remaining solely connected with the anterior cardinals and their
| |
| derivatives, constitute the superior venae cavae. The inferior
| |
| cava receives the hepatic veins.
| |
| | |
| Apart from the non-development of the subintestinal vein
| |
| the visceral section of the venous system is very similar to that
| |
| in Fishes.
| |
| | |
| The further changes in the venous system must be dealt
| |
| with separately for each group.
| |
| | |
| Amphibia. In Amphibia (Gotte, No. 296) the anterior and posterior
| |
| cardinal veins arise as in Pisces. From the former the internal jugular vein
| |
| arises as a branch ; the external jugular constituting the main stem. The
| |
| subclavian with its large cutaneous branch also springs from the system of
| |
| the anterior cardinal. The common trunk formed by the junction of these
| |
| three veins falls into the ductus Cuvieri.
| |
| | |
| The posterior cardinal veins occupy the same position as in Pisces, and
| |
| unite behind with the caudal veins, which Gotte has shewn to be originally
| |
| situated below the post-anal gut. The iliac veins unite with the posterior
| |
| cardinal veins, where the latter fall into the caudal vein. The original
| |
| piscine condition of the veins is not long retained. It is first of all disturbed
| |
| by the development of the anterior part of the important unpaired venous
| |
| trunk which forms in the adult the vena cava inferior. This is developed
| |
| independently, but unites behind with the right posterior cardinal. From
| |
| this point backwards the two cardinal veins coalesce for some distance, to
| |
| give rise to the posterior section of the vena cava inferior, situated between
| |
| the kidneys 1 . The anterior sections of the cardinal veins subsequently
| |
| atrophy. The posterior part of the cardinal veins, from their junction with
| |
| the vena cava inferior to the caudal veins, forms a rhomboidal figure. The
| |
| iliac vein joins the outer angle of this figure, and is thus in direct communication with the inferior vena cava, but it is also connected with a longitu
| |
| 1 This statement of Gotte's is opposed to that of Rathke for the Amniota, and
| |
| cannot be considered as completely established.
| |
| | |
| | |
| | |
| 656 VEINS OF THE SNAKE.
| |
| | |
| dinal vessel on the outer border of the kidneys, which receives transverse
| |
| vertebral veins and transmits their blood to the kidneys, thus forming a
| |
| renal portal system. The anterior limbs of the rhomboid formed by the
| |
| cardinal veins soon atrophy, so that the blood from the hind limbs can only
| |
| pass to the inferior vena cava through the renal portal system. The
| |
| posterior parts of the two cardinal veins (uniting in the Urodela directly
| |
| with the unpaired caudal vein) still persist. The iliac veins also become
| |
| directly connected with a new vein, the anterior abdominal vein, which
| |
| has meanwhile become developed. Thus the iliac veins become united
| |
| with the system of the vena cava inferior through the vena renalis advehens
| |
| on the outer border of the kidney, and with the anterior abdominal veins by
| |
| the epigastric veins.
| |
| | |
| The visceral venous system begins with the development of two vitelline
| |
| veins, which at first join the sinus venosus directly. They soon become
| |
| enveloped in the liver, where they break up into a capillary system, which
| |
| is also joined by the other veins from the viscera. The hepatic system has
| |
| in fact the same relations as in Fishes. Into this system the anterior
| |
| abdominal vein also pours itself in the adult. This vein is originally
| |
| formed of two vessels, which at first fall directly into the sinus venosus,
| |
| uniting close to their opening into the sinus with a vein from the truncus
| |
| arteriosus. They become prolonged backwards, and after receiving the
| |
| epigastric veins above mentioned from the iliac veins, and also veins from
| |
| the allantoic bladder, unite behind into a single vessel. Anteriorly the
| |
| right vein atrophies and the left continues forward the unpaired posterior
| |
| section.
| |
| | |
| A secondary connection becomes established between the anterior abdominal vein and the portal system ; so that the blood originally transported
| |
| by the former vein to the heart becomes diverted so as to fall into the liver.
| |
| A remnant of the primitive connection is still retained in the adult in the
| |
| form of a small vein, the so-called vena bulbi posterior, which brings the
| |
| blood from the walls of the truncus arteriosus directly into the anterior
| |
| abdominal vein.
| |
| | |
| The pulmonary veins grow directly from the heart to the lungs.
| |
| | |
| For our knowledge of the development of the venous system of the
| |
| Amniota we are mainly indebted to Rathke.
| |
| | |
| Reptilia. As an example of the Reptilia the Snake may be selected,
| |
| its venous system having been fully worked out by Rathke in his important
| |
| memoir on its development (No. 300).
| |
| | |
| The anterior (external jugular) and posterior cardinal veins are formed in
| |
| the embryo as in all other types (fig. 370, vj and vc] ; and the anterior
| |
| cardinal, after giving rise to the anterior vertebral and to the cephalic veins,
| |
| persists with but slight modifications in the adult ; while the two ductus
| |
| Cuvieri constitute the superior venos cavas.
| |
| | |
| The two posterior cardinals unite behind with the caudal veins. They
| |
| are placed in the usual situation on the dorsal and outer border of the
| |
| kidneys.
| |
| | |
| | |
| | |
| THE VENOUS SYSTEM.
| |
| | |
| | |
| | |
| 657
| |
| | |
| | |
| | |
| | |
| U
| |
| FIG. 370. ANTERIOR
| |
| PORTION OF THE VENOUS
| |
| SYSTEM OF AN EMBRYONIC
| |
| SNAKE. (From Gegenbaur;
| |
| after Rathke.)
| |
| | |
| vc. posterior cardinal
| |
| vein; vj. jugular vein; DC.
| |
| ductus Cuvieri ; vu. allantoic vein ; v. ventricle ; ba.
| |
| truncus arteriosus ; a. visceral clefts ; /. auditory
| |
| vesicle.
| |
| | |
| | |
| | |
| With the development of the vena cava inferior, to be described below,
| |
| the blood from the kidneys becomes mainly
| |
| transported by this vessel to the heart ; and the
| |
| section of the posterior cardinals opening into
| |
| the ductus Cuvieri gradually atrophies, their
| |
| posterior parts remaining however on the outer
| |
| border of the kidneys as the vena? renales
| |
| advehentes 1 .
| |
| | |
| While the front part of the posterior cardinal
| |
| veins is undergoing atrophy, the intercostal veins,
| |
| which originally poured their blood into the
| |
| posterior cardinal veins, become also connected
| |
| with two longitudinal veins the posterior vertebral veins which are homologous with the
| |
| azygos and hemiazygos veins of Man ; and bear
| |
| the same relation to the anterior vertebral veins
| |
| that the anterior and posterior cardinals do to
| |
| each other.
| |
| | |
| These veins are at first connected by trans
| |
| verse anastomoses with the posterior cardinals,
| |
| but, on the disappearance of the front part of the
| |
| latter, the whole of the blood from the intercostal veins falls into the
| |
| posterior vertebral veins. They are united in front with the anterior vertebral veins, and the common trunk of the two veins on each side falls into
| |
| the jugular vein.
| |
| | |
| The posterior vertebral veins are at first symmetrical, but after becoming
| |
| connected by transverse anastomoses, the right becomes the more important
| |
| of the two.
| |
| | |
| The vena cava inferior, though considerably later in its development
| |
| than the cardinals, arises fairly early. It constitutes in front an unpaired
| |
| trunk, at first very small, opening into the right allantoic vein, close to the
| |
| heart. Posteriorly it is continuous with two veins placed on the inner
| |
| border of the kidneys 2 .
| |
| | |
| The vena cava inferior passes through the dorsal part of the liver, and in
| |
| doing so receives the hepatic veins.
| |
| | |
| The portal system is at first constituted by the vitelline vein, which is
| |
| directly continuous with the venous end of the heart, and at first receives
| |
| the two ductus Cuvieri, but at a later period unites with the left ductus.
| |
| | |
| 1 Rathke's account of the vena renalis advehens is thus entirely opposed to that
| |
| which Gotte gives for the Frog, but my own observations on the Lizard incline me to
| |
| accept Rathke's statements, for the Amniota at any rate.
| |
| | |
| 2 The vena cava inferior does not according to Rathke's account unite behind with
| |
| the posterior cardinal veins, as it is stated by Gotte to do in the Anura. Gb'tte
| |
| questions the accuracy of Rathke's statements on this head, but my own observations
| |
| are entirely in favour of Rathke's observations, and lend no support whatever to
| |
| Gotte's views.
| |
| | |
| | |
| | |
| B. III.
| |
| | |
| | |
| | |
| 658 VEINS OF THE CHICK.
| |
| | |
| It soon receives a mesenteric vein bringing the blood from the viscera,
| |
| which is small at first but rapidly increases in importance.
| |
| | |
| The common trunk of the vitelline and mesenteric veins, which may be
| |
| called the portal vein, becomes early enveloped by the liver, and gives off
| |
| branches to this organ, the blood from which passes by the hepatic veins
| |
| to the vena cava inferior. As the branches in the liver become more
| |
| important, less and less blood is directly transported to the heart, and finally
| |
| the part of the original vitelline vein in front of the liver is absorbed, and the
| |
| whole of the blood from the portal system passes from the liver into the
| |
| vena cava inferior.
| |
| | |
| The last section of the venous system to be dealt with is that of the
| |
| anterior abdominal vein. There are originally, as in the Anura, two veins
| |
| belonging to this system, which owing to the precocious development of the
| |
| bladder to form the allantois, constitute the allantoic veins (fig. 370, vu}.
| |
| | |
| These veins, running along the anterior abdominal wall, are formed
| |
| somewhat later than the vitelline vein, and fall into the two ductus Cuvieri.
| |
| They unite with two epigastric veins (homologous with those in the Anura),
| |
| which connect them with the system of the posterior cardinal veins. The
| |
| left of the two eventually atrophies, so that there is formed an unpaired
| |
| allantoic vein. This vein at first receives the vena cava inferior close to the
| |
| heart, but eventually the junction of the two takes place in the region of the
| |
| liver, and finally the anterior abdominal vein (as it comes to be after the
| |
| atrophy of the allantois) joins the portal system and breaks up into capillaries
| |
| in the liver 1 .
| |
| | |
| In Lizards the iliac veins join the posterior cardinals, and so pour part of
| |
| their blood into the kidneys ; they also become connected by the epigastric
| |
| veins with the system of the anterior abdominal or allantoic vein. The
| |
| subclavian veins join the system of the superior venae cavas.
| |
| | |
| The venous system of Birds and Mammals differs in two important
| |
| points from that of Reptilia and Amphibia. Firstly the anterior abdominal
| |
| vein is only a foetal vessel, forming during foetal life the allantoic vein ;
| |
| and secondly a direct connection is established between the vena cava
| |
| inferior and the veins of the hind limbs and posterior parts of the cardinal
| |
| veins, so that there is no renal portal system.
| |
| | |
| Aves. The Chick may be taken to illustrate the development of the
| |
| venous system in Birds.
| |
| | |
| On the third day, nearly the whole of the venous blood from the body
| |
| of the embryo is carried back to the heart by two main venous trunks,
| |
| the anterior (fig. 125, S.Ca.V) and posterior (V.Ca) cardinal veins, joining on
| |
| each side to form the short transverse ductus Cuvieri (DC), both of which
| |
| unite with the sinus venosus close to the heart. As the head and neck
| |
| continue to enlarge, and the wings become developed, the single anterior
| |
| | |
| 1 The junction between the portal system and the anterior abdominal vein is
| |
| apparently denied by Rathke (No. 300, p. 173), hut this must he an error on
| |
| his part.
| |
| | |
| | |
| | |
| THE VENOUS SYSTEM.
| |
| | |
| | |
| | |
| 659
| |
| | |
| | |
| | |
| | |
| V.C.L
| |
| | |
| | |
| | |
| cardinal or jugular vein (fig. 371, /), of each side, is joined by two new
| |
| veins : the vertebral vein, bringing back blood from the head and neck, and
| |
| the subclavian vein from the wing (W\
| |
| | |
| On the third day the posterior cardinal veins are the only veins which
| |
| return the blood from the hinder part of the body of the embryo.
| |
| | |
| About the fourth or fifth day, however, the vena cava inferior (fig. 371,
| |
| V.C.L) makes its appearance. This, starting
| |
| from the sinus venosus not far from the heart,
| |
| is on the fifth day a short trunk running backward in the middle line below the aorta, and
| |
| speedily losing itself in the tissues of the
| |
| Wolffian bodies. When the true kidneys are
| |
| formed it also receives blood from them, and
| |
| thenceforward enlarging rapidly becomes the
| |
| channel by which the greater part of the blood
| |
| from the hinder part of the body finds its way
| |
| to the heart. In proportion as the vena cava
| |
| inferior increases in size, the posterior cardinal
| |
| veins diminish.
| |
| | |
| The blood originally coming to them from
| |
| the posterior part of the spinal cord and trunk
| |
| is transported into two posterior vertebral veins,
| |
| similar to those in Reptilia, which are however
| |
| placed dorsally to the heads of the ribs, and
| |
| join the anterior vertebral veins. With their
| |
| appearance the anterior parts of the posterior
| |
| cardinals disappear. The blood from the hind
| |
| limbs becomes transported directly through the
| |
| kidney into the vena cava inferior, without
| |
| forming a renal portal system 1 .
| |
| | |
| On the third day the course of the vessels from the yolk-sack is very
| |
| simple. The two vitelline veins, of which the right is already the smaller,
| |
| form the ductus venosus, from which, as it passes through the liver on its
| |
| way to the heart, are given off the two sets of vena advehentes and vena
| |
| revehentes (fig. 371).
| |
| | |
| With the appearance of the allantois on the fourth day, a new feature is
| |
| introduced. From the ductus venosus there is given off a vein which
| |
| quickly divides into two branches. These, running along the ventral walls
| |
| of the body from which they receive some amount of blood, pass to the
| |
| allantois. They are the allantoic veins (fig. 371, U] homologous with the
| |
| anterior abdominal vein of the lower types. They unite in front to form a
| |
| single vein, which becomes, by reason of the rapid growth of the allantois,
| |
| very long. The right branch soon diminishes in size and finally disappears.
| |
| Meanwhile the left on reaching the allantois bifurcates ; and, its two
| |
| | |
| | |
| | |
| FIG. 371. DIAGRAM OF
| |
| THE VENOUS CIRCULATION
| |
| IN THE CHICK AT THE COMMENCEMENT OF THE FIFTH
| |
| | |
| DAY.
| |
| | |
| H. heart ; d. c. ductus Cuvieri. Into the ductus Cuvieri
| |
| of each side fall/, the jugular
| |
| vein, W. the vein from the
| |
| wing, and c. the inferior cardinal vein ; S. V. sinus venosus ;
| |
| Of. vitelline vein ; U. allantoic vein, which at this stage
| |
| gives off branches to the bodywalls ; V.C.l. inferior vena
| |
| cava ; /. liver.
| |
| | |
| | |
| | |
| The mode in which this is effected requires further investigation.
| |
| | |
| 42 2
| |
| | |
| | |
| | |
| 66o
| |
| | |
| | |
| | |
| VEINS OF THE CHICK.
| |
| | |
| | |
| | |
| | |
| branches becoming large and conspicuous, there still appear to be two
| |
| main allantoic veins. At its first appearance the allantoic vein seems to be
| |
| but a small branch of the vitelline, but as the allantois grows rapidly,
| |
| and the yolk-sack dwindles, this state of things is reversed, and the less conspicuous vitelline appears as a branch of the larger allantoic vein.
| |
| | |
| On the third day the blood returning from the walls of the intestine is
| |
| insignificant in amount. As however the
| |
| intestine becomes more and more developed, it acquires a distinct venous system,
| |
| and its blood is returned by veins which
| |
| form a trunk, the mesenteric vein (fig. 372,
| |
| M") falling into the vitelline vein at its
| |
| junction with the allantoic vein.
| |
| | |
| These three great veins, in fact, form a
| |
| large common trunk, which enters at once
| |
| into the liver, and which we may now call
| |
| the portal vein (fig. 372, P. V}. This, at its
| |
| entrance into the liver, partly breaks up
| |
| into the vena advehentes, and partly continues as the ductus venosus (D.V}
| |
| straight through the liver, emerging from
| |
| which it joins the vena cava inferior. Before
| |
| the establishment of the vena cava inferior,
| |
| the venas revehentes, carrying back the
| |
| blood which circulates through the hepatic
| |
| capillaries, join the ductus venosus close to
| |
| its exit from the liver. By the time however that the vena cava has become a large
| |
| and important vessel it is found that the
| |
| venae revehentes, or as we may now call
| |
| them the hepatic veins, have shifted their
| |
| embouchment, and now fall directly into
| |
| that vein, the ductus venosus making a separate junction rather higher up (fig. 372).
| |
| | |
| This state of things continues with but slight changes till near the end
| |
| of incubation, when the chick begins to breathe the air in the air-chamber
| |
| of the shell, and respiration is no longer carried on by the allantois. Blood
| |
| then ceases to flow along the allantoic vessels ; they become obliterated.
| |
| The vitelline vein, which as the yolk becomes gradually absorbed proportionately diminishes in size and importance, comes to appear as a mere
| |
| branch of the portal vein. The ductus venosus becomes obliterated ; and
| |
| hence the whole of the blood coming through the portal vein flows into the
| |
| substance of the liver, and so by the hepatic veins into the vena cava.
| |
| | |
| Although the allantoic (anterior abdominal) vein is obliterated in the
| |
| adult, there is nevertheless established an anastomosis between the portal
| |
| system and the veins bringing the blood from the limbs to the vena cava
| |
| | |
| | |
| | |
| FIG. 372. DIAGRAM OF THE
| |
| VENOUS CIRCULATION IN THE
| |
| CHICK DURING THE LATER DAYS
| |
| OF INCUBATION.
| |
| | |
| H. heart ; V.S.R. right vena
| |
| cava superior; V.S.L. left vena cava
| |
| superior. The two venas cavrc
| |
| superiores are the original 'ductus
| |
| Cuvieri,' they open into the sinus
| |
| venosus. J. jugular vein; Su.V.
| |
| anterior vertebral vein ; In. V. inferior vertebral vein ; W. subclavian; V.C.I, vena cava inferior;
| |
| D. V. ductus venosus ; P. V. portal
| |
| vein ; M. mesenteric vein bringing
| |
| blood from the intestines into the
| |
| portal vein ; O.f. vitelline vein ; U.
| |
| allantoic vein. The three last mentioned veins unite together to form
| |
| the portal vein ; /. liver.
| |
| | |
| | |
| | |
| THE VENOUS SYSTEM.
| |
| | |
| | |
| | |
| 66l
| |
| | |
| | |
| | |
| inferior, in that the caudal vein and posterior pelvic veins open into a
| |
| vessel, known as the coccygeo-mesenteric vein, which joins the portal
| |
| vein ; while at the same time the posterior pelvic veins are connected with
| |
| the common iliac veins by a vessel which unites with them close to their
| |
| junction with the coccygeo-mesenteric vein.
| |
| | |
| Mammalia. In Mammals the same venous trunks are developed in
| |
| the embryo as in other types (fig. 373 A). The anterior cardinals or
| |
| external jugulars form the primitive veins of the anterior part of the body,
| |
| and the internal jugulars and anterior vertebrals are subsequently formed.
| |
| The subclavians (fig. 373 A, j), developed on the formation of the anterior
| |
| limbs, also pour their blood into these primitive trunks. In the lower
| |
| Mammalia (Monotremata, Marsupialia, Insectivora, some Rodentia, etc.,
| |
| the two ductus Cuvieri remain as the two superior venae cavae, but more
| |
| usually an anastomosis arises between the right and left innominate veins,
| |
| and eventually the whole of the blood of the left superior cava is carried to
| |
| the right side, and there is left only a single superior cava (fig. 373 B and C).
| |
| | |
| | |
| | |
| | |
| | |
| F IG - 373- DIAGRAM OF THE DEVELOPMENT OF THE PAIRED VENOUS SYSTEM OF
| |
| | |
| MAMMALS (MAN). (From Gegenbaur.)
| |
| | |
| j. jugular vein ; cs. vena cava superior; s. subclavian veins; c. posterior cardinal
| |
| vein ; v. vertebral vein ; az. azygos vein ; cor. coronary vein.
| |
| | |
| A. Stage in which the cardinal veins have already disappeared. Their position
| |
| is indicated by dotted lines.
| |
| | |
| B. Later stage when the blood from the left jugular vein is carried into the right
| |
| to form the single vena cava superior ; a remnant of the left superior cava being however still left.
| |
| | |
| C. Stage after the left vertebral vein has disappeared; the right vertebral
| |
| remaining as the azygos vein. The coronary vein remains as the last remnant of the
| |
| left superior vena cava.
| |
| | |
| A small rudiment of the left superior cava remains however as the sinus
| |
| coronartus and receives the coronary vein from the heart (figs. 373 C,
| |
| cor and 374, cs).
| |
| | |
| The posterior cardinal veins form at first the only veins receiving the
| |
| | |
| | |
| | |
| 662
| |
| | |
| | |
| | |
| THE VEINS OF MAMMALIA.
| |
| | |
| | |
| | |
| blood from the posterior part of the trunk and kidneys ; and on the
| |
| development of the hind limbs receive the blood from them also.
| |
| | |
| As in the types already described
| |
| an unpaired vena cava inferior becomes
| |
| eventually developed, and gradually
| |
| carries off a larger and larger portion
| |
| of the blood originally returned by the
| |
| posterior cardinals. It unites with the
| |
| common stem of the allantoic and
| |
| vitelline veins in front of the liver.
| |
| | |
| At a later period a pair of trunks
| |
| is established bringing the blood from
| |
| the posterior part of the cardinal veins
| |
| and the crural veins directly into the
| |
| vena cava inferior (fig. 374, il}. These
| |
| vessels, whose development has not
| |
| been adequately investigated, form the
| |
| common iliac veins, while the posterior
| |
| ends of the cardinal veins which join
| |
| them become the hypogastric veins (fig.
| |
| 374, hy). Owing to the development of
| |
| the common iliac veins there is no renal
| |
| portal system like that of the Reptilia
| |
| and Amphibia.
| |
| | |
| Posterior vertebral veins, similar to
| |
| those of Reptilia and Birds, are established in connection with the intercostal
| |
| and lumbar veins, and unite anteriorly
| |
| with the front part of the posterior
| |
| | |
| | |
| | |
| | |
| FIG. 374. DIAGRAM OF THE CHIEF
| |
| | |
| VENOUS TRUNKS OF MAN. (From
| |
| Gegenbaur.)
| |
| | |
| cs. vena cava superior ; s. subclavian vein ; ji. internal jugular ; je.
| |
| external jugular ; az. azygos vein ; ha.
| |
| hemiazygos vein ; c. clotted line shewing previous position of cardinal veins ;
| |
| ci. vena cava inferior ; r. renal veins ;
| |
| il. iliac ; hy. hypogastric veins ; h.
| |
| hepatic veins.
| |
| | |
| The dotted lines shew the position
| |
| of embryonic vessels aborted in the
| |
| adult.
| |
| | |
| | |
| | |
| cardinal veins (fig. 373 A) 1 .
| |
| | |
| On the formation of the posterior vertebral veins, and as the inferior
| |
| vena cava becomes more important, the middle part of the posterior cardinals becomes completely aborted (fig. 374, f), the anterior and posterior
| |
| parts still persisting, the former as the continuations of the posterior
| |
| vertebrals into the anterior vena cava (az\ the latter as the hypogastric veins
| |
| (Ay).
| |
| | |
| Though in a few Mammalia both the posterior vertebrals persist, a
| |
| transverse connection is usually established between them, and the one (the
| |
| right) becoming the more important constitutes the azygos vein (fig. 374, az),
| |
| the persisting part of the left forming the hemiazygos vein (ha}.
| |
| | |
| The remainder of the venous system is formed in the embryo of the
| |
| vitelline and allantoic veins, the former being eventually joined by the
| |
| mesenteric vein so as to constitute the portal vein.
| |
| | |
| 1 Rathke, as mentioned above, holds that in the Snake the front part of the
| |
| posterior cardinals completely aborts. Further investigations are required to shew
| |
| whether there really is a difference between Mammalia and Reptilia in this matter.
| |
| | |
| | |
| | |
| | |
| | |
| | |
| THE VENOUS SYSTEM. 663
| |
| | |
| The vitelline vein is the first part of this system established, and divides
| |
| near the heart into two veins bringing back the blood from the yolk-sack
| |
| (umbilical vesicle). The right vein soon however aborts.
| |
| | |
| The allantoic (anterior abdominal) veins are originally paired. They
| |
| are developed very early, and at first course along the still widely open
| |
| somatic walls of the body, and fall into the single vitelline trunk in front.
| |
| The right allantoic vein disappears before long, and the common trunk
| |
| formed by the junction of the vitelline and allantoic veins becomes considerably elongated. This trunk is soon enveloped by the liver.
| |
| | |
| The succeeding changes have been somewhat differently described by
| |
| Kolliker and Rathke. According to the former the common trunk of the
| |
| allantoic and vitelline veins in its passage through the liver gives off
| |
| branches to the liver, and also receives branches from this organ near its
| |
| anterior exit. The main trunk is however never completely aborted, as in
| |
| the embryos of other types, but remains as the ductus venosus Arantii.
| |
| | |
| With the development of the placenta the allantoic vein becomes the
| |
| main source of the ductus venosus, and the vitelline or portal vein, as it may
| |
| perhaps be now conveniently called, ceases to join it directly, but falls into
| |
| one of its branches in the liver.
| |
| | |
| The vena cava inferior joins the continuation of the ductus venosus in
| |
| front of the liver, and, as it becomes more important, it receives directly
| |
| the hepatic veins which originally brought back blood into the ductus
| |
| venosus. The ductus venosus becomes moreover merely a small branch of
| |
| the vena cava.
| |
| | |
| At the close of foetal life the allantoic vein becomes obliterated up to its
| |
| place of entrance into the liver ; the ductus venosus becomes a solid cord
| |
| the so-called round ligament and the whole of the venous blood is brought
| |
| to the liver by the portal vein 1 .
| |
| | |
| Owing to the allantoic (anterior abdominal) vein having merely a fcetal
| |
| existence an anastomosis between the iliac veins and the portal system by
| |
| means of the anterior abdominal vein is not established.
| |
| | |
| | |
| | |
| BIBLIOGRAPHY of the Venous System.
| |
| | |
| (498) J. Marshall. "On the development of the great anterior veins." Phil.
| |
| Trans., 1859.
| |
| | |
| (499) H. Rathke. " Ueb. d. Bildung d. Pfortader u. d. Lebervenen b. Saugethieren." MeckeVs Archiv, 1830.
| |
| | |
| (500) H. Rathke. "Ueb. d. Bau u. d. Entwick. d. Venensystems d. Wirbelthiere." Bericht. Jib. d. natttrh. Seminar, d. Univ. Konigsberg, 1838.
| |
| | |
| Vide also Von Baer (No. 291), Gotte (No. 296), Kolliker (No. 298), and Rathke
| |
| (Nos. 299, 300, and 301).
| |
| | |
| 1 According to Rathke the original trunk connecting the allantoic vein directly
| |
| with the heart through the liver is aborted, and the ductus venosus Arantii is a
| |
| secondary connection established in the latter part of foetal life.
| |
| | |
| | |
| | |
| 664 LYMPHATIC SYSTEM.
| |
| | |
| | |
| | |
| Lymphatic System.
| |
| | |
| The lymphatic system arises from spaces in the general parenchyma of
| |
| the body, independent in their origin of the true body cavity, though communicating both with this cavity and with the vascular system.
| |
| | |
| In all the true Vertebrata certain parts of the system form definite trunks
| |
| communicating with the venous system ; and in the higher types the walls of
| |
| the main lymphatic trunks become quite distinct.
| |
| | |
| But little is known with reference to the ontogeny of the lymphatic vessels,
| |
| but they originate late in larval life, and have at first the form of simple
| |
| intercellular spaces.
| |
| | |
| The lymphatic glands appear to originate from lymphatic plexuses, the
| |
| cells of which produce lymph corpuscles. It is only in Birds and Mammals,
| |
| and especially in the latter, that the lymphatic glands form definite structures.
| |
| | |
| The Spleen. The spleen, from its structure, must be classed with the
| |
| lymphatic glands, though it has definite relations to the vascular system.
| |
| It is developed in the mesoblast of the mesogastrium, usually about the
| |
| same time and in close connection with the pancreas.
| |
| | |
| According to Miiller and Peremeschko the mass of mesoblast which
| |
| forms the spleen becomes early separated by a groove on the one side from
| |
| the pancreas and on the other from the mesentery. Some of its cells
| |
| become elongated, and send out processes which uniting with like processes
| |
| from other cells form the trabecular system. From the remainder of the
| |
| tissue are derived the cells of the spleen pulp, which frequently contain more
| |
| than one nucleus. Especial accumulations of these cells take place at a
| |
| later period to form the so-called Malpighian corpuscles of the spleen.
| |
| | |
| BIBLIOGRAPHY of Spleen.
| |
| | |
| (501) W. Miiller. "The Spleen." Strieker's Histology.
| |
| | |
| (502) Peremeschko. " Ueb. d. Entwick. d. Milz." Sitz. d. Wuti. Akad.
| |
| Wiss., Vol. LVI. 1867.
| |
| | |
| Suprarenal ^bodies.
| |
| | |
| In Elasmobranch Fishes two distinct sets of structures are found, both of
| |
| which have been called suprarenal bodies. As shewn in the sequel both of
| |
| these structures probably unite in the higher types to form the suprarenal
| |
| bodies.
| |
| | |
| One of them consists of a series of paired bodies, situated on the
| |
| branches of the dorsal aorta, segmentally arranged, and forming a chain
| |
| extending from close behind the heart to the hinder end of the body cavity.
| |
| Each body is formed of a series of lobes, and exhibits a well-marked
| |
| distinction into a cortical layer of columnar cells, and a medullary substance
| |
| formed of irregular polygonal cells. As first shewn by Leydig, they are
| |
| | |
| | |
| | |
| SUPRARENAL BODIES. 665
| |
| | |
| closely connected with the sympathetic ganglia, and usually contain numerous
| |
| ganglion cells distributed amongst the proper cells of the body.
| |
| | |
| The second body consists of an unpaired column of cells placed between
| |
| the dorsal aorta and unpaired caudal vein, and bounded on each side by the
| |
| posterior parts of the kidney. I propose to call it the interrenal body.
| |
| In front it overlaps the paired suprarenal bodies, but does not unite with
| |
| them. It is formed of a series of well-marked lobules, etc. In the fresh
| |
| state Leydig (No. 506) finds that "fat molecules form the chief mass of the
| |
| body, and one finds freely imbedded in them clear vesicular nuclei." As
| |
| may easily be made out from hardened specimens it is invested by a tunica
| |
| propria, which gives off septa dividing it into well-marked areas filled with
| |
| polygonal cells. These cells constitute the true parenchyma of the body.
| |
| By the ordinary methods of hardening, the oil globules, with which they are
| |
| filled in the fresh state, completely disappear.
| |
| | |
| The paired suprarenal bodies (Balfour, No. 292, pp. 242 244) are developed from the sympathetic ganglia. These ganglia, shewn in an early
| |
| stage in fig. 380, sy.g, become gradually divided into a ganglionic part and a
| |
| glandular part. The former constitutes the sympathetic ganglia of the adult ;
| |
| the latter the true paired suprarenal bodies. The interrenal body is however
| |
| developed (Balfour, No. 292, pp. 245 247) from indifferent mesoblast cells
| |
| between the two kidneys, in the same situation as in the adult.
| |
| | |
| The development of the suprarenal bodies in the Amniota has been most
| |
| fully studied by Braun (No. 503) in the Reptilia.
| |
| | |
| In Lacertilia they consist of a pair of elongated yellowish bodies, placed
| |
| between the vena renalis revehens and the generative glands.
| |
| | |
| They are formed of two constituents, viz. (i) masses of brown cells placed
| |
| on the dorsal side of the organ, which stain deeply with chromic acid, like
| |
| certain of the cells of the suprarenals of Mammalia, and (2) irregular cords,
| |
| in part provided with a lumen, filled with fat-like globules l , amongst which
| |
| are nuclei. On treatment with chromic acid the fat globules disappear, and
| |
| the cords break up into bodies resembling columnar cells.
| |
| | |
| The dorsal masses of brown cells are developed from the sympathetic
| |
| ganglia in the same way as the paired suprarenal bodies of the Elasmobranchii, while the cords filled with fat-like globules are formed of indifferent
| |
| mesoblast cells as a thickening in the lateral walls of the inferior vena cava,
| |
| and the cardinal veins continuous with it. The observations of Brunn (No.
| |
| 504) on the Chick, and Kolliker (No. 298, pp. 953955) n the Mammal,
| |
| add but little to those of Braun. They shew that the greater part of the
| |
| gland (the cortical substance) in these two types is derived from the mesoblast,
| |
| and that the glands are closely connected with sympathetic ganglia ; while
| |
| Kolliker also states that the posterior part of the organ is unpaired in the
| |
| embryo rabbit of 1 6 or 17 days.
| |
| | |
| The structure and development of what I have called the interrenal body
| |
| | |
| 1 These globules are not formed of a true fatty substance, and this is also probably
| |
| true for the similar globules of the interrenal bodies of Elasmobranchii.
| |
| | |
| | |
| | |
| 666 SUPRARENAL BODIES.
| |
| | |
| in Elasmobranchii so closely correspond with that of the mesoblastic part of
| |
| the suprarenal bodies of the Reptilia, that I have very little hesitation in
| |
| regarding them as homologous 1 ; while the paired bodies in Elasmobranchii,
| |
| derived from the sympathetic ganglia, clearly correspond with the part of the
| |
| suprarenals of Reptilia having a similar origin ; although the anterior parts
| |
| of the paired suprarenal bodies of Fishes have clearly become aborted in the
| |
| higher types.
| |
| | |
| In Elasmobranch Fishes we thus have (i) a series of paired
| |
| bodies, derived from the sympathetic ganglia, and (2) an unpaired body of mesoblastic origin. In the Amniota these bodies
| |
| unite to form the compound suprarenal bodies, the two constituents of which remain, however, distinct in their development.
| |
| The mesoblastic constituent appears to form the cortical part of
| |
| the adult suprarenal body, and the nervous constituent the
| |
| medullary part.
| |
| | |
| BIBLIOGRAPHY of the Suprarenal bodies,
| |
| | |
| (503) M. Braun. "Bau u. Entwick. d. Nebennieren bei Reptilien. " Arbeit,
| |
| a. d. zool.-zoot. Institut Wurzlttrg, Vol. V. 1879.
| |
| | |
| (504) A. v. Brunn. "Ein Beitrag z. Kenntniss d. feinern Baues u. d. Entwick.
| |
| d. Nebennieren." Archiv f. mikr. Anat., Vol. VIII. 1872.
| |
| | |
| (505) Fr. Leydig. Untersiich. iib. Fische u. fieptilten. Berlin, 1853.
| |
| | |
| (506) Fr. Leydig. Rochen u. Haie. Leipzig, 1852.
| |
| | |
| Vide also F. M. Balfour (No. 292), Kolliker (No. 298), Remak (No. 302), etc.
| |
| | |
| 1 The fact of the organ being unpaired in Elasmobranchii and paired in the
| |
| Amniota is of no importance, as is shewn by the fact that part of the organ is unpaired
| |
| in the Rabbit.
| |
| | |
| | |
| | |
| CHAPTER XXII.
| |
| | |
| | |
| | |
| THE MUSCULAR SYSTEM.
| |
| | |
| | |
| | |
| | |
| IN all the Ccelenterata, except the Ctenophora, the contractile elements of the body wall consist of filiform processes of
| |
| ectodermal or entodermal epithelial cells (figs. 375 and 376 B).
| |
| The elements provided with these processes, which were first
| |
| discovered by Kleinenberg, are known as myo-epithelial
| |
| cells. Their contractile parts may either be striated (fig. 376)
| |
| or non-striated (fig. 375). In some
| |
| instances the epithelial part of the
| |
| cell may nearly abort, its nucleus
| |
| alone remaining (fig. 376 A) ; and
| |
| in this way a layer of muscles lying
| |
| completely below the surface may
| |
| be established.
| |
| | |
| There is embryological evidence
| |
| of the derivation of the voluntary
| |
| muscular system of a large number of types from myo-epithelial
| |
| cells of this kind. The more important of these groups are the
| |
| Chaetopoda, the Gephyrea, the Chaetognatha, the Nematoda, and
| |
| the Vertebrata 1 .
| |
| | |
| While there is clear evidence that the muscular system of a
| |
| large number of types is composed of cells which had their
| |
| origin in myo-epithelial cells, the mode of evolution of the
| |
| | |
| 1 If recent statements of Metschnikoff are to be trusted, the Echinodermata must
| |
| be added to these groups. The amoeboid cells stated in the first volume of this
| |
| treatise to form the muscles in this group, on the authority of Selenka, give rise,
| |
| according to Metschnikoff, only to the cutis, while the same naturalist states the
| |
| epithelial cells of the vasoperitoneal vesicles are provided with muscular tails.
| |
| | |
| | |
| | |
| FIG. 375. MYO-EPITHELIAL
| |
| CELLS OF HYDRA. (From Gegenbaur ; after Kleinenberg.)
| |
| | |
| m. contractile fibres.
| |
| | |
| | |
| | |
| 668 THE MUSCULAR FIBRES.
| |
| | |
| muscular system of other types is still very obscure. The
| |
| muscles may arise in the embryo from amoeboid or indifferent
| |
| cells, and the Hertwigs 1 hold that in many of these instances the
| |
| muscles have also phylogenetically taken their origin from
| |
| indifferent connective-tissue cells. The subject is however beset
| |
| with very serious difficulties, and to discuss it here would carry
| |
| me too far into the region of pure histology.
| |
| | |
| The voluntary muscular system of the CJiordata.
| |
| | |
| The muscular fibres. The muscular elements of the
| |
| Chordata undoubtedly belong to the myo-epithelial type. The
| |
| embryonic muscle-cells are at first simple epithelial cells, but
| |
| | |
| | |
| | |
| | |
| FIG. 376. MUSCLE-CELLS OF LIZZIA KOLLIKERI. (From Lankester ; after
| |
| O. and R. Hertwig.)
| |
| | |
| A. Muscle-cell from the circular fibres of the subumbrella.
| |
| | |
| B. Myo-epithelial cells from the base of a tentacle.
| |
| | |
| soon become spindle-shaped : part of their protoplasm becomes
| |
| differentiated into longitudinally placed striated muscular fibrils,
| |
| while part, enclosing the nucleus, remains indifferent, and constitutes the epithelial element of the cells. The muscular
| |
| fibrils are either placed at one side of the epithelial part of the
| |
| cell, or in other instances (the Lamprey, the Newt, the Sturgeon,
| |
| the Rabbit) surround it. The latter arrangement is shewn for
| |
| the Sturgeon in fig. 57.
| |
| | |
| The number of the fibrils of each cell gradually increases,
| |
| and the protoplasm diminishes, so that eventually only the
| |
| nucleus, or nuclei resulting from its division, are left. The
| |
| products of each cell probably give rise, in conjunction with a
| |
| further division of the nucleus, to a primitive bundle, which,
| |
| | |
| 1 O. and R. Hertwig, Die Calomthcorie. Jena, 1881.
| |
| | |
| | |
| | |
| THE MUSCULAR SYSTEM.
| |
| | |
| | |
| | |
| 669
| |
| | |
| | |
| | |
| t>r
| |
| | |
| | |
| | |
| | |
| except in Amphioxus, Petromyzon, etc., is surrounded by a
| |
| special investment of sarcolemma.
| |
| | |
| The voluntary muscular system. For the purposes of
| |
| description the muscular system of the Vertebrata may conveniently be divided into two sections, viz. that of the head and
| |
| that of the trunk. The main part, if
| |
| not the whole, of the muscular system
| |
| of the trunk is derived from certain
| |
| structures, known as the muscle-plates,
| |
| which take their origin from part of
| |
| the primitive mesoblastic somites.
| |
| | |
| It has already been stated (pp.
| |
| 292 ^296) that the mesoblastic somites
| |
| are derived from the dorsal segmented
| |
| part of the primitive mesoblastic plates.
| |
| Since the history of these bodies is
| |
| presented in its simplest form in Elasmobranchii it will be convenient to
| |
| commence with this group. Each
| |
| somite is composed of two layers a
| |
| somatic and a splanchnic both formed
| |
| of a single row of columnar cells.
| |
| Between these two layers is a cavity,
| |
| which is at first directly continuous
| |
| with the general body cavity, of which
| |
| indeed it merely forms a specialised
| |
| part (fig. 377). Before long the cavity
| |
| becomes however completely constricted off from the permanent body cavity.
| |
| | |
| Very early (fig. 377) the inner or splanchnic wall of the
| |
| somites loses its simple constitution, owing to the middle part of
| |
| it undergoing peculiar changes. The meaning of the changes is
| |
| at once shewn by longitudinal horizontal sections, which prove
| |
| (% 378) that the cells in this situation (mp') have become
| |
| extended in a longitudinal direction, and, in fact, form typical
| |
| spindle-shaped embryonic muscle-cells, each with a large
| |
| nucleus. Every muscle-cell extends for the whole length of a
| |
| somite. The inner layer of each somite, immediately within
| |
| the muscle-band just described, begins to proliferate, and produce
| |
| | |
| | |
| | |
| FIG. 377. TRANSVERSE
| |
| SECTION THROUGH THETRUNK
| |
| OF AN EMBRYO SLIGHTLY
| |
| OLDER THAN FIG. 28 E.
| |
| | |
| nc. neural canal ; pr. posterior root of spinal nerve ; x.
| |
| subnotochordal rod ; ao. aorta ;
| |
| sc. somatic mesoblast ; sf>.
| |
| splanchnic mesoblast ; mp.
| |
| muscle-plate ; mp', portion of
| |
| muscle-plate converted into
| |
| muscle ; Vr. portion of the
| |
| vertebral plate which will give
| |
| rise to the vertebral bodies ; al.
| |
| alimentary tract.
| |
| | |
| | |
| | |
| THE MUSCLE-PLATES.
| |
| | |
| | |
| | |
| a mass of cells, placed between the muscles and the notochord
| |
| ( Vr\ These cells form the commencing vertebral bodies, and
| |
| have at first (fig. 378) the same segmentation as the somites
| |
| from which they sprang.
| |
| | |
| After the separation of the vertebral bodies from the somites
| |
| the remaining parts of the somites may be called muscle-plates ;
| |
| since they become directly converted into the whole voluntary
| |
| muscular system of the trunk (fig. 379, mp}.
| |
| | |
| According to the statements of Bambeke and Go'tte, the Amphibians
| |
| present some noticeable peculiarities in the development of their muscular
| |
| system, in that such distinct muscle-plates as those of other vertebrate types
| |
| are not developed. Each side-plate of mesoblast is divided into a somatic
| |
| and a splanchnic layer, continuous throughout the vertebral and parietal
| |
| portions of the plate. The vertebral portions (somites) of the plates soon
| |
| become separated from the parietal, and form independent masses of cells
| |
| constituted of two layers, which were originally continuous with the
| |
| somatic and splanchnic layers of the parietal plates (fig. 79). The outer or
| |
| somatic layer of the vertebral plates is formed of a single row of cells, but
| |
| the inner or splanchnic layer is made up of a kernel of cells on the side of
| |
| the somatic layer and an inner layer. The kernel of the splanchnic layer
| |
| and the outer or somatic layer together correspond to a muscle- plate of other
| |
| Vertebrata, and exhibit a similar segmentation.
| |
| | |
| Osseous Fishes are stated to agree with Amphibians in the development
| |
| of their somites and muscular
| |
| system 1 , but further observations
| |
| on this point are required.
| |
| | |
| In Birds the horizontal splitting of the mesoblast extends at
| |
| first to the dorsal summit of the
| |
| mesoblastic plates, but after the
| |
| isolation of the somites the split
| |
| between the somatic and splanchnic layers becomes to a large extent obliterated, though in the anterior somites it appears in part
| |
| to persist. The somites on the
| |
| second day, as seen in a transverse section (fig. 115, P.?'.), are
| |
| somewhat quadrilateral in form
| |
| but broader than they are deep.
| |
| | |
| Each at that time consists of
| |
| a somewhat thick cortex of radi
| |
| | |
| | |
| | |
| FlG. 378. HORIZONTALSECTION THROUGH
| |
| THE TRUNK OF AN EMBRYO OF SCYLL1UM
| |
| CONSIDERABLY YOUNGER THAN 28 F.
| |
| | |
| | |
| | |
| The section is taken at the level of the
| |
| notochord, and shews the separation of the
| |
| cells to form the vertebral bodies from the
| |
| muscle-plates.
| |
| | |
| ch. notochord ; ep. epiblast ; Vr, rudiment
| |
| of vertebral body ; mp. muscle- plate ; mp' .
| |
| portion of muscle-plate already differentiated
| |
| into longitudinal muscles.
| |
| | |
| | |
| | |
| 1 Ehrlich, " Ueber den peripher. Theil d. Urwirbel." Archiv f. mikr. Anal.,
| |
| Vol. XI.
| |
| | |
| | |
| | |
| THE MUSCULAR SYSTEM. 671
| |
| | |
| ating rather granular columnar cells, enclosing a small kernel of spherical
| |
| cells. They are not, as may be seen in the above figure, completely
| |
| separated from the ventral (or lateral as they are at this period) parts of the
| |
| mesoblastic plate, and the dorsal and outer layer of the cortex of the
| |
| somites is continuous with the somatic layer of mesoblast, the remainder of
| |
| the cortex, with the central kernel, being continuous with the splanchnic
| |
| layer. Towards the end of the second and beginning of the third day the
| |
| upper and outer layer of the cortex, together probably with some of the
| |
| central cells of the kernel, becomes separated off as a muscle-plate (fig. 1 16).
| |
| The muscle-plate when formed (fig. 117) is found to consist of two layers,
| |
| an inner and an outer, which enclose between them an almost obliterated
| |
| central cavity ; and no sooner is the muscle-plate formed than the middle
| |
| portion of the inner layer becomes converted into longitudinal muscles.
| |
| The avian muscle-plates have, in fact, precisely the same constitution as
| |
| those of Elasmobranchii. The central space is clearly a remnant of the
| |
| vertebral portion of the body cavity, which, though it wholly or partially
| |
| disappears in a previous stage, reappears again on the formation of the
| |
| muscle-plate.
| |
| | |
| The remainder of the somite, after the formation of the muscle-plate,
| |
| is of very considerable bulk ; the cells of the cortex belonging to it lose
| |
| their distinctive characters, and the major part of it becomes the vertebral
| |
| rudiment.
| |
| | |
| In Mammalia the history appears to be generally the same as in Elasmobranchii. The split which gives rise to the body cavity is continued to
| |
| the dorsal summit of the mesoblastic plates, and the dorsal portions of the
| |
| plates with their contained cavities become divided into somites, and are
| |
| then separated off from the ventral. The later development of the somites
| |
| has not been worked out with the requisite care, but it would seem that they
| |
| form somewhat cubical bodies in which all trace of the primitive slit is lost.
| |
| The further development resembles that in Birds.
| |
| | |
| The first changes of the mesoblastic somites and the formation of the muscle-plates do not, according to existing statements,
| |
| take place on quite the same type throughout the Vertebrata,
| |
| yet the comparison which has been instituted between Elasmobranchs and other Vertebrates appears to prove that there are
| |
| important common features in their development, which may be
| |
| regarded as primitive, and as having been inherited from the
| |
| ancestors of Vertebrates. These features are (i) the extension
| |
| of the body cavity into the vertebral plates, and subsequent
| |
| enclosure of this cavity between the two layers of the muscleplates ; (2) the primitive division of the vertebral plate into an
| |
| outer (somatic) and an inner (splanchnic) layer, and the formation
| |
| of a large part of the voluntary muscular system out of the inner
| |
| | |
| | |
| | |
| THE MUSCLE-PLATES.
| |
| | |
| | |
| | |
| sp.c
| |
| | |
| | |
| | |
| layer, which in all cases is converted into muscles earlier than
| |
| the outer layer.
| |
| | |
| The conversion of the muscle-plates into muscles. It
| |
| | |
| will be convenient to commence this subject with a description
| |
| of the changes which take place in
| |
| such a simple type as that of the
| |
| Elasmobranchii.
| |
| | |
| At the time when the muscleplates have become independent
| |
| structures they form flat two-layered
| |
| oblong bodies enclosing a slit-like
| |
| central cavity (fig. 379, mp). The
| |
| outer or somatic wall is formed of
| |
| simple epithelial -like cells. The
| |
| inner or splanchnic wall has however a somewhat complicated structure. It is composed dorsally and
| |
| ventrally of a columnar epithelium,
| |
| but in its middle portion of the
| |
| muscle-cells previously spoken of.
| |
| Between these and the central cavity
| |
| of the plates the epithelium forming
| |
| the remainder of the layer commences to insert itself; so that between the first-formed muscle and
| |
| the cavity of the muscle-plate there
| |
| appears a thin layer of cells, not
| |
| however continuous throughout.
| |
| | |
| When first formed the muscleplates, as viewed from the exterior,
| |
| have nearly straight edges ; soon
| |
| however they become bent in the middle, so that the edges have
| |
| an obtusely angular form, the apex of the angle being directed
| |
| forwards. They are so arranged that the anterior edge of the
| |
| one plate fits into the posterior edge of the one in front. In the
| |
| lines of junction between the plates layers of connective-tissue
| |
| cells appear, which form the commencements of the intermuscular
| |
| septa.
| |
| | |
| The growth of the plates is very rapid, and their upper ends
| |
| | |
| | |
| | |
| | |
| FIG. 379. SECTION THROUGH
| |
| THE TRUNK OF A SCYLLIUM EMBRYO SLIGHTLY YOUNGER THAN
| |
| | |
| 28 F.
| |
| | |
| sp.c. spinal canal ; W. white
| |
| matter of spinal cord ; pr. posterior nerve-roots ; ch. notochord ;
| |
| x. sub-notochordal rod ; ao. aorta ;
| |
| mp. muscle-plate; mp' . inner layer
| |
| of muscle-plate already converted
| |
| into muscles ; Vr. rudiment of
| |
| vertebral body ; si. segmental
| |
| tube ; sd. segmental duct ; sp.v.
| |
| spiral valve ; z/. subintestinal vein ;
| |
| P.O. primitive generative cells.
| |
| | |
| | |
| | |
| THE MUSCULAR SYSTEM. 673
| |
| | |
| soon extend to the summit of the neural canal, and their lower
| |
| ones nearly meet in the median ventral line. The original band
| |
| of muscles, whose growth at first is very slow, now increases
| |
| with great rapidity, and forms the nucleus of the whole voluntary muscular system (fig. 380, mp'). It extends upwards and
| |
| downwards by the continuous conversion of fresh cells of the
| |
| splanchnic layer into muscle-cells. At the same time it grows
| |
| rapidly in thickness by the addition of fresh spindle-shaped
| |
| muscle-cells from the somatic layer as well as by the division of
| |
| the already existing cells.
| |
| | |
| Thus both layers of the muscle-plate are concerned in forming
| |
| the great longitudinal lateral muscles, though the splanchnic layer
| |
| is converted into muscles very much sooner than the somatic 1 .
| |
| | |
| Each muscle-plate is at first a continuous structure, extending
| |
| from the dorsal to the ventral surface, but after a time it becomes
| |
| divided by a layer of connective tissue, which becomes developed
| |
| nearly on a level with the lateral line, into a dorso-lateral and
| |
| a ventro-lateral section. The ends of the muscle-plates
| |
| continue for a long time to be formed of undifferentiated
| |
| columnar cells. The complicated outlines of the inter-muscular
| |
| septa become gradually established during the later stages of
| |
| development, causing the well-known appearances of the muscles
| |
| in transverse sections, which require no special notice here.
| |
| | |
| The muscles of the limbs. The limb muscles are formed
| |
| in Elasmobranchii, coincidently with the cartilaginous skeleton,
| |
| as two bands of longitudinal fibres on the dorsal and ventral
| |
| surfaces of the limbs (fig. 346). The cells, from which these
| |
| muscles originate, are derived from the muscle-plates. When
| |
| the ends of the muscle-plates reach the level of the limbs they
| |
| bend outwards and enter the tissue of the limbs (fig. 380).
| |
| Small portions of several muscle-plates (m.pl) come in this way
| |
| to be situated within the limbs, and are very soon segmented
| |
| off from the remainder of the muscle-plates. The portions of
| |
| the muscle-plates thus introduced soon lose their original dis
| |
| 1 The brothers Hertwig have recently maintained that only the inner layer of the
| |
| muscle-plates is converted into muscles. In the Elasmobranchs it is easy to demonstrate the incorrectness of this view, and in Acipenser (vide fig. 57, mp) the two layers
| |
| of the muscle-plate retain their original relations after the cells of both of them have
| |
| become converted into muscles.
| |
| | |
| B. in. 43
| |
| | |
| | |
| | |
| 674
| |
| | |
| | |
| | |
| THE MUSCLE-PLATES.
| |
| | |
| | |
| | |
| 3,-n,
| |
| | |
| | |
| | |
| | |
| FIG. 380. TRANSVERSE SECTION THROUGH THE ANTERIOR PART OF THE TRUNK
| |
| OF AN EMBRYO OF SCYLLIUM SLIGHTLY OLDER THAN FIG. 29 B.
| |
| | |
| The section is diagrammatic in so far that the anterior nerve-roots have been
| |
| inserted for the whole length ; whereas they join the spinal cord half-way between
| |
| two posterior roots.
| |
| | |
| sp.c. spinal cord; sp.g. ganglion of posterior root; ar. anterior root; dn. dorsally
| |
| directed nerve springing from posterior root; nip. muscle-plate; mp'. part of muscleplate already converted into muscles; vi.pl. part of muscle-plate which gives rise to
| |
| the muscles of the limbs; /. nervus lateralis; ao. aorta; ch. notochord; sy.g. sympathetic ganglion; ca.v. cardinal vein; sp.n. spinal nerve; sd. segmental (archinephric)
| |
| duct; st. segmental tube; du. duodenum; pan. pancreas; hp.d. point of junction of
| |
| hepatic duct with duodenum ; umc. umbilical canal.
| |
| | |
| | |
| | |
| THE MUSCULAR SYSTEM. 675
| |
| | |
| tinctness. There can however be but little doubt that they
| |
| supply the tissue for the muscles of the limbs. The muscleplates themselves, after giving off buds to the limbs, grow
| |
| downwards, and soon cease to shew any trace of having given
| |
| off these buds.
| |
| | |
| In addition to the longitudinal muscles of the trunk just described,
| |
| which are generally characteristic of Fishes, there is found in Amphioxus a
| |
| peculiar transverse abdominal muscle, extending from the mouth to the
| |
| abdominal pore, the origin of which has not been made out.
| |
| | |
| It has already been shewn that in all the higher Vertebrata
| |
| muscle-plates appear, which closely resemble those in Elasmobranchii; so that all the higher Vertebrata pass through, with
| |
| reference to their muscular system, a fish- like stage. The
| |
| middle portion of the inner layers of their muscle-plates becomes, as in Elasmobranchii, converted into muscles at a very
| |
| early period, and the outer layer for a long time remains formed
| |
| of indifferent cells. That these muscle-plates give rise to the
| |
| main muscular system of the trunk, at any rate to the episkeletal
| |
| muscles of Huxley, is practically certain, but the details of the
| |
| process have not been made out.
| |
| | |
| In the Perennibranchiata the fish-like arrangement of muscles is retained through life in the tail and in the dorso-lateral parts of the trunk.
| |
| In the tail of the Amniotic Vertebrata the primitive arrangement is also
| |
| more or less retained, and the same holds good for the dorso-lateral trunk
| |
| muscles of the Lacertilia. In the other Amniota and the Anura the
| |
| dorso-lateral muscles have become divided up into a series of separate
| |
| muscles, which are arranged in two main layers. It is probable that the
| |
| intercostal muscles belong to the same group as the dorso-lateral muscles.
| |
| | |
| The abdominal muscles of the trunk, even in the lowest Amphibia,
| |
| exhibit a division into several layers. The recti abdominis are the least
| |
| altered part of this system, and usually retain indications of the primitive
| |
| inter-muscular septa, which in many Amphibia and Lacertilia are also
| |
| to some extent preserved in the other abdominal muscles.
| |
| | |
| In the Amniotic Vertebrates there is formed underneath the vertebral
| |
| column and the transverse processes a system of muscles, forming part
| |
| of the hyposkeletal system of Huxley, and called by Gegenbaur the subvertebral muscles. The development of this system has not been worked
| |
| out, but on the whole I am inclined to believe that it is derived from
| |
| the muscle-plates. Kolliker, Huxley and other embryologists believe
| |
| however that these muscles are independent of the muscle-plates in their
| |
| origin.
| |
| | |
| 432
| |
| | |
| | |
| | |
| 676 THE HEAD-CAVITIES.
| |
| | |
| | |
| | |
| Whether the muscle of the diaphragm is to be placed in the same
| |
| category as the hyposkeletal muscles has not been made out.
| |
| | |
| It is probable that the cutaneous muscles of the trunk are derived
| |
| from the cells given off from the muscle-plates. Kolliker however believes
| |
| that they have an independent origin.
| |
| | |
| The limb-muscles, both extrinsic and intrinsic, as may be concluded
| |
| from their development in Elasmobranchii, are derived from the muscleplates. Kleinenberg found in Lacertilia a growth of the muscle-plates
| |
| into the limbs, and in Amphibia Gotte finds that the outer layer of the
| |
| muscle-plates gives rise to the muscles of the limbs.
| |
| | |
| In the higher Vertebrata on the other hand the entrance of the muscleplates into the limbs has not been made out (Kolliker). It seems therefore
| |
| probable that by an embryological modification, of which instances are so
| |
| frequent, the cells which give rise to the muscles of the limbs in the higher
| |
| Vertebrata can no longer be traced into a direct connection with the muscleplates.
| |
| | |
| TJte Somites and muscular system of the head.
| |
| | |
| The extension of the somites to the anterior end of the body
| |
| in Amphioxus clearly proves that somites, similar to those of
| |
| the trunk, were originally present in a region, which in the
| |
| higher Vertebrata has become differentiated into the head. In
| |
| the adult condition no true Vertebrate exhibits indications of
| |
| such somites, but in the embryos of several of the lower Vertebrata structures have been found, which are probably equivalent
| |
| to the somites of the trunk : they have been frequently alluded
| |
| to in the previous chapters of this volume. These structures
| |
| have been most fully worked out in Elasmobranchii.
| |
| | |
| The mesoblast in Elasmobranch embryos becomes first split
| |
| into somatic and splanchnic layers in the region of the head ;
| |
| and between these layers there are formed two cavities, one on
| |
| each side, which end in front opposite the blind anterior extremity of the alimentary canal ; and are continuous behind
| |
| with the general body-cavity (fig. 20 A, vp}. I propose calling
| |
| them the head-cavities. The cavities of the two sides have
| |
| no communication with each other.
| |
| | |
| Coincidently with the formation of an outgrowth from the
| |
| throat to form the first visceral cleft, the head-cavity on each
| |
| side becomes divided into a section in front of the cleft and a
| |
| section behind the cleft ; and at a later period it becomes, owing
| |
| to the formation of a second cleft, divided into three sections :
| |
| | |
| | |
| | |
| THE MUSCULAR SYSTEM.
| |
| | |
| | |
| | |
| 677
| |
| | |
| | |
| | |
| vn~.
| |
| | |
| | |
| | |
| | |
| (i) a section in front of the first or hyomandibular cleft; (2) a
| |
| section in the hyoid arch between the hyomandibular cleft and
| |
| the hyobranchial or first branchial cleft ; (3) a section behind
| |
| the first branchial cleft.
| |
| | |
| The front section of the head-cavity grows forward, and soon
| |
| becomes divided, without the intervention of a visceral cleft, into
| |
| an anterior and posterior division.
| |
| The anterior lies close to the eye,
| |
| and in front of the commencing
| |
| mouth involution. The posterior
| |
| part lies completely within the mandibular arch.
| |
| | |
| As the rudiments of the successive visceral clefts are formed, the
| |
| posterior part of the head-cavity becomes divided into successive sections, there being one section for
| |
| each arch. Thus the whole headcavity becomes on each side divided
| |
| into (i) a premandibular section ; (2)
| |
| a mandibular section (vide fig. 29 A,
| |
| PP] > (3) a hyoid section ; (4) sections
| |
| in each of the branchial arches.
| |
| | |
| The first of these divisions forms
| |
| a space of a considerable size, with
| |
| epithelial walls of somewhat short
| |
| columnar cells (fig. 381, ipp}. It is
| |
| situated close to the eye, and presents a rounded or sometimes a
| |
| triangular figure in section. The
| |
| two halves of the cavity are prolonged ventralwards, and meet below
| |
| the base of the fore-brain. The
| |
| connection between them appears to last for a considerable time.
| |
| These two cavities are the only parts of the body-cavity within
| |
| the head which unite ventrally. The section of the head-cavity
| |
| just described is so similar to the remaining sections that it
| |
| must be considered as serially homologous with them.
| |
| | |
| The next division of the head-cavity, which from its position
| |
| | |
| | |
| | |
| FIG. 381. TRANSVERSE SECTION THROUGH THE FRONT PART
| |
| OF THE HEAD OF A YOUNG PRISTIURUS EMBRYO.
| |
| | |
| The section, owing to the cranial flexure, cuts both the foreand the hind-brain. It shews the
| |
| premandibular and mandibular
| |
| head-cavities ipp and ipp, etc.
| |
| The section is moreover somewhat
| |
| oblique from side to side.
| |
| | |
| fb. fore-brain ; /. lens of eye ;
| |
| m. mouth ; pt. upper end of mouth,
| |
| forming pituitary involution; lao.
| |
| mandibular aortic arch; ipp. and
| |
| ipp. first and second head-cavities;
| |
| \vc. first visceral cleft; V. fifth
| |
| nerve ; aim. auditory nerve ; VII.
| |
| seventh nerve ; aa. dorsal aorta ;
| |
| acv. anterior cardinal vein ; ch,
| |
| notochord.
| |
| | |
| | |
| | |
| 678 THE HEAD-CAVITIES.
| |
| | |
| may be called the mandibular cavity, presents a spatulate shape,
| |
| being dilated dorsally, and produced ventrally into a long thin
| |
| process parallel to the hyomandibular gill-cleft (fig. 20, pp}.
| |
| Like the previous space it is lined by a short columnar epithelium.
| |
| | |
| The mandibular aortic arch is situated close to its inner side
| |
| (fig. 381, 2pp). After becoming separated from the lower part
| |
| (Marshall), the upper part of the cavity atrophies about the time
| |
| of the appearance of the external gills. Its lower part also
| |
| becomes much narrowed, but its walls of columnar cells persist.
| |
| The outer or somatic wall becomes very thin indeed, the
| |
| splanchnic wall, on the other hand, thickens and forms a layer
| |
| of several rows of elongated cells. In each of the remaining
| |
| arches there is a segment of the original body-cavity fundamentally similar to that in the mandibular arch (fig. 382). A dorsal
| |
| dilated portion appears, however, to be present in the third or
| |
| hyoid section alone (fig. 20), and even
| |
| there disappears very soon, after being
| |
| segmented off from the lower part
| |
| (Marshall). The cavities in the posterior parts of the head become much
| |
| reduced like those in its anterior part,
| |
| though at rather a later period. FlG . 382 . HORIZONTAL
| |
| | |
| It has been shewn that the divi- SECTION THROUGH THE PENULTIMATE VISCERAL ARCH OF
| |
| | |
| sions of the body-cavity in the head, AN EMBRYO OF PRISTIURUS.
| |
| with the exception of the anterior, e p. epiblast; vc. pouch of
| |
| early become atrophied, not so how- hypoblast which will form the
| |
| | |
| walls of a visceral cleit ; //.
| |
| CVer their walls. The cells forming segment of body-cavity in vis
| |
| the walls both of the dorsal and ven- ceral arch ; aa ' aortic arch '
| |
| tral sections of these cavities become elongated, and finally
| |
| become converted into muscles. Their exact history has not
| |
| been followed in its details, but they almost unquestionably
| |
| become the musculus contrictor superficialis and musculus interbranchialis 1 ; and probably also musculus levator mandibuli and
| |
| other muscles of the front part of the head.
| |
| | |
| The anterior cavity close to the eye remains unaltered much
| |
| longer than the remaining cavities.
| |
| | |
| 1 Vide Vetter, " Die Kiemen und Kiefermusculatur d. Fische." Jenaische Zcltschrift, Vol. vn.
| |
| | |
| | |
| | |
| | |
| THE MUSCULAR SYSTEM.
| |
| | |
| | |
| | |
| 679
| |
| | |
| | |
| | |
| Its further history is very interesting. In my original account
| |
| of this cavity (No. 292, p. 208) I stated my belief that its walls
| |
| gave rise to the eye-muscles, and the history of this process has
| |
| been to some extent worked out by Marshall in his important
| |
| memoir (No. 509).
| |
| | |
| Marshall finds that the ventral portion of this cavity, where
| |
| its two halves meet, becomes separated from the remainder.
| |
| The eventual fate of this part has not however been followed.
| |
| Each dorsal section acquires a cup-like form, investing the
| |
| posterior and inner surface of the eye. The cells of its outer
| |
| wall subsequently give rise to three sets of muscles. The middle
| |
| of these, partly also derived from the inner walls of the cup,
| |
| becomes the rectus internus of the eye, the dorsal set forms the
| |
| rectus superior, and the ventral the rectus inferior. The obliquus
| |
| inferior appears also to be in part developed from the walls of
| |
| this cavity.
| |
| | |
| Marshall brings evidence to shew that the rectus externus (as
| |
| might be anticipated from its nerve supply) has no connection
| |
| with the walls of the premandibular head-cavity, and finds that
| |
| it arises close to the position originally occupied by the second
| |
| and third cavities. Marshall has not satisfactorily made out the
| |
| mode of development of the obliquus superior.
| |
| | |
| The walls of the cavities, whose history has just been recorded, have definite relations with the cranial nerves, an account
| |
| of which has already been given at p. 461.
| |
| | |
| Head-cavities, in the main similar to those of Elasmobranchii, have been found in the embryo of Petromyzon (fig. 45,
| |
| /ic\ the Newt (Osborn and Scott), and various Reptilia (Parker).
| |
| | |
| BIBLIOGRAPHY.
| |
| | |
| (507) G.M.Humphry. " Muscles in Vertebrate Animals." Journ. of Anat.
| |
| and Phys., Vol. vi. 1872.
| |
| | |
| (508) J. Miiller. " Vergleichende Anatomic d. Myxinoiden. Part I. Osteologie
| |
| u. Myologie." Akad. Wiss., Berlin, 1834.
| |
| | |
| (509) A. M. Marshall. "On the head cavities and associated nerves of
| |
| Elasmobranchs." Quart. J. of Micr. Science, Vol. xxi. 1881.
| |
| | |
| (510) A. Schneider. " Anat. u. Entwick. d. Muskelsystems d. Wirbelthierc."
| |
| Silz. d. Oberhessischen Gesellschaft, 1873.
| |
| | |
| (511) A. Schneider. Beitrdge z. vergleich. Anat. . Entwick. d. Wirbelthiere.
| |
| Berlin, 1879.
| |
| | |
| Vide 2^0 Gotte (No. 296), Kolliker (N o. 298), Balfour (No. 292), Huxley, etc.
| |
| | |
| | |
| | |
| CHAPTER XXIII.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| EXCRETORY organs consist of coiled or branched and often
| |
| ciliated tubes, with an excretory pore opening on the outer surface
| |
| of the body, and as a rule an internal ciliated orifice placed in the
| |
| body-cavity. In forms provided with a true vascular system,
| |
| there is a special development of capillaries around the glandular
| |
| part of the excretory organs. In many instances the glandular
| |
| cells of the organs are filled with concretions of uric acid or some
| |
| similar product of nitrogenous waste.
| |
| | |
| There is a very great morphological and physiological similarity between almost all the forms of excretory organ found in
| |
| the animal kingdom, but although there is not a little to be said
| |
| for holding all these organs to be derived from some common
| |
| prototype, the attempt to establish definite homologies between
| |
| them is beset with very great difficulties.
| |
| | |
| Platyelminthes. Throughout the whole of the Platyelminthes these organs are constructed on a well-defined type, and
| |
| in the Rotifera excretory organs of a similar form to those of the
| |
| Platyelminthes are also present.
| |
| | |
| These organs (Fraipont, No. 513) are more or less distinctly
| |
| paired, and consist of a system of wide canals, often united into a
| |
| network, which open on the one hand into a pair of large tubes
| |
| leading to the exterior, and on the other into fine canals which
| |
| terminate by ciliated openings, either in spaces between the
| |
| connective-tissue cells (Platyelminthes), or in the body-cavity
| |
| (Rotifera). The fine canals open directly into the larger ones,
| |
| without first uniting into canals of an intermediate size.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 68 1
| |
| | |
| | |
| | |
| The two large tubes open to the exterior, either by means of
| |
| a median posteriorly placed contractile vesicle, or by a pair of
| |
| vesicles, which have a ventral and anterior position. The former
| |
| type is characteristic of the majority of the Trematoda, Cestoda.
| |
| and Rotifera, and the latter of the Nemertea and some Trematoda.
| |
| In the Turbellaria the position of the external openings of the
| |
| system is variable, and in a few Cestoda (Wagner) there are
| |
| lateral openings on each of the successive proglottides, in addition
| |
| to the terminal openings. The mode of development of these
| |
| organs is unfortunately not known.
| |
| | |
| Mollusca. In the Mollusca there are usually present two
| |
| independent pairs of excretory organs one found in a certain
| |
| number of forms during early larval life only 1 , and the other
| |
| always present in the adult.
| |
| | |
| The larval excretory organ has been found in the pulmonate
| |
| Gasteropoda (Gegenbaur, Fol 2 , Rabl), in Teredo (Hatschek), and
| |
| possibly also in Paludina. It is placed in the anterior region of
| |
| the body, and opens ventrally on each side, a short way behind
| |
| the velum. It is purely a larval organ, disappearing before the
| |
| close of the veliger stage. In the aquatic Pulmonata, where it is
| |
| best developed, it consists on each side of a V-shaped tube, with
| |
| a dorsally-placed apex, containing an enlargement of the lumen.
| |
| There is a ciliated cephalic limb, lined by cells with concretions,
| |
| and terminating by an internal opening near the eye, and a nonciliated pedal limb opening to the exterior 3 .
| |
| | |
| Two irreconcilable views are held as to the development of
| |
| this system. Rabl (Vol. II. No. 268) and Hatschek hold that it
| |
| is developed in the mesoblast ; and Rabl states that in Planorbis
| |
| it is formed from the anterior mesoblast cells of the mesoblastic
| |
| bands. A special mesoblast cell on each side elongates into two
| |
| processes, the commencing limbs of the future organ. A lumen
| |
| is developed in this cell, which is continued into each limb, while
| |
| | |
| 1 I leave out of consideration an external renal organ found in many marine
| |
| Gasteropod larvte, vide Vol. II. p. 280.
| |
| | |
| 2 H. Fol, "Etudes sur le devel. d. Mollusques. " Mem. Hi. Archiv d. Zool.
| |
| exfJr. et gener., Vol. VIII.
| |
| | |
| 3 The careful observations of Fol seem to me nearly conclusive in favour of this
| |
| limb having an external opening, and the statement to the reverse effect on p. 280 of
| |
| Vol. ii. of this treatise, made on the authority of Rabl and Biitschli, must probably be
| |
| corrected.
| |
| | |
| | |
| | |
| 682 POLYZOA.
| |
| | |
| the continuations of the two limbs are formed by perforated
| |
| mesoblast cells.
| |
| | |
| According to Fol these organs originate in aquatic Pulmonata
| |
| as a pair of invaginations of the epiblast, slightly behind the
| |
| mouth. Each invagination grows in a dorsal direction, and after
| |
| a time suddenly bends on itself, and grows ventralwards and
| |
| forwards. It thus acquires its V-shaped form.
| |
| | |
| In the terrestrial Pulmonata the provisional excretory organs
| |
| are, according to Fol, formed as epiblastic invaginations, in the
| |
| same way as those in the aquatic Pulmonata, but have the form
| |
| of simple non-ciliated sacks, without internal openings.
| |
| | |
| The permanent renal organ of the Mollusca consists typically
| |
| of a pair of tubes, although in the majority of the Gasteropoda
| |
| one of the two tubes is not developed. It is placed considerably
| |
| behind the provisional renal organ.
| |
| | |
| Each tube, in its most typical form, opens by a ciliated funnel
| |
| into the pericardial cavity, and has its external opening at the
| |
| side of the foot. The pericardial funnel leads into a glandular
| |
| section of the organ, the lining cells of which are filled with
| |
| concretions. This section is followed by a ciliated section, from
| |
| which a narrow duct leads to the exterior.
| |
| | |
| As to the development of this organ the same divergence of
| |
| opinion exists as in the case of the provisional renal organ.
| |
| | |
| Rabl's careful observations on Planorbis (Vol. II. No. 268) tend
| |
| to shew that it is developed from a mass of mesoblast cells, near
| |
| the end of the intestine. The mass becomes hollow, and,
| |
| attaching itself to the epiblast on the left side of the anus,
| |
| acquires an opening to the exterior. Its internal opening is not
| |
| established till after the formation of the heart. Fol gives an
| |
| equally precise account, but states that the first rudiment of the
| |
| organ arises as a solid mass of epiblast cells. Lankester finds
| |
| that this organ is developed as a paired invagination of the.
| |
| epiblast in Pisidium, and Bobretzky also derives it from the
| |
| epiblast in marine Prosobranchiata. In Cephalopoda on the
| |
| other hand Bobretzky's observations (I conclude this from his
| |
| figures) indicate that the excretory sacks of the renal organs are
| |
| derived from the mesoblast.
| |
| | |
| Polyzoa. Simple excretory organs, consisting of a pair of
| |
| ciliated canals, opening between the mouth and the anus, have
| |
| | |
| | |
| | |
| EXCRETORY ORGAN>.
| |
| | |
| | |
| | |
| 68 3
| |
| | |
| | |
| | |
| been found by Hatschek and Joliet in the Entoproctous Polyzoa,
| |
| and are developed, according to Hatschek, by whom they were
| |
| first found in the larva, from the mesoblast
| |
| | |
| Brachiopoda. One or rarely two (Rhynchonella) pairs of
| |
| canals, with both peritoneal and external openings, are found in
| |
| the Brachiopoda. They undoubtedly serve as genital ducts, but
| |
| from their structure are clearly of the same nature as the
| |
| excretory organs of the Chaetopoda described below. Their
| |
| development has not been worked out.
| |
| | |
| Chaetopoda. Two forms of excretory organ have been met
| |
| with in the Chaetopoda. The one form is universally or nearly
| |
| universally present in the adult, and typically consists of a pair
| |
| of coiled tubes repeated in every segment. Each tube has an
| |
| internal opening, placed as a rule in the segment in front of that
| |
| in which the greater part of the organ and the external opening
| |
| are situated.
| |
| | |
| There are great variations in the structure of these organs,
| |
| which cannot be dealt with here. It may be noted however that
| |
| the internal opening may be absent, and that there may be
| |
| several internal openings for each organ (Polynoe). In the
| |
| Capitellidae moreover several pairs of excretory tubes have been
| |
| shewn by Eisig (No. 512) to be present in each of the posterior
| |
| segments.
| |
| | |
| The second form of excretory organ has as yet only been
| |
| found in the larva of Polygordius, and will be more conveniently
| |
| dealt with in connection with the development of the excretory
| |
| system of this form.
| |
| | |
| There is still considerable doubt as to the mode of formation
| |
| of the excretory tubes of the Chaetopoda. Kowalevsky (No. 277),
| |
| from his observations on the Oligochasta, holds that they develop
| |
| as outgrowths of the epithelial layer covering the posterior side
| |
| of the dissepiments, and secondarily become connected with the
| |
| epidermis.
| |
| | |
| Hatschek finds that in Criodrilus they arise from a continuous
| |
| linear thickening of the somatic mesoblast, immediately beneath
| |
| the epidermis, and dorsal to the ventral band of longitudinal
| |
| muscles. They break up into S-shaped cords, the anterior end
| |
| of each of which is situated in front of a dissepiment, and is
| |
| formed at first of a single large cell, while the posterior part is
| |
| | |
| | |
| | |
| 684 CHvETOPODA.
| |
| | |
| | |
| | |
| continued into the segment behind. The cords are covered by
| |
| a peritoneal lining, which still envelopes them, when in the
| |
| succeeding stage they are carried into the body-cavity. They
| |
| subsequently become hollow, and their hinder ends acquire
| |
| openings to the exterior. The formation of their internal
| |
| openings has not been followed.
| |
| | |
| Kleinenberg is inclined to believe that the excretory tubes
| |
| take their origin from the epiblast, but states that he has not
| |
| satisfactorily worked out their development.
| |
| | |
| The observations of Risig (No. 512) on the Capitellidae
| |
| support Kowalevsky's view that the excretory tubes originate
| |
| from the lining of the peritoneal cavity.
| |
| | |
| Hatschek (No. 514) has given a very interesting account of
| |
| the development of the excretory system in Polygordius.
| |
| | |
| The excretory system begins to be formed, while the larva is
| |
| still in the trochospere stage (fig. 383, npli), and consists of a
| |
| provisional excretory organ, which is placed in front of the future
| |
| segmented part of the body, and occupies a position very
| |
| similar to that of the provisional excretory organ found in some Molluscan
| |
| larvae (vide p. 68 1).
| |
| | |
| Hatschek, with some shew of reason, holds that the provisional excretory organs of Polygordius are homologous with those of the Mollusca.
| |
| | |
| In its earliest stage the provisional
| |
| excretory organ of Polygordius consists of a pair of simple ciliated tubes, FIG. 383. POLYOORDIUS
| |
| | |
| , . , r 11-1 LARVA. (After Hatschek.)
| |
| | |
| each with an anterior funnel-like open- m _ moulh . ^ supraKBSO .
| |
| | |
| ing situated in the midst of the meSO- phageal ganglion ; nph. nephri11 11 . , dion ; ine.p. mesoblastic band;
| |
| | |
| blast cells, and a posterior external an _ anus 5 oL stomach .
| |
| opening. The latter is placed immediately in front of what afterwards becomes the segmented region
| |
| of the embryo. While the larva is still unsegmented, a second
| |
| internal opening is formed for each tube (fig. 383, np/i) and the
| |
| two openings so formed may eventually become divided into
| |
| five (fig. 384 A), all communicating by a single pore with the
| |
| exterior.
| |
| | |
| When the posterior region of the embryo becomes segmented,
| |
| | |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 685
| |
| | |
| | |
| | |
| paired excretory organs are formed in each of the posterior
| |
| segments, but the account of their development, as given by
| |
| Hatschek, is so remarkable that I do not think it can be
| |
| definitely accepted without further confirmation.
| |
| | |
| From the point of junction of the two main branches of the
| |
| larval kidney there grows backwards (fig. 384 B), to the hind
| |
| end of the first segment, a very delicate tube, only indicated by
| |
| its ciliated lumen, its walls not being differentiated. Near the
| |
| front end of this tube a funnel, leading into the larval body
| |
| cavity of the head, is formed, and subsequently the posterior end
| |
| of the tube acquires an external opening, and the tube distinct
| |
| walls. The communication with the provisional excretory organ
| |
| is then lost, and thus the excretory tube of the first segment is
| |
| established.
| |
| | |
| The excretory tubes in the second and succeeding segments
| |
| are formed in the same way as in the first, i.e. by the continuation of the lumen of the hind end of the excretory tube from
| |
| the preceding segment, and the subsequent separation of this
| |
| part as a separate tube.
| |
| | |
| The tube may be continued with a sinuous course through
| |
| | |
| | |
| | |
| | |
| | |
| A
| |
| A
| |
| | |
| A
| |
| +
| |
| | |
| A.
| |
| | |
| | |
| | |
| Y
| |
| | |
| Y
| |
| Y
| |
| Y
| |
| Y
| |
| | |
| | |
| | |
| J)
| |
| | |
| | |
| | |
| FIG. 384. DIAGRAM ILLUSTRATING THE DEVELOPMENT OF THE EXCRETORY
| |
| SYSTEM OF POLYGORDIUS. (After Hatschek.)
| |
| | |
| several segments without a distinct wall. The external and
| |
| internal openings of the permanent excretory tubes are thus
| |
| secondarily acquired. The internal openings communicate with
| |
| the permanent body-cavity. The development of the perma
| |
| | |
| | |
| 686 GEPHYREA.
| |
| | |
| | |
| | |
| nent excretory tubes is diagrammatically represented in fig.
| |
| 384 C and D.
| |
| | |
| The provisional excretory organ atrophies during larval life.
| |
| | |
| If Hatschek's account of the development of the excretory system of
| |
| Polygordius is correct, it is clear that important secondary modifications
| |
| must have taken place in it, because his description implies that there sprouts
| |
| from the anterior excretory organ, while it has its own external opening, a
| |
| posterior duct, which does not communicate either with the exterior or with
| |
| the body-cavity! Such a duct could have no function. It is intelligible
| |
| either (i) that the anterior excretory organ should lead into a longitudinal
| |
| duct, opening posteriorly ; that then a series of secondary openings into the
| |
| body-cavity should attach themselves to this, that for each internal opening
| |
| an external should subsequently arise, and the whole break up into separate
| |
| tubes ; or (2) that behind an anterior provisional excretory organ a series of
| |
| secondary independent segmental tubes should be formed. But from Hatschek's account neither of these modes of evolution can be deduced.
| |
| | |
| Gephyrea. The Gephyrea may have three forms of excretory organs, two of which are found in the adult, and one,
| |
| similar in position and sometimes also in structure, to the
| |
| provisional excretory organ of Polygordius, has so far only been
| |
| found in the larvae of Echiurus and Bonellia.
| |
| | |
| In all the Gephyrea the so-called 'brown tubes' are
| |
| apparently homologous with the segmented excretory tubes of
| |
| Chaetopods. Their main function appears to be the transportation of the generative products to the exterior. There is but a
| |
| single highly modified tube in Bonellia, forming the oviduct and
| |
| uterus ; a pair of tubes in the Gephyrea inermia, and two or
| |
| three pairs in most Gephyrea armata, except Bonellia. Their
| |
| development has not been studied.
| |
| | |
| In the Gephyrea armata there is always present a pair of
| |
| posteriorly placed excretory organs, opening in the adult into
| |
| the anal extremity of the alimentary tract, and provided with
| |
| numerous ciliated peritoneal funnels. These organs were stated
| |
| by Spengel to arise in Bonellia as outgrowths of the gut ; but in
| |
| Echinrus Hatschek (No. 515) finds that they are developed from
| |
| the somatic mesoblast of the terminal part of the trunk. They
| |
| soon become hollow, and after attaching themselves to the
| |
| epiblast on each side of the anus, acquire external openings.
| |
| They are not at first provided with peritoneal funnels, but these
| |
| parts of the organs become developed from a ring of cells at
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 687
| |
| | |
| | |
| | |
| their inner extremities ; and there is at first but a single funnel
| |
| for each vesicle. The mode of increase of the funnels has not
| |
| been observed, nor has it been made out how the organs themselves become attached to the hind-gut.
| |
| | |
| The provisional excretory organ of Echiurus is developed at
| |
| an early larval stage, and is functional during the whole of
| |
| larval life. It at first forms a ciliated tube on each side, placed
| |
| in front of that part of the larva which becomes the trunk of the
| |
| adult. It opens to the exterior by a fine pore on the ventral
| |
| side, immediately in front of one of the mesoblastic bands, and
| |
| appears to be formed of perforated cells. It terminates internally in a slight swelling, which represents the normal internal
| |
| ciliated funnel. The primitively simple excretory organ becomes
| |
| eventually highly complex by the formation of numerous
| |
| branches, each ending in a slightly swollen extremity. These
| |
| branches, in the later larval stages, actually form a network, and
| |
| the inner end of each main branch divides into a bunch of fine
| |
| tubes. The whole organ resembles in many respects the excretory organ of the Platyelminthes.
| |
| | |
| In the larva of Bonellia Spengel has described a pair of
| |
| provisional excretory tubes, opening near the anterior end of
| |
| the body, which are probably homologous with the provisional
| |
| excretory organs of Echiurus (vide Vol. II., fig. 162 C, se).
| |
| | |
| Discophora. As in many of the types already spoken of,
| |
| permanent and provisional excretory organs may be present in
| |
| the Discophora. The former are usually segmentally arranged,
| |
| and resemble in many respects the excretory tubes of the
| |
| Chaetopoda. They may either be provided with a peritoneal
| |
| funnel (Nephelis, Clepsine) or have no internal opening
| |
| (Hirudo).
| |
| | |
| Bourne 1 has shewn that the cells surrounding the main duct
| |
| in the medicinal Leech are perforated by a very remarkable
| |
| network of ductules, and the structure of these organs in the
| |
| Leech is so peculiar that it is permissible to state with due reserve
| |
| their homology with the excretory organs of the Chaetopoda.
| |
| | |
| The excretory tubes of Clepsine are held by Whitman to be
| |
| developed in the mesoblast.
| |
| | |
| 1 "On the Structure of the Nephridia of the Medicinal Leech." Quart. J. of
| |
| Micr. Science, Vol. XX. 1880.
| |
| | |
| | |
| | |
| 688 ARTHROPODA.
| |
| | |
| | |
| | |
| There are found in the embryos of Nephelis and Hirudo
| |
| certain remarkable provisional excretory organs the origin and
| |
| history of which are not yet fully made out. In Nephelis they
| |
| appear as one (according to Robin), or (according to Biitschli)
| |
| as two successive pairs of convoluted tubes on the dorsal side of
| |
| the embryo, which are stated by the latter author to develop
| |
| from the scattered mesoblast cells underneath the skin. At
| |
| their fullest development they extend, according to Robin, from
| |
| close to the head to near the ventral sucker. Each of them is
| |
| U-shaped, with the open end of the U forwards, each limb of the
| |
| U being formed by two tubes united in front. No external
| |
| opening has been clearly made out. Fiirbringer is inclined from
| |
| his own researches to believe that they open laterally. They
| |
| contain a clear fluid.
| |
| | |
| In Hirudo, Leuckart has described three similar pairs of
| |
| organs, the structure of which he has fully elucidated. They
| |
| are situated in the posterior part of the body, and each of them
| |
| commences with an enlargement, from which a convoluted tube
| |
| is continued for some distance backwards; the tube then turns
| |
| forwards again, and after bending again upon itself opens to the
| |
| exterior. The anterior part is broken up into a kind of
| |
| labyrinthic network.
| |
| | |
| The provisional excretory organs of the Leeches cannot be
| |
| identified with the anterior provisional organs of Polygordius
| |
| and Echiurus.
| |
| | |
| Arthropoda. Amongst the Arthropoda Peripatus is the
| |
| only form with excretory organs of the type of the segmental
| |
| excretory organs of the Chsetopoda 1 .
| |
| | |
| These organs are placed at the bases of the feet, in the
| |
| lateral divisions of the body-cavity, shut off from the main
| |
| median division of the body-cavity by longitudinal septa of
| |
| transverse muscles.
| |
| | |
| Each fully developed organ consists of three parts :
| |
| | |
| (i) A dilated vesicle opening externally at the base of a
| |
| foot. (2) A coiled glandular tube connected with this, and
| |
| subdivided again into several minor divisions. (3) A short
| |
| terminal portion opening at one extremity into the coiled tube
| |
| | |
| 1 Vide F. M. Balfour, " On some points in the Anatomy of Peripatus Capensis."
| |
| Quart. J, of Micr. Science, Vol. XIX. 1879.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 689
| |
| | |
| | |
| | |
| and at the other, as I believe, into the body cavity. This
| |
| section becomes very conspicuous, in stained preparations, by
| |
| the intensity with which the nuclei of its walls absorb the
| |
| colouring matter.
| |
| | |
| In the majority of the Tracheata the excretory organs have
| |
| the form of the so-called Malpighian tubes, which always (vide
| |
| Vol. II.) originate as a pair of outgrowths of the epiblastic
| |
| proctodaeum. From their mode of development they admit of
| |
| comparison with the anal vesicles of the Gephyrea, though in
| |
| the present state of our knowledge this comparison must be
| |
| regarded as somewhat hypothetical.
| |
| | |
| The antennary and shell-glands of the Crustacea, and
| |
| possibly also the so-called dorsal organ of various Crustacean
| |
| larvae appear to be excretory, and the two former have been
| |
| regarded by Claus and Grobben as belonging to the same
| |
| system as the segmental excretory tubes of the Chaetopoda.
| |
| | |
| Nematoda. Paired excretory tubes, running for the whole
| |
| length of the body in the so-called lateral line, and opening in
| |
| front by a common ventral pore, are present in the Nematoda.
| |
| They do not appear to communicate with the body cavity, and
| |
| their development has not been studied.
| |
| | |
| Very little is known with reference either to the structure or
| |
| development of excretory organs in the Echinodermata and the
| |
| other Invertebrate types of which no mention has been so far
| |
| made in this Chapter.
| |
| | |
| Excretory organs and generative ducts of the Craniata.
| |
| | |
| Although it would be convenient to separate, if possible, the
| |
| history of the excretory organs from that of the generative
| |
| ducts, yet these parts are so closely related in the Vertebrata, in
| |
| some cases the same duct having at once a generative and a
| |
| urinary function, that it is not possible to do so.
| |
| | |
| The excretory organs of the Vertebrata consist of three
| |
| distinct glandular bodies and of their ducts. These are (i) a
| |
| small glandular body, usually with one or more ciliated funnels
| |
| opening into the body cavity, near the opening of which there
| |
| projects into the body cavity a vascular glomerulus. It is
| |
| situated very far forwards, and is usually known as the head
| |
| 44
| |
| | |
| | |
| | |
| 690 ELASMOBRANCHII.
| |
| | |
| | |
| | |
| kidney, though it may perhaps be more suitably called, adopting
| |
| Lankester's nomenclature, the pronepliros. Its duct, which forms
| |
| the basis for the generative and urinary ducts, will be called the
| |
| segmented duct.
| |
| | |
| (2) The Wolffian body, which may be also called the
| |
| mesonepJiros. It consists of a series of, at first, segmentally
| |
| (with a few exceptions) arranged glandular canals (segmental
| |
| tubes) primitively opening at one extremity by funnel-shaped
| |
| apertures into the body cavity, and at the other into the
| |
| segmental duct. This duct becomes in many forms divided
| |
| longitudinally into two parts, one of which then remains
| |
| attached to the segmental tubes and forms the Wolffian or
| |
| mesonepJiric duct, while the other is known as the Milllerian
| |
| dnct.
| |
| | |
| (3) The kidney proper or metanephros. This organ is only
| |
| found in a completely differentiated form in the amniotic Vertebrata. Its duct is an outgrowth from the Wolrfian duct.
| |
| | |
| The above parts do not coexist in full activity in any living
| |
| adult member of the Vertebrata, though all of them are found
| |
| together in certain embryos. They are so intimately connected
| |
| that they cannot be satisfactorily dealt with separately.
| |
| | |
| Elasmobranchii. The excretory system of the Elasmobranchii is by no means the most primitive known, but at the
| |
| same time it forms a convenient starting point for studying the
| |
| modifications of the system in other groups. The most remarkable peculiarity it presents is the absence of a pronephros.
| |
| The development of the Elasmobranch excretory system has
| |
| been mainly studied by Semper and myself.
| |
| | |
| The first trace of the system makes its appearance as a knob
| |
| of mesoblast, springing from the intermediate cell-mass near the
| |
| level of the hind end of the heart (fig. 385 K,pd). This knob is
| |
| the rudiment of the abdominal opening of the segmental duct,
| |
| and from it there grows backwards to the level of the anus a
| |
| solid column of cells, which constitutes the rudiment of the
| |
| segmental duct itself (fig. 385 B, pd). The knob projects
| |
| towards the epiblast, and the column connected with it lies
| |
| between the mesoblast and epiblast. The knob and column do
| |
| not long remain solid, but the former acquires an opening into
| |
| the body cavity (fig. 421, sd) continuous with a lumen, which
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 691
| |
| | |
| | |
| | |
| makes its appearance in the column (fig. 386, sd). The knob
| |
| forms the only structure which can be regarded as a rudiment of
| |
| the pronephros.
| |
| | |
| | |
| | |
| spn
| |
| | |
| | |
| | |
| spn
| |
| | |
| | |
| | |
| | |
| FlG. 385. TWO SECTIONS OF A PRISTIURUS EMBRYO WITH THREE VISCERAL
| |
| | |
| CLEFTS.
| |
| | |
| The sections illustrate the development of the segmental duct (pd) or primitive
| |
| duct of the pronephros. In A (the anterior of the two sections) this appears as a
| |
| solid knob (pd) projecting towards the epiblast. In B is seen a section of the column
| |
| which has grown backwards from the knob in A.
| |
| | |
| spn. rudiment of a spinal nerve; me. medullary canal; ch. notochord; X. subnotochordal rod; mp. muscle-plate; mp' . specially developed portion of muscle-plate;
| |
| ao. dorsal aorta ; pd. segmental duct ; so. somatopleure ; sp. splanchnopleure ; //.
| |
| body cavity; ep. epiblast; al. alimentary canal.
| |
| | |
| While the lumen is gradually being formed, the segmental
| |
| tubes of the mesonephros become established. They appear to
| |
| arise as differentiations of the parts of the primitive lateral plates
| |
| of mesoblast, placed between the dorsal end of the body cavity
| |
| and the muscle-plate (fig. 386, st) 1 , which are usually known as
| |
| the intermediate cell-masses.
| |
| | |
| The lumen of the segmental tubes, though at first very small,
| |
| soon becomes of a considerable size. It appears to be established
| |
| in the position of the section of the body cavity in the intermediate cell-mass, which at first unites the part of the body
| |
| cavity in the muscle-plates with the permanent body cavity.
| |
| The lumen of each tube opens at its lower end into the dorsal
| |
| part of the body cavity (fig. 386, st}, and each tube curls obliquely
| |
| | |
| 1 In my original account of the development I held these tubes to be invaginations
| |
| of the peritoneal epithelium. Sedgwick (No. 549) was led to doubt the accuracy of
| |
| my original statement from his investigations on the chick ; and from a re-examination of my specimens he arrived at the results stated above, and which I am now
| |
| myself inclined to adopt.
| |
| | |
| 442
| |
| | |
| | |
| | |
| 692
| |
| | |
| | |
| | |
| ELASMOBRANCHII.
| |
| | |
| | |
| | |
| sp.c
| |
| | |
| | |
| | |
| | |
| backwards round the inner and dorsal side of the segmental
| |
| duct, near which it at first ends blindly.
| |
| | |
| One segmental tube makes its
| |
| appearance for each somite (fig. 265),
| |
| commencing with that immediately
| |
| behind the abdominal opening of the
| |
| segmental duct, the last tube being
| |
| situated a few segments behind the
| |
| anus. Soon after their formation
| |
| the blind ends of the segmental tubes
| |
| come in contact with, and open into
| |
| the segmental duct, and each of them
| |
| becomes divided into four parts.
| |
| These are (i) a section carrying the
| |
| peritoneal opening, known as the
| |
| peritoneal funnel, (2) a dilated vesicle
| |
| into which this opens, (3) a coiled
| |
| tubulus proceeding from (2), and
| |
| terminating in (4) a wider portion
| |
| opening into the segmental duct. At
| |
| the same time, or shortly before this,
| |
| each segmental duct unites with and
| |
| opens into one of the horns of the
| |
| cloaca, and also retires from its
| |
| primitive position between the epiblast and mesoblast, and assumes a
| |
| position close to the epithelium lining
| |
| the body cavity (fig. 380, sd}. The
| |
| general features of the excretory
| |
| organs at this period are diagrammatically represented in the
| |
| woodcut (fig. 387). In this fig. pd is the segmental duct and
| |
| o its abdominal opening; s.t points to the segmental tubes,
| |
| the finer details of whose structure are not represented in the
| |
| diagram. The mesonephros thus forms at this period an elongated gland composed of a series of isolated coiled tubes, one
| |
| extremity of each of which opens into the body cavity, and the
| |
| other into the segmental duct, which forms the only duct of the
| |
| system, and communicates at its front end with the body cavity,
| |
| and behind with the cloaca.
| |
| | |
| | |
| | |
| FIG. 386. SECTION THROUGH
| |
| THE TRUNK OF A SCYLLIUM EMBRYO SLIGHTLY YOUNGER THAN
| |
| | |
| 28 F.
| |
| | |
| sp.c. spinal canal; W. white
| |
| matter of spinal cord ; pr. posterior nerve-roots ; ch. notochord ;
| |
| x. sub-notochordal rod ; ao. aorta ;
| |
| nip, muscle-plate ; nip', inner layer
| |
| of muscle-plate already converted
| |
| into muscles ; Vr, rudiment of
| |
| vertebral body ; st. segmental
| |
| tube; sd. segmental duct; sp.v.
| |
| spiral valve ; v. subintestinal vein ;
| |
| p.o. primitive generative cells.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 693
| |
| | |
| | |
| | |
| The next important change concerns the segmental duct,
| |
| which becomes longitudinally split into two complete ducts in
| |
| the female, and one complete duct and parts of a second duct in
| |
| the male. The manner in which this takes place is diagrammatically represented in fig. 387 by the clear line x, and in
| |
| transverse section in figs. 388 and 389. The resulting ducts are
| |
| (i) the Wolffian duct or mesonephric duct (wd\ dorsally, which
| |
| remains continuous with the excretory tubules of the mesonephros, and ventrally (2) the oviduct or Miillerian duct in the
| |
| female, and the rudiments of this duct in the male. In the
| |
| | |
| | |
| | |
| | |
| | |
| FIG. 387. DIAGRAM OF THE PRIMITIVE CONDITION OF THE KIDNEY IN AN
| |
| | |
| ELASMOBRANCH EMBRYO.
| |
| | |
| pd. segmental duct. It opens at o into the body cavity and at its other extremity
| |
| into the cloaca; x. line along which the division appears which separates the segmental
| |
| duct into the Wolffian duct above and the Miillerian duct below; s.t. segmental
| |
| tubes. They open at one end into the body cavity, and at the other into the segmental duct.
| |
| | |
| female the formation of these ducts takes place (fig. 389) by a
| |
| nearly solid rod of cells being gradually split off from the
| |
| ventral side of all but the foremost part of the original segmental
| |
| duct. This nearly solid cord is the Miillerian duct (pd}. A
| |
| very small portion of the lumen of the original segmental duct
| |
| is perhaps continued into it, but in any case it very soon acquires
| |
| a wide lumen (fig. 389 A). The anterior part of the segmental
| |
| duct is not divided, but remains continuous with the Mullerian
| |
| duct, of which its anterior pore forms the permanent peritoneal
| |
| opening 1 (fig. 387). The remainder of the segmental duct (after
| |
| the loss of its anterior section, and the part split off from its
| |
| ventral side) forms the Wolffian duct. The process of formation
| |
| of these ducts in the male differs from that in the female chiefly
| |
| | |
| 1 Five or six segmental tubes belong to the region of the undivided anterior part
| |
| of the segmental duct, which forms the front end of the Mullerian duct ; but they appear to atrophy very early, without acquiring a definite attachment to the segmental
| |
| duct.
| |
| | |
| | |
| | |
| 694
| |
| | |
| | |
| | |
| ELASMOBRANCHIL
| |
| | |
| | |
| | |
| in the fact of the anterior undivided part of the segmental duct,
| |
| which forms the front end of the Miillerian duct, being shorter,
| |
| | |
| | |
| | |
| | |
| trd/
| |
| | |
| | |
| | |
| | |
| FIG. 389. FOUR SECTIONS
| |
| THROUGH THE ANTERIOR
| |
| I'ART OF THE SEGMENTAL
| |
| DUCT OF A FEMALE EMBRYO
| |
| OF SCYLLIUM CANICULA.
| |
| | |
| The figure shews how the
| |
| segmental duct becomes split
| |
| into the Wolffian or mesonephric duct above, and Miillerian duct or oviduct below.
| |
| | |
| wd. Wolffian or mesonephric duct; od. Miillerian
| |
| duct or oviduct ; sd. segmental duct.
| |
| | |
| | |
| | |
| FIG. 388. DIAGRAMMATIC REPRESENTATION OF A TRANSVERSE SECTION OF A
| |
| | |
| SCYLLIUM EMBRYO ILLUSTRATING THE
| |
| FORMATION OF THE WOLFFIAN AND MlJLLERIAN DUCTS BY THE LONGITUDINAL
| |
| SPLITTING OF THE SEGMENTAL DUCT.
| |
| | |
| me. medullary canal; mp. muscle-plate;
| |
| ch. notochord; ao. aorta; cav. cardinal
| |
| vein; st. segmental tube. On the left side
| |
| the section passes through the opening of
| |
| a segmental tube into the body cavity. On
| |
| the right this opening is represented by
| |
| dotted lines, and the opening of the segmental tube into the Wolffian duct has
| |
| been cut through; iv.d. Wolffian duct;
| |
| m.d. Miillerian duct. The section is taken
| |
| through the point where the segmental
| |
| duct and Wolffian duct have just become
| |
| separate; gr. the germinal ridge with the
| |
| thickened germinal epithelium ; /. liver ;
| |
| i. intestine with spiral valve.
| |
| | |
| and in the column of cells with which it is continuous being
| |
| from the first incomplete.
| |
| | |
| The segmental tubes of the mesonephros undergo further
| |
| important changes. The vesicle at the termination of each peritoneal funnel sends a bud forwards towards the preceding
| |
| tubulus, which joins the fourth section of it close to the opening
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 695
| |
| | |
| | |
| | |
| | |
| into the Wolffian duct (fig. 390, px). The remainder of the
| |
| vesicle becomes converted
| |
| into a Malpighian body (mg}.
| |
| | |
| By the first of these changes 10^-4 M @W>f
| |
| a tube is established connecting each pair of segments
| |
| of the mesonephros, and
| |
| though this tube is in part
| |
| aborted (or only represented
| |
| by a fibrous band) in the
| |
| anterior part of the excretory
| |
| organs in the adult, and most
| |
| probably in the hinder part,
| |
| yet it seems almost certain
| |
| that the secondary and tertiary Malpighian bodies of
| |
| the majority of segments are
| |
| developed from its persisting
| |
| blind end. Each of these
| |
| | |
| | |
| | |
| FIG. 390. LONGITUDINAL VERTICAL
| |
| SECTION THROUGH PART OF THE MESONEPHROS OF AN EMBRYO OF SCYLLIUM.
| |
| | |
| The figure contains two examples of the
| |
| budding of the vesicle of a segmental tube
| |
| (which forms a Malpighian body in its own
| |
| segment) to unite with the tubulus in the
| |
| preceding segment close to its opening into
| |
| the Wolffian (mesonephric) duct.
| |
| | |
| ge. epithelium of body-cavity; st. peritoneal funnel of segmental tube with its
| |
| peritoneal opening; mg. Malpighian body;
| |
| px. bud from Malphigian body uniting with
| |
| preceding segment.
| |
| | |
| | |
| | |
| secondary and tertiary Malpighian bodies is connected with a
| |
| convoluted tubulus (fig. 391, a.mg), which is also developed from
| |
| the tube connecting each pair of segmental tubes, and therefore
| |
| falls into the primary tubulus close to its junction with the
| |
| | |
| | |
| | |
| st.c
| |
| | |
| | |
| | |
| | |
| w.d
| |
| | |
| | |
| | |
| FIG. 391. THREE SEGMENTS OF THE ANTERIOR PART OF THE MESONEPHROS OF A
| |
| NEARLY RIPE EMBRYO OF SCYLLIUM CANICULA AS A TRANSPARENT OBJECT.
| |
| The figure shews a fibrous band passing from the primary to the secondary Malpighian bodies in two segments, which is the remains of the outgrowth from the
| |
| primary Malpighian body.
| |
| | |
| sf.o. peritoneal funnel; p. ing. primary Malpighian body; a.mg. accessory Malpighian body; w.d. mesonephric (Wolffian) duct.
| |
| | |
| | |
| | |
| 696 ELASMOBRANCI1II.
| |
| | |
| | |
| | |
| segmental duct. Owing to the formation of the accessory tubuli
| |
| the segments of the mesonephros acquire a compound character.
| |
| | |
| The third section of each tubulus becomes by continuous
| |
| growth, especially in the hinder segments, very bulky and
| |
| convoluted.
| |
| | |
| The general character of a slightly developed segment of
| |
| the mesonephros at its full growth may be gathered from fig.
| |
| 391. It commences with (i) a peritoneal opening, somewhat
| |
| oval in form (st.d) and leading directly into (2) a narrow tube,
| |
| the segmental tube, which takes a more or less oblique course
| |
| backwards, and, passing superficially to the Wolffian duct (w.d},
| |
| opens into (3) a Malpighian body (p.mg) at the anterior extremity of an isolated coil of glandular tubuli. This coil forms
| |
| the third section of each segment, and starts from the Malpighian body. It consists of a considerable number of rather
| |
| definite convolutions, and after uniting with tubuli from one,
| |
| two, or more (according to the size of the segment) accessory
| |
| Malpighian bodies (a.mg) smaller than the one into which the
| |
| segmental tube falls, eventually opens by (4) a narrowish
| |
| collecting tube into the Wolffian duct at the posterior end of
| |
| the segment. Each segment is probably completely isolated
| |
| from the adjoining segments, and never has more than one
| |
| peritoneal funnel and one communication with the Wolffian duct.
| |
| | |
| Up to this time there has been no distinction between the
| |
| anterior and posterior tubuli of the mesonephros, which alike
| |
| open into the Wolffian duct. The collecting tubes of a considerable number of the hindermost tubuli (ten or eleven in
| |
| Scyllium canicula), either in some species elongate, overlap,
| |
| while at the same time their openings travel backward so that
| |
| they eventually open by apertures (not usually so numerous as
| |
| the separate tubes), on nearly the same level, into the hindermost section of the Wolffian duct in the female, or into the
| |
| urinogenital cloaca, formed by the coalesced terminal parts of
| |
| the Wolffian ducts, in the male; or in other species become
| |
| modified, by a peculiar process of splitting from the Wolnian
| |
| duct, so as to pour their secretion into a single duct on each
| |
| side, which opens in a position corresponding with the numerous
| |
| ducts of the other species (fig. 392). In both cases the modified
| |
| posterior kidney-segments are probably equivalent to the per
| |
| | |
| | |
| EXCRETORY ORGANS. 697
| |
| | |
| | |
| | |
| manent kidney or metanephros of the amniotic Vertebrates, and
| |
| for this reason the numerous collecting tubes or single collecting
| |
| tube, as the case may be, will be spoken of as ureters. The
| |
| anterior tubuli of the primitive excretory organ retain their early
| |
| relation to the Wolffian duct, and form the permanent Wolffian
| |
| body or mesonephros.
| |
| | |
| The originally separate terminal extremities of the Wolffian
| |
| ducts always coalesce, and form a urinal cloaca, opening by a
| |
| single aperture, situated at the extremity of the median papilla
| |
| behind the anus. Some of the peritoneal openings of the segmental tubes in Scyllium, or in other cases all the openings,
| |
| become obliterated.
| |
| | |
| In the male the anterior segmental tubes undergo remarkable modifications, and become connected with the testes.
| |
| Branches appear to grow from the first three or four or more of
| |
| them (though probably not from their peritoneal openings),
| |
| which pass to the base of the testis, and there uniting into a
| |
| longitudinal canal, form a network, and receive the secretion of
| |
| the testicular ampullae (fig. 393, nf). These ducts, the vasa
| |
| efferent ia, carry the semen to the Wolffian body, but before
| |
| opening into the tubuli of this body they unite into a canal
| |
| known as the longitudinal canal of the Wolffian body (l.c\ from
| |
| which pass off ducts equal in number to the vasa efferentia,
| |
| each of which normally ends in a Malpighian corpuscle. From
| |
| the Malpighian corpuscles so connected there spring the convoluted tubuli, forming the generative segments of the Wolffian
| |
| body, along which the semen is conveyed to the Wolffian duct
| |
| (v.d). The Wolffian duct itself becomes much contorted and
| |
| acts as vas deferens.
| |
| | |
| Figs. 392 and 393 are diagrammatic representations of the
| |
| chief constituents of the adult urinogenital organs in the two
| |
| sexes. In the adult female (fig. 392), there are present the
| |
| following parts :
| |
| | |
| (1) The oviduct or Mullerian duct (m.d) split off from the
| |
| segmental duct of the kidneys. Each oviduct opens at its
| |
| anterior extremity into the body cavity, and behind the two
| |
| oviducts have independent communications with the general
| |
| cloaca.
| |
| | |
| (2) The mesonephric ducts (w.d), the other product of the
| |
| | |
| | |
| | |
| 698
| |
| | |
| | |
| | |
| ELASMOBRANCHII.
| |
| | |
| | |
| | |
| segmental ducts of the kidneys. They end in front by becoming continuous with the tubulus of the anterior persisting
| |
| segment of the mesonephros on each side, and unite behind to
| |
| | |
| | |
| | |
| | |
| FIG. 392. DIAGRAM OF THE ARRANGEMENT OF THE URINOGENITAL ORGANS
| |
| | |
| IN AN ADULT FEMALE ELASMOBRANCH.
| |
| | |
| m.d. Miillerian duct; w.d. Wolffian duct; s.t. segmental tubes; five of them are
| |
| represented with openings into the body cavity, the posterior segmental tubes form
| |
| the mesonephros ; ov. ovary.
| |
| | |
| open by a common papilla into the cloaca. The mesonephric
| |
| duct receives the secretion of the anterior tubuli of the primitive
| |
| mesonephros.
| |
| | |
| (3) The ureter which carries off the secretion of the kidney
| |
| proper or metanephros. It is represented in my diagram in its
| |
| most rare and differentiated condition as a single duct connected
| |
| with the posterior segmental tubes.
| |
| | |
| (4) The segmental tubes (.$-./) some of which retain their
| |
| | |
| | |
| | |
| -S.t:
| |
| | |
| | |
| | |
| | |
| FIG. 393. DIAGRAM OF THE ARRANGEMENT OF THE URINOGENITAL ORGANS
| |
| | |
| IN AN ADULT MALE ELASMOBRANCH.
| |
| | |
| m.d. rudiment of Miillerian duct; w.d. Wolffian duct, marked vd in front and
| |
| serving as vas deferens; s.t. segmental tubes; two of them are represented with openings into the body cavity; d. ureter; /. testis; nt. canal at the base of the testis;
| |
| VE, vasa efferentia; Ic. longitudinal canal of the Wolffian body.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 699
| |
| | |
| | |
| | |
| original openings into the body cavity, and others are without
| |
| them. They are divided into two groups, an anterior forming
| |
| the mesonephros or Wolffian body, which pours its secretion
| |
| into the Wolffian duct ; and a posterior group forming a gland
| |
| which is probably equivalent to the kidney proper of amniotic
| |
| Craniata, and is connected with the ureter.
| |
| | |
| In the male the following parts are present (fig. 393):
| |
| | |
| (1) The Mlillerian duct (m.d], consisting of a small rudiment attached to the liver, representing the foremost end of the
| |
| oviduct of the female.
| |
| | |
| (2) The mesonephric duct (w.d] which precisely corresponds
| |
| to the mesonephric duct of the female, but, in addition to
| |
| serving as the duct of the Wolffian body, also acts as a vas
| |
| deferens (vd}. In the adult male its foremost part has a very
| |
| tortuous course.
| |
| | |
| (3) The ureter (d\ which has the same fundamental constitution as in the female.
| |
| | |
| (4) The segmental tubes (s.t). The posterior tubes have
| |
| the same arrangement in both sexes, but in the male modifications take place in connection with the anterior tubes to fit them
| |
| to act as transporters of the semen.
| |
| | |
| Connected with the anterior tubes there are present (i) the
| |
| vasa efferentia (VE], united on the one hand with (2) the
| |
| central canal in the base of the testis (/), and on the other with
| |
| the longitudinal canal of the Wolffian body (/<?). From the
| |
| latter are seen passing off the successive tubuli of the anterior
| |
| segments of the Wolffian body, in connection with which Malpighian bodies are typically present, though not represented in
| |
| my diagram.
| |
| | |
| Apart from the absence of the pronephros the points which
| |
| deserve notice in the Elasmobranch excretory system are (i)
| |
| The splitting of the segmental duct into Wolffian (mesonephric)
| |
| and Mullerian ducts. (2) The connection of the former with
| |
| the mesonephros, and of the latter with the abdominal opening
| |
| of the segmental duct which represents the pronephros of other
| |
| types. (3) The fact that the Mullerian duct serves as oviduct,
| |
| and the Wolffian duct as vas deferens. (4) The differentiation
| |
| of a posterior section of the mesonephros into a special gland
| |
| foreshadowing the metanephros of the Amniota.
| |
| | |
| | |
| | |
| /OO CYCLOSTOMATA.
| |
| | |
| | |
| | |
| Cyclostomata. The development of the excretory system
| |
| amongst the Cyclostomata has only been studied in Petromyzon
| |
| (Miiller, Furbringer, and Scott).
| |
| | |
| The first part of the system developed is the segmental duct.
| |
| It appears in the embryo of about 14 days (Scott) as a solid
| |
| cord of cells, differentiated from the somatic mesoblast near the
| |
| dorsal end of the body cavity. This cord is at first placed
| |
| immediately below the epiblast, and grows backwards by a
| |
| continuous process of differentiation of fresh mesoblast cells. It
| |
| soon acquires a lumen, and joins the cloacal section of the
| |
| alimentary tract before the close of foetal life. Before this
| |
| communication is established, the front end of the duct sends a
| |
| process towards the body cavity, the blind end of which acquires
| |
| a ciliated opening into the latter. A series of about four or five
| |
| successively formed outgrowths from the duct, one behind the
| |
| other, give rise to as many ciliated funnels opening into the body
| |
| cavity, and each communicating by a more or less elongated
| |
| tube with the segmental duct. These funnels, which have a
| |
| metameric arrangement, constitute the pronephros, the whole
| |
| of which is situated in the pericardial region of the body
| |
| cavity.
| |
| | |
| On the inner side of the peritoneal openings of each pronephros there is formed a vascular glomerulus, projecting into
| |
| the body cavity, and covered by peritoneal epithelium. For a
| |
| considerable period the pronephros constitutes the sole functional part of the excretory system.
| |
| | |
| A mesonephros is formed (Furbringer) relatively late in
| |
| larval life, as a segmentally arranged series of solid cords,
| |
| derived from the peritoneal epithelium. These cords constitute
| |
| the rudiments of the segmental tubes. They are present for a
| |
| considerable portion of the body cavity, extending backwards
| |
| from a point shortly behind the pronephros. They soon separate
| |
| from the peritoneal epithelium, become hollowed out into canals,
| |
| and join the segmental duct. At their blind extremity (that
| |
| originally connected with the peritoneal epithelium) a Malpighian
| |
| body is formed.
| |
| | |
| The pronephros is only a provisional excretory organ, the
| |
| atrophy of which commences during larval life, and is nearly
| |
| completed when the Ammoccete has reached 180 mm. in length.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 70 1
| |
| | |
| Further changes take place in connection with the excretory
| |
| system on the conversion of the Ammoccete into the adult.
| |
| | |
| The segmental ducts in the adult fall into a common urinogenital cloaca, which opens on a papilla behind the anus. This
| |
| cloaca also communicates by two apertures (abdominal pores)
| |
| with the body cavity. The generative products are carried into
| |
| the cloaca by these pores ; so that their transportation outwards
| |
| is not performed by any part of the primitive urinary system.
| |
| The urinogenital cloaca is formed by the separation of the portion
| |
| of the primitive cloaca containing the openings of the segmental
| |
| ducts from that connected with the alimentary tract.
| |
| | |
| The mesonephros of the Ammoccete undergoes at the metamorphosis complete atrophy, and is physiologically replaced by
| |
| a posterior series of segmental tubes, opening into the hindermost portion of the segmental duct (Schneider).
| |
| | |
| In Myxine the excretory system consists (i) of a highly developed pronephros with a bunch of ciliated peritoneal funnels opening into the pericardial section of the body cavity. The coiled and branched tubes of which
| |
| the pronephros is composed open on the ventral side of the anterior portion
| |
| of the segmental duct, which in old individuals is cut off from the posterior
| |
| section of the duct. On the dorsal side of the portion of the segmental duct
| |
| belonging to the pronephros there are present a small number of diverticula,
| |
| terminating in glomeruli : they are probably to be regarded as anterior
| |
| segmental tubes. (2) Of a mesonephros, which commences a considerable
| |
| distance behind the pronephros, and is formed of straight extremely simple
| |
| segmental tubes opening into the segmental duct (fig. 385).
| |
| | |
| The excretory system of Myxine clearly retains the characters of the
| |
| system as it exists in the larva of Petromyzon.
| |
| | |
| Teleostei. In most Teleostei the pronephros and mesonephros coexist through life, and their products are carried off by
| |
| a duct, the nature of which is somewhat doubtful, but which is
| |
| probably homologous with the mesonephric duct of other types.
| |
| | |
| The system commences in the embryo (Rosenberg, Oellacher,
| |
| Gotte, Furbringer) with the formation of a groove-like fold of the
| |
| somatic layer of peritoneal epithelium, which becomes gradually
| |
| constricted into a canal; the process of constriction commencing
| |
| in the middle and extending in both directions. The canal does
| |
| not however close anteriorly, but remains open to the body
| |
| cavity, thus giving rise to a funnel equivalent to the pronephric
| |
| funnels of Petromyzon and Myxine. On the inner side of this
| |
| | |
| | |
| | |
| 702
| |
| | |
| | |
| | |
| TELEOSTEI.
| |
| | |
| | |
| | |
| funnel there is formed a glomerulus, projecting into the body
| |
| | |
| cavity ; and at the same time that
| |
| | |
| this is being formed the anterior end
| |
| | |
| of the canal becomes elongated and
| |
| | |
| convoluted. The above structures
| |
| | |
| constitute a pronephros, while the
| |
| | |
| posterior part of the primitive canal
| |
| | |
| forms the segmental duct.
| |
| | |
| The portion of the body cavity
| |
| with the glomerulus and peritoneal
| |
| funnel of the pronephros (fig. 395,
| |
| po) soon becomes completely isolated from the remainder, so as to
| |
| form a closed cavity (gl). The
| |
| development of the mesonephros
| |
| does not take place till long after
| |
| that of the pronephros. The segmental tubes which form it are
| |
| stated by Fiirbringer to arise from
| |
| solid ingrowths of peritoneal epithelium, developed successively from
| |
| before backwards, but Sedgwick
| |
| informs me that they arise as differentiations of the mesoblastic cells
| |
| near the peritoneal epithelium. They
| |
| soon become hollow, and unite with
| |
| the segmental duct. Malpighian
| |
| bodies are developed on their median
| |
| portions. They grow very greatly
| |
| in length, and become much convoluted, but the details of this
| |
| process have not been followed out.
| |
| | |
| The foremost segmental tubes are situated close behind the
| |
| pronephros, while the hindermost are in many cases developed
| |
| in the post-anal continuations of the body cavity. The pronephros appears to form the swollen cephalic portion of the kidney
| |
| of the adult, and the mesonephros the remainder ; the so-called
| |
| caudal portion, where present, being derived (?) from the postanal segmental tubes.
| |
| | |
| In some cases the cephalic portion of the kidneys is absent
| |
| | |
| | |
| | |
| | |
| FIG. 394. PORTIONS OF THE
| |
| MESONEPHROS OF MYXINE. (From
| |
| Gegenbaur; after J. Miiller.)
| |
| | |
| a. segmental duct ; b. segmental tube; c. glomerulus ; d. afferent,
| |
| e. efferent artery.
| |
| | |
| B represents a portion of A
| |
| highly magnified.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 703
| |
| | |
| | |
| | |
| in the adult, which probably implies the atrophy of the pronephros ; in other instances the cephalic portion of the kidneys is
| |
| the only part developed. Its relation to the embryonic proncphros requires however further elucidation.
| |
| | |
| In the adult the ducts in the lower part of the kidneys lie as
| |
| a rule on their outer borders, and almost invariably open into a
| |
| | |
| | |
| | |
| | |
| pr
| |
| | |
| | |
| | |
| FIG. 395. SECTION THROUGH THE PRONEPHROS OF A TROUT AND ADJACENT
| |
| PARTS TEN DAYS BEFORE HATCHING.
| |
| | |
| pr.n. pronephros ; po. opening of pronephros into the isolated portion of the body
| |
| cavity containing the glomerulus ; gl. glomerulus ; ao. aorta ; ch. notochord ; x.
| |
| subnotochordal rod ; al. alimentary tract.
| |
| | |
| urinary bladder, which usually opens in its turn on the urinogenital papilla immediately behind the genital pore, but in a few
| |
| instances there is a common urinogenital pore.
| |
| | |
| In most Osseous Fish there are true generative ducts continuous with the investment of the generative organs. It
| |
| appears to me most probable, from the analogy of Lepidostcus,
| |
| to be described in the next section, that these ducts are split off
| |
| from the primitive segmental duct, and correspond with the
| |
| Miillerian ducts of Elasmobranchii, etc. ; though on this point
| |
| we have at present no positive embryological evidence (vide
| |
| general considerations at the end of the Chapter). In the
| |
| female Salmon and the male and female Eel the generative
| |
| products are carried to the exterior by abdominal pores. It is
| |
| possible that this may represent a primitive condition, though it
| |
| | |
| | |
| | |
| 704
| |
| | |
| | |
| | |
| GANOIDEI.
| |
| | |
| | |
| | |
| is more probably a case of degeneration, as is indicated by the
| |
| presence of ducts in the male Salmon and in forms nearly allied
| |
| to the Salmonidae.
| |
| | |
| The coexistence of abdominal pores and generative ducts in
| |
| Mormyrus appears to me to demonstrate that the generative
| |
| ducts in Teleostei cannot be derived from the coalescence of the
| |
| investment of the generative organs with the abdominal pores.
| |
| | |
| Ganoidei. The true excretory gland of the adult Ganoidei
| |
| resembles on the whole that of Teleostei, consisting of an
| |
| elongated band on each side the mesonephros an anterior
| |
| dilatation of which probably represents the pronephros.
| |
| | |
| There is in both sexes a Mullerian duct, provided, except
| |
| in Lepidosteus, with an abdominal funnel, which is however
| |
| situated relatively very far back in the abdominal cavity. The
| |
| Mullerian ducts appear to serve as generative canals in both sexes.
| |
| In Lepidosteus they are continuous with the investment of the
| |
| generative glands, and thus a relation between the generative ducts
| |
| and glands, very similar to that in Teleostei, is brought about.
| |
| | |
| Posteriorly the Mullerian ducts and the ducts of the mesonephros remain united. The common duct so formed on each
| |
| side is clearly the primitive segmental duct. It receives the
| |
| secretion of a certain number of the posterior mesonephric
| |
| tubules, and usually unites with its fellow to form a kind of
| |
| bladder, opening by a single
| |
| pore into the cloaca, behind
| |
| the anus. The duct which
| |
| receives the secretion of the
| |
| anterior mesonephric tubules
| |
| is the true mesonephric or
| |
| Wolffian duct.
| |
| | |
| The development of the
| |
| excretory system, which has
| |
| been partially worked out in
| |
| Acipenscr and Lepidosteus 1 ,
| |
| is on the whole very similar
| |
| to that in the Teleostei. The
| |
| first portion of the system to
| |
| | |
| | |
| | |
| | |
| FIG. 396. SECTION THROUGH THE
| |
| TRUNK OF A LEPIDOSTEUS EMBRYO ON
| |
| THE SIXTH DAY AFTER IMPREGNATION.
| |
| | |
| me. medullary cord ; ms. mesoblast ; sg.
| |
| segmental duct ; ch. notochord ; .r. subnotochordal rod; hy. hypoblast.
| |
| | |
| | |
| | |
| 1 Acipenser has been investigated by Fiirbringer, Salensky, Sedgwick, and also
| |
| by myself, and Lepidosteus by W. N. Parker and myself.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 705
| |
| | |
| | |
| | |
| be formed is the segmental duct. In Lepidosteus this duct is
| |
| formed as a groove-like invagination of the somatic peritoneal
| |
| epithelium, precisely as in Teleostei, and shortly afterwards
| |
| forms a duct lying between the mesoblast and the epiblast
| |
| (fig. 396, sg}. In Acipenser (Salensky) however it is formed as
| |
| | |
| | |
| | |
| | |
| FIG. 397. TRANSVERSE SECTION THROUGH THE ANTERIOR PART OF AN ACIPENSER
| |
| | |
| EMBRYO. (After Salensky.)
| |
| | |
| Rf. medullary groove ; Alp. medullary plate ; Wg. segmental duct ; Ch. notochord ; En. hypoblast ; Sgp. mesoblastic somite ; Sp. parietal part of mesoblastic
| |
| plate.
| |
| | |
| a solid ridge of the somatic mesoblast, as in Petromyzon and
| |
| Elasmobranchii (fig. 397, Wg).
| |
| | |
| In both forms the ducts unite behind with the cloaca, and a
| |
| pronephros of the Teleostean type appears to be developed.
| |
| This gland is provided with but one 1 peritoneal opening, which
| |
| together with the glomerulus belonging to it becomes encapsuled
| |
| in a special section of the body cavity. The opening of the
| |
| pronephros of Acipenser into this cavity is shewn in fig. ^<^>,pr.n.
| |
| At this early stage of Acipenser (larva of 5 mm.) I could find
| |
| no glomerulus.
| |
| | |
| The mesonephros is formed some distance behind, and some
| |
| time after the pronephros, both in Acipenser and Lepidosteus,
| |
| so that in the larvae of both these genera the pronephros is for
| |
| a considerable period the only excretory organ. In Lepidosteus
| |
| especially the development of the mesonephros occurs very
| |
| late.
| |
| | |
| The development of the mesonephros has not been worked
| |
| out in Lepidosteus, but in Acipenser the anterior segmental
| |
| tubes become first established as (I believe) solid cords of cells,
| |
| attached at one extremity to the peritoneal epithelium on each
| |
| | |
| 1 I have not fully proved this point, but have never found more than one
| |
| opening.
| |
| | |
| | |
| | |
| B. III.
| |
| | |
| | |
| | |
| 45
| |
| | |
| | |
| | |
| GANOIDEI.
| |
| | |
| | |
| | |
| side of the insertion of the mesentery, and extending upwards
| |
| and outwards round the segmental duct 1 . The posterior segmental tubes arise later than the anterior, and (as far as can be
| |
| determined from the sections in my possession) they are formed
| |
| independently of the peritoneal epithelium, on the dorsal side
| |
| of the segmental duct.
| |
| | |
| In later stages (larvae of 7 10 mm.) the anterior segmental
| |
| tubes gradually lose their attachment to the peritoneal epithelium. The extremity near the peritoneal epithelium forms a
| |
| Malpighian body, and the other end unites with the segmental
| |
| duct. At a still later stage wide peritoneal funnels are es
| |
| | |
| | |
| sjy.c
| |
| | |
| | |
| | |
| mjo
| |
| | |
| | |
| | |
| pr.n
| |
| | |
| | |
| | |
| | |
| FIG. 398. TRANSVERSE SECTION THROUGH THE REGION OF THE STOMACH OF A
| |
| | |
| LARVA OF ACIPENSER 5 MM. IN LENGTH.
| |
| | |
| st. epithelium of stomach ; yk. yolk ; ch. notochord, below which is a subnotochordal rod; pr.n. pronephros ; ao. aorta; mf. muscle-plate formed of large cells,
| |
| the outer parts of which are differentiated into contractile fibres ; sp.c. spinal cord ;
| |
| b.c. body cavity.
| |
| | |
| tablished, for at any rate a considerable number of the tubes,
| |
| leading from the body cavity to the Malpighian bodies. These
| |
| | |
| 1 Whether the segmental tubes are formed as ingrowths of the peritoneal
| |
| epithelium, or in situ, could not be determined.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 707
| |
| | |
| funnels have been noticed by Furbringer, Salensky and myself,
| |
| but their mode of development has not, so far as I know, been
| |
| made out. The funnels appear to be no longer present in the
| |
| adult. The development of the Mullerian ducts has not been
| |
| worked out.
| |
| | |
| Dipnoi. The excretory system of the Dipnoi is only known in the
| |
| adult, but though in some respects intermediate in character between that of
| |
| the Ganoidei and Amphibia, it resembles that of the Ganoidei in the
| |
| important feature of the Mullerian ducts serving as genital ducts in both
| |
| sexes.
| |
| | |
| Amphibia. In Amphibia (Gotte, Furbringer) the development of the excretory system commences, as in Teleostei, by
| |
| the formation of the segmental duct from a groove formed by a
| |
| fold of the somatic layer of the peritoneal epithelium, near the
| |
| dorsal border of the body cavity (fig. 399, u). The anterior end
| |
| of the groove is placed immediately behind the branchial
| |
| region. Its posterior part soon becomes converted into a canal
| |
| by a constriction which commences a short way from the front
| |
| end of the groove, and thence extends backwards. This canal
| |
| at first ends blindly close to the cloaca, into which however it
| |
| soon opens.
| |
| | |
| The anterior open part of the groove in front of the constriction (fig. 399, n] becomes differentiated into a longitudinal
| |
| duct, which remains in open communication with the body
| |
| cavity by two (many Urodela) three (many Anura) or four
| |
| (Cceciliidae) canals. This constitutes the dorsal part of the
| |
| pronephros. The ventral part of the gland is formed from the
| |
| section of the duct immediately behind the longitudinal canal.
| |
| This part grows in length, and, assuming an S-shaped curvature,
| |
| becomes placed on the ventral side of the first formed part of
| |
| the pronephros. By continuous growth in a limited space the
| |
| convolutions of the canal of the pronephros become more numerous, and the complexity of the gland is further increased by the
| |
| outgrowth of blindly ending diverticula.
| |
| | |
| At the root of the mesentery, opposite the peritoneal openings
| |
| of the pronephros, a longitudinal fold, lined by peritoneal epithelium, and attached by a narrow band of tissue, makes its
| |
| appearance. It soon becomes highly vascular, and constitutes a
| |
| glomerulus homologous with that in Petromyzon and Teleostei.
| |
| | |
| 452
| |
| | |
| | |
| | |
| AMPHIBIA.
| |
| | |
| | |
| | |
| a*'
| |
| | |
| | |
| | |
| The section of the body cavity which contains the openings
| |
| of the pronephros and the glomerulus,
| |
| becomes dilated, and then temporarily
| |
| shut off from the remainder. At a
| |
| later period it forms a special though
| |
| not completely isolated compartment.
| |
| For a long time the pronephros and
| |
| its duct form the only excretory organs
| |
| of larval Amphibia. Eventually however the formation of the mesonephros
| |
| commences, and is followed by the
| |
| atrophy of the pronephros. The mesonephros is composed, as in other
| |
| types, of a series of segmental tubes,
| |
| but these, except in Cceciliidae, no
| |
| longer correspond in number with the
| |
| myotomes, but are in all instances
| |
| more numerous. Moreover, in the
| |
| posterior part of the mesonephros in
| |
| the Urodeles, and through the whole
| |
| length of the gland in other types,
| |
| secondary and tertiary segmental tubes
| |
| are formed in addition to the primary
| |
| tubes.
| |
| | |
| | |
| | |
| | |
| FIG. 399. TRANSVERSE SECTION THROUGH A VERY YOUNG
| |
| TADPOLE OF BOMBINATOR AT
| |
| THE LEVEL OF THE ANTERIOR
| |
| END OF THE YOLK-SACK. (After
| |
| | |
| Gotte.)
| |
| | |
| a. fold of epiblast continuous
| |
| with the dorsal fin; is", neural
| |
| cord; m. lateral muscle; as 1 .
| |
| outer layer of muscle-plate; s.
| |
| lateral plate of mesoblast ; b.
| |
| mesentery ; u. open end of the
| |
| segmental duct, which forms the
| |
| pronephros ; f. alimentary tract ;
| |
| f. ventral diverticulum which
| |
| becomes the liver; e. junction of
| |
| yolk cells and hypoblast cells ;
| |
| d. yolk cells.
| |
| | |
| | |
| | |
| The development of the mesonephros
| |
| commences in Salamandra (Fiirbringer) with
| |
| the formation of a series of solid cords, which
| |
| in the anterior myotomes spring from the
| |
| peritoneal epithelium on the inner side of the
| |
| segmental duct, but posteriorly arise independently of this epithelium in the adjoining
| |
| mesoblast. Sedgwick informs me that in the
| |
| | |
| Frog the segmental tubes are throughout developed in the mesoblast, independently of the peritoneal epithelium. These cords next become detached
| |
| from the peritoneal epithelium (in so far as they are primitively united to it),
| |
| and after first assuming a vesicular form, grow out into coiled tubes, with a
| |
| median limb the blind end of which assists in forming a Malpighian body,
| |
| and a lateral limb which comes in contact with and opens into the segmental
| |
| duct, and an intermediate portion connecting the two. At the junction of
| |
| the median with the intermediate portion, and therefore at the neck of the
| |
| Malpighian body, a canal grows out in a ventral direction, which meets the
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 709
| |
| | |
| peritoneal epithelium, and then develops a funnel-shaped opening into the
| |
| body cavity, which subsequently becomes ciliated. In this way the peritoneal
| |
| funnels which are present in the adult are established.
| |
| | |
| The median and lateral sections of the segmental tubes become highly
| |
| convoluted, and the separate tubes soon come into such close proximity that
| |
| their primitive distinctness is lost.
| |
| | |
| The first fully developed segmental tube is formed in Salamandra maculata in about the sixth myotome behind the pronephros. But in the region
| |
| between the two structures rudimentary segmental tubes are developed.
| |
| | |
| The number of primary segmental tubes in the separate myotomes of
| |
| Salamandra is as follows :
| |
| | |
| In the 6th myotome (i.e. the first with a true
| |
| | |
| segmental tube) 12 segmental tubes
| |
| | |
| yth roth myotome 23
| |
| | |
| IIth ... 34
| |
| | |
| I2th 3 4 or 4 5
| |
| | |
| I3th y> 45
| |
| | |
| 1 3th i6th 56
| |
| | |
| It thus appears that the segmental tubes are not only more numerous than
| |
| the myotomes, but that the number in each myotome increases from before
| |
| backwards. In the case of Salamandra there are formed in the region of
| |
| the posterior (10 16) myotomes secondary, tertiary, etc. segmental tubes out
| |
| of independent solid cords, which arise in the mesoblast dorsally to the tubes
| |
| already established.
| |
| | |
| The secondary segmental tubes appear to develop out of these cords
| |
| exactly in the same way as the primary ones, except that they do not join the
| |
| segmental duct directly, but unite with the primary segmental tubes shortly
| |
| before the junction of the latter with the segmental duct. In this way compound segmental tubes are established with a common collecting tube, but
| |
| with numerous Malpighian bodies and ciliated peritoneal openings. The
| |
| difference in the mode of origin of these compound tubes and of those in
| |
| Elasmobranchii is very striking.
| |
| | |
| The later stages in the development of the segmental tubes have not been
| |
| studied in the other Amphibian types.
| |
| | |
| In Cceciliidas the earliest stages are not known, but the tubes present in
| |
| the adult (Spengel) a truly segmental arrangement, and in the young each of
| |
| them is single, and provided with only a single peritoneal funnel. In the
| |
| adult however many of the segmental organs become compound, and may
| |
| have as many as twenty funnels, etc. Both simple and compound segmental
| |
| tubes occur in all parts of the mesonephros, and are arranged in no definite
| |
| order.
| |
| | |
| In the Anura (Spengel) all the segmental tubes are compound, and an
| |
| enormous number of peritoneal funnels are present on the ventral surface,
| |
| but it has not yet been definitely determined into what part of the segmental
| |
| tubes they open.
| |
| | |
| | |
| | |
| 710 AMPHIBIA.
| |
| | |
| | |
| | |
| Before dealing with the further changes of the Wolffian body
| |
| it is necessary to return to the segmental duct, which, at the
| |
| time when the pronephros is undergoing atrophy, becomes split
| |
| into a dorsal Wolffian and ventral Mullerian duct. The process
| |
| in Salamandra (Fiirbringer) has much the same character as in
| |
| Elasmobranchii, the Mullerian duct being formed by the gradual
| |
| separation, from before backwards, of a solid row of cells from
| |
| the ventral side of the segmental duct, the remainder of the duct
| |
| constituting the Wolffian duct. During the formation of the
| |
| Mullerian duct its anterior part becomes hollow, and attaching
| |
| itself in front to the peritoneal epithelium acquires an opening
| |
| into the body cavity. The process of hollowing is continued
| |
| backwards pari passu with the splitting of the segmental duct.
| |
| In the female the process is continued till the Mullerian duct
| |
| opens, close to the Wolffian duct, into the cloaca. In the male
| |
| the duct usually ends blindly. It is important to notice that
| |
| the abdominal opening of the Mullerian duct in the Amphibia
| |
| (Salamandra) is a formation independent of the pronephros, and
| |
| placed slightly behind it ; and that the undivided anterior part
| |
| of the segmental duct (with the pronephros) is not, as in Elasmobranchii, united with the Mullerian duct, but remains connected
| |
| with the Wolffian duct.
| |
| | |
| The development of the Mullerian duct has not been satisfactorily
| |
| studied in other forms besides Salamandra. In Cceciliidae its abdominal
| |
| opening is on a level with the anterior end of the Wolffian body. In other
| |
| forms it is usually placed very far forwards, close to the root of the lungs
| |
| (except in Proteus and Batrachoseps, where it is placed somewhat further
| |
| back), and some distance in front of the Wolffian body.
| |
| | |
| The Mullerian duct is always well developed in the female, and serves as
| |
| oviduct. In the male it does not (except possibly in Alytes) assist in the
| |
| transportation of the genital products, and is always more or less rudimentary, and in Anura may be completely absent.
| |
| | |
| After the formation of the Mullerian duct, the Wolffian duct
| |
| remains as the excretory channel for the Wolffian body, and, till
| |
| the atrophy of the pronephros, for this gland also. Its anterior
| |
| section, in front of the Wolffian body, undergoes a more or less
| |
| complete atrophy.
| |
| | |
| The further changes of the excretory system concern (i) the
| |
| junction in the male of the anterior part of the Wolffian body
| |
| with the testis ; (2) certain changes in the collecting tubes of the
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 711
| |
| | |
| | |
| | |
| posterior part of the mesonephros. The first of these processes
| |
| results in the division of the Wolffian body into a sexual and a
| |
| non-sexual part, and in Salamandra and other Urodeles the
| |
| division corresponds with the distribution of the simple and
| |
| compound segmental tubes.
| |
| | |
| Since the development of the canals connecting the testes with
| |
| the sexual part of the Wolffian body has not been in all points
| |
| satisfactorily elucidated, it will be convenient to commence with a
| |
| description of the adult arrangement of the parts (fig. 400 B). In
| |
| most instances a non-segmental system of canals the vasa effcrentia (ve) coming from the testis, fall into a canal known as the
| |
| longitudinal canal of the Wolffian body, from which there pass off
| |
| transverse canals, which fall into, and are equal in number to, the
| |
| primary Malpighian bodies of the sexual part of the gland. The
| |
| spermatozoa, brought to the Malpighian bodies, are thence transported along the segmental tubes to the Wolffian duct, and so to
| |
| the exterior. The system of canals connecting the testis with
| |
| the Malpighian bodies is known as the testicular network. The
| |
| number of segmental tubes connected with the testis varies
| |
| very greatly. In Siredon there are as many as from 30 32
| |
| (Spengel).
| |
| | |
| The longitudinal canal of the Wolffian body is in rare instances
| |
| (Spelerpes, etc.) absent, where the sexual part of the Wolffian body is
| |
| slightly developed. In the Urodela the testes are united with the anterior
| |
| part of the Wolffian body. In the Cceciliidas the junction takes place in an
| |
| homologous part of the Wolffian body, but, owing to the development of the
| |
| anterior segmental tubes, which are rudimentary in the Urodela, it is
| |
| situated some way behind the front end. Amongst the Anura the connection
| |
| of the testis with the tubules of the Wolffian body is subject to considerable
| |
| variations. In Bufo cinereus the normal Urodele type is preserved, and in
| |
| Bombinator the same arrangement is found in a rudimentary condition, in
| |
| that there are transverse trunks from the longitudinal canal of the Wolffian
| |
| body, which end blindly, while the semen is carried into the Wolffian
| |
| duct by canals in front of the Wolffian body. In Alytes and Discoglossus
| |
| the semen is carried away by a similar direct continuation of the longitudinal canal in front of the Wolffian body, but there are no rudimentary transverse canals passing into the Wolffian body, as in Bombinator. In Rana the transverse ducts which pass off from the longitudinal
| |
| canal of the Wolffian body, after dilating to form (?) rudimentary Malpighian
| |
| bodies, enter directly into the collecting tubes near their opening into the
| |
| Wolffian duct.
| |
| | |
| | |
| | |
| 712 AMPHIBIA.
| |
| | |
| | |
| | |
| In most Urodeles the peritoneal openings connected with the primary
| |
| generative Malpighian bodies atrophy, but in Spelerpes they persist. In
| |
| the Cceciliidie they also remain in the adult state.
| |
| | |
| With reference to the development of these parts little is
| |
| known except that the testicular network grows out from the
| |
| primary Malpighian bodies, and becomes united with the testis.
| |
| Embryological evidence, as well as the fact of the persistence of
| |
| the peritoneal funnels of the generative region in the adults
| |
| of some forms, proves that the testicular network is not developed
| |
| from the peritoneal funnels.
| |
| | |
| Rudiments of the testicular network are found in the female Cceciliidae
| |
| and in the females of many Urodela (Salamandra, Triton). These rudiments may in their fullest development consist of a longitudinal canal and
| |
| of transverse canals passing from this to the Malpighian bodies, together
| |
| with some branches passing into the mesovarium.
| |
| | |
| Amongst the Urodela the collecting tubes of the hinder non-sexual part
| |
| of the Wolffian body, which probably represents a rudimentary metanephros,
| |
| undergo in the male sex a change similar to that which they usually undergo
| |
| in Elasmobranchii. Their points of junction with the Wolffian duct are
| |
| carried back to the hindermost end of the duct (fig. 400 B), and the collecting
| |
| tubes themselves unite together into one or more short ducts (ureters) before
| |
| joining the Wolffian duct.
| |
| | |
| In Batrachoseps only the first collecting tube becomes split off in
| |
| this way ; and it forms a single elongated ureter which receives all the
| |
| collecting tubes of the posterior segmental tubes. In the female and in
| |
| the male of Proteus, Menobranchus, and Siren the collecting tubes retain
| |
| their primitive transverse course and open laterally into the Wolffian duct.
| |
| In rare cases (Ellipsoglossus, Spengel} the ureters open directly into the
| |
| cloaca.
| |
| | |
| The urinary bladder of the Amphibia is an outgrowth of the
| |
| ventral wall of the cloacal section of the alimentary tract, and is
| |
| homologous with the allantois of the amniotic Vertebrata.
| |
| | |
| The subjoined diagram (fig. 400) of the urogenital system of
| |
| Triton illustrates the more important points of the preceding
| |
| description.
| |
| | |
| In the female (A) the following parts are present :
| |
| | |
| (1) The Mullerian duct or oviduct (od) derived from the
| |
| splitting of the segmental duct.
| |
| | |
| (2) The Wolffian duct (sug) constituting the portion of the
| |
| segmental duct left after the formation of the Mullerian duct.
| |
| | |
| (3) The mesonephros (r), divided into an anterior sexual part
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 7'3
| |
| | |
| | |
| | |
| connected with a rudimentary testicular network, and a posterior
| |
| part. The collecting tubes from both
| |
| parts fall transversely into the Wolffian duct.
| |
| | |
| (4) The ovary (ov).
| |
| | |
| (5) The rudimentary testicular
| |
| network.
| |
| | |
| In the male (B) the following
| |
| parts are present :
| |
| | |
| (1) The functionless though fairly
| |
| developed Miillerian duct (;).
| |
| | |
| (2) The Wolffian duct (sug).
| |
| | |
| (3) The mesonephros (r) divided
| |
| into a true sexual part, through the
| |
| segmental tubes of which the semen
| |
| passes, and a non-sexual part. The
| |
| collecting tubes of the latter do not
| |
| enter the Wolffian duct directly, but
| |
| bend obliquely backwards and only
| |
| fall into it close to its cloacal aperture, after uniting to form one or two
| |
| primary tubes (ureters).
| |
| | |
| (4) The testicular network (ve)
| |
| consisting of (i) transverse ducts
| |
| from the testes, falling into (2) the
| |
| longitudinal canal of the Wolffian
| |
| body, from which (3) transverse canals are again given off to the Malpighian bodies.
| |
| | |
| Amniota. The amniotic Vertebrata agree, so far as is known, very
| |
| closely amongst themselves in the
| |
| formation of the urinogenital system.
| |
| | |
| The most characteristic feature of the system is the full
| |
| development of a metanephros, which constitutes the functional
| |
| kidney on the atrophy of the mesonephros or Wolffian body,
| |
| which is a purely embryonic organ. The first part of the
| |
| system to develop is a duct, which is usually spoken of as the
| |
| Wolffian duct, but which is really the homologue of the seg
| |
| | |
| | |
| | |
| FIG. 400. DIAGRAM OF THE
| |
| URINOGENITAL SYSTEM OF TRITON. (From Gegenbaur ; after
| |
| Spengel.)
| |
| | |
| A. Female. B. Male.
| |
| r. mesonephros, on the surface
| |
| of which numerous peritoneal funnels are visible ; sug. mesonephric
| |
| or Wolffian duct; od. oviduct
| |
| (Miillerian duct); in. Miillerian
| |
| duct of male ; ve. vasa efferentia of
| |
| testis ; t. testis ; ov. ovary ; up.
| |
| urinogenital pore.
| |
| | |
| | |
| | |
| 714 AMNIOTA.
| |
| | |
| | |
| | |
| mental duct. It apparently develops in all the Amniota nearly
| |
| on the Elasmobranch type, as a solid rod, primarily derived
| |
| from the somatic mesoblast of the intermediate cell mass (fig.
| |
| 401 W.d}\
| |
| | |
| The first trace of it is visible in an embryo Chick with eight
| |
| somites, as a ridge projecting from the intermediate cell mass towards the epiblast in the region of the seventh somite. In the
| |
| course of further development it continues to constitute such a
| |
| ridge as far as the eleventh somite (Sedgwick), but from this
| |
| point it grows backwards in the space between the epiblast and
| |
| mesoblast In an embryo with fourteen somites a small lumen
| |
| has appeared in its middle part and in front it is connected with
| |
| rudimentary Wolffian tubules, which develop in continuity with
| |
| it (Sedgwick). In the succeeding stages the lumen of the duct
| |
| gradually extends backwards and forwards, and the duct itself
| |
| also passes inwards relatively to the epiblast (fig. 402). Its hindend elongates till it comes into connection with, and opens into,
| |
| the cloacal section of the hind-gut' 2 .
| |
| | |
| It might have been anticipated that, as in the lower types,
| |
| the anterior end of the segmental duct would either open into
| |
| the body cavity, or come into connection with a pronephros.
| |
| Neither of these occurrences takes place, though in some types
| |
| (the Fowl) a structure, which is probably the rudiment of a
| |
| pronephros, is developed ; it does not however appear till a later
| |
| stage, and is then unconnected with the segmental duct. The
| |
| next part of the system to appear is the mesonephros or
| |
| Wolffian body.
| |
| | |
| This is formed in all Amniota as a series of segmental tubes,
| |
| which in Lacertilia (Braun) correspond with the myotomes, but
| |
| in Birds and Mammalia are more numerous.
| |
| | |
| In Reptilia (Braun, No. 542), the mesonephric tubes develop as segmentally-arranged masses on the inner side of the Wolffian duct, and
| |
| appear to be at first united with the peritoneal epithelium. Each mass soon
| |
| becomes an oval vesicle, probably opening for a very short period into the
| |
| | |
| 1 Dansky and Kostenitsch (No. 543) describe the Wolffian duct in the Chick as
| |
| developing from a groove opening to the peritoneal cavity, which subsequently
| |
| becomes constricted into a duct. I have never met with specimens such as those
| |
| figured by these authors.
| |
| | |
| 2 The foremost extremity of the segmental duct presents, according to Gasser,
| |
| curious irregularities and an anterior completely isolated portion is often present.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 715
| |
| | |
| | |
| | |
| peritoneal cavity by a peritoneal funnel. The vesicles become very early
| |
| detached from the peritoneal epithelium, and lateral outgrowths from them
| |
| give rise to the main parts of the segmental tubes, which soon unite with the
| |
| segmental duct.
| |
| | |
| In Birds the development of the segmental tubes is more complicated 1 .
| |
| | |
| The tubules of the Wolffian body are derived from the intermediate cell
| |
| mass, shewn in fig. 401, between the upper end of the body cavity and the
| |
| | |
| | |
| | |
| g.o.
| |
| | |
| | |
| | |
| | |
| FIG. 401. TRANSVERSE SECTION THROUGH THE DORSAL REGION OF AN
| |
| | |
| EMBRYO CHICK OF 45 HOURS.
| |
| | |
| M.c. medullary canal ; P.v. mesoblastic somite ; W.d. Wolffian duct which is in
| |
| contact with the intermediate cell mass ; So. somatopleure ; S.p. splanchnopleure ;
| |
| p.p. pleuroperitoneal cavity ; ch. notochord ; op. boundary of area opaca; v. bloodvessel.
| |
| | |
| muscle-plate. In the Chick the mode of development of this mass into the
| |
| segmental tubules is different in the regions in front of and behind about the
| |
| sixteenth segment. In front of about the sixteenth segment the intermediate
| |
| cell mass becomes detached from the peritoneal epithelium at certain points,
| |
| remaining attached to it at other points, there being several such to each
| |
| segment. The parts of the intermediate cell mass attached to the peritoneal
| |
| epithelium become converted into S-shaped cords (fig. 402, st] which soon
| |
| unite with the segmental duct (wd}. Into the commencement of each
| |
| of these cords the lumen of the body cavity is for a short distance
| |
| prolonged, so that this part constitutes a rudimentary peritoneal funnel.
| |
| | |
| 1 Correct figures of the early stages of these structures were first given by
| |
| Kolliker, but the correct interpretation of them and the first satisfactory account of
| |
| the development of the excretory organs of Birds was given by Sedgwick (No. 549).
| |
| | |
| | |
| | |
| 716
| |
| | |
| | |
| | |
| AMNIOTA.
| |
| | |
| | |
| | |
| In the Duck the attachment of the intermediate cell mass to the peritoneal
| |
| epithelium is prolonged further back than in the Chick.
| |
| | |
| In the foremost segmental tubes, which never reach a very complete
| |
| development, the peritoneal funnels widen considerably, while at the same
| |
| time they acquire a distinct lumen. The section of the tube adjoining
| |
| the wide peritoneal funnel becomes partially invaginated by the formation of
| |
| a glomerulus, and this glomerulus soon grows to such an extent as to project
| |
| through the peritoneal funnel, the neck of which it completely fills, into the
| |
| body cavity (fig. 403, gl). There is thus formed a series of free peritoneal
| |
| glomeruli belonging to the anterior Wolfnan tubuli 1 . These tubuli become
| |
| however early aborted.
| |
| | |
| In the case of the remaining tubules developed from the S-shaped cords
| |
| the attachment to the peritoneal epithelium is very soon lost. The cords
| |
| acquire a lumen, and open into the segmental duct. Their blind extremities
| |
| constitute the rudiments of Malpighian bodies.
| |
| | |
| | |
| | |
| am
| |
| | |
| | |
| | |
| | |
| FIG. 402. TRANSVERSE SECTION THROUGH THE TRUNK OF A DUCK EMBRYO WITH
| |
| | |
| ABOUT TWENTY-FOUR MESOBLASTIC SOMITES.
| |
| | |
| am. amnion ; so. somatopleure ; sp. splanchnopleure ; ivd. Wolffian duct ; st. segmental tube; ca.v. cardinal vein; m.s. muscle-plate; sp.g. spinal ganglion; sp.c.
| |
| spinal cord ; ch. notochord ; ao. aorta ; hy. hypoblast.
| |
| | |
| 1 These external glomeruli were originally mistaken by me (No. 539) for the
| |
| glomeralus of the pronephros, from their resemblance to the glomerulus of the
| |
| Amphibian pronephros. Their true meaning was made out by Sedgwick (No.
| |
| 550).
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 717
| |
| | |
| | |
| | |
| In the posterior part of the Wolffian body of the Chick the intermediate
| |
| cell mass becomes very early detached from the peritoneal epithelium, and
| |
| at a considerably later period breaks up into oval vesicles similar to those of
| |
| the Reptilia, which form the rudiments of the segmental tubes.
| |
| | |
| Secondary and tertiary segmental tubules are formed in the Chick, on the
| |
| dorsal side of the primary tubules,
| |
| as direct differentiations of the mesoblast. They open independently into
| |
| the Wolffian duct.
| |
| | |
| In Mammalia the segmental tubules (Egli) are formed as solid masses
| |
| in the same situation as in Birds and
| |
| Reptiles. It is not known whether
| |
| they are united with the peritoneal
| |
| epithelium. They soon become oval
| |
| vesicles, which develop into complete
| |
| tubules in the manner already indicated.
| |
| | |
| | |
| | |
| | |
| After the establishment of
| |
| the Wolffian body there is formed
| |
| in both sexes in all the Amniota
| |
| a duct, which in the female
| |
| becomes the oviduct, but which
| |
| is functionless and disappears
| |
| more or less completely in the
| |
| male. This duct, in spite of certain peculiarities in its development, is without doubt homologous with the Mullerian duct of
| |
| | |
| | |
| | |
| FIG. 403. SECTION THROUGH THE
| |
| EXTERNAL GLOMERULUS OF ONE OF
| |
| THE ANTERIOR SEGMENTAL TUBES OF
| |
| AN EMBRYO CHICK OF ABOUT IOO H.
| |
| | |
| gl. glomerulus ; ge. peritoneal epithelium ; Wd. Wolffian duct ; ao.
| |
| aorta ; me. mesentery. The segmental
| |
| tube, and the connection between the
| |
| external and internal parts of the glomerulus are not shewn in this figure.
| |
| | |
| | |
| | |
| | |
| FIG. 404. SECTIONS SHEWING TWO OF THE PERITONEAL INVAGINATIONS WHICH
| |
| GIVE RISE TO THE ANTERIOR PART OF THE MULLERIAN DUCT (PRONEPHROS).
| |
| (After Balfour and Sedgwick. )
| |
| | |
| A is the nth section of the series.
| |
| B i 5th
| |
| | |
| C i8th ,, ,,
| |
| | |
| gri. second groove ; gr$. third groove ; ri. second ridge ; wit. Wolffian duct.
| |
| | |
| | |
| | |
| 7 i8
| |
| | |
| | |
| | |
| AMNIOTA.
| |
| | |
| | |
| | |
| the Ichthyopsida. In connection with its anterior extremity
| |
| certain structures have been found in the Fowl, which are
| |
| probably, on grounds to be hereafter stated, homologous with
| |
| the pronephros (Balfour and Sedgwick).
| |
| | |
| The pronephros, as I shall call it, consists of a slightly
| |
| convoluted longitudinal canal with three or more peritoneal
| |
| openings. In the earliest condition, it consists of three successive
| |
| open involutions of the peritoneal epithelium, connected together
| |
| by more or less well-defined ridge-like thickenings of the
| |
| epithelium. It takes its origin from the layer of thickened
| |
| peritoneal epithelium situated near the dorsal angle of the body
| |
| cavity, and is situated some considerable distance behind the
| |
| front end of the Wolfifian duct.
| |
| | |
| In a slightly later stage the ridges connecting the grooves
| |
| become partially constricted off from the peritoneal epithelium,
| |
| | |
| | |
| | |
| | |
| FIG. 405. SECTION OF THE WOLFFIAN BODY DEVELOPING PRONEPHROS AND
| |
| GENITAL GLAND OF THE FOURTH DAY. (After Waldeyer.) Magnified 160 times.
| |
| m. mesentery; Z. somatopleure ; a', portion of the germinal epithelium from
| |
| which the involution (2) to form the pronephros (anterior part of Miillerian duct) takes
| |
| place; a. thickened portion of the germinal epithelium in which the primitive
| |
| germinal cells C and o are lying ; E. modified mesoblast which will form the stroma
| |
| of the ovary ; WK. Wolffian body ; y. Wolffian duct.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 719
| |
| | |
| and develop a lumen. The condition of the structure at this
| |
| stage is illustrated by fig. 404, representing three transverse
| |
| sections through two grooves, and through the ridge connecting
| |
| them.
| |
| | |
| The pronephros may in fact now be described as a slightly
| |
| convoluted duct, opening into the body cavity by three groovelike apertures, and continuous behind with the rudiment of the
| |
| true Miillerian duct.
| |
| | |
| The stage just described is that of the fullest development
| |
| of the pronephros. In it, as in all the previous stages, there
| |
| appear to be only three main openings into the body cavity ; but
| |
| in some sections there are indications of the possible presence of
| |
| one or two additional rudimentary grooves.
| |
| | |
| In an embryo not very much older than the one last
| |
| described the pronephros atrophies as such, its two posterior
| |
| openings vanishing, and its anterior opening remaining as the
| |
| permanent opening of the Miillerian duct.
| |
| | |
| The pronephros is an extremely transitory structure, and its
| |
| development and atrophy are completed between the QOth and
| |
| i2Oth hours of incubation.
| |
| | |
| The position of the pronephros in relation to the Wolffian
| |
| body is shewn in fig. 405, which probably passes through a
| |
| region between two of the peritoneal openings. As long as the
| |
| pronephros persists, the Mullerian duct consists merely of a very
| |
| | |
| | |
| | |
| | |
| FlG. 406. TWO SECTIONS SHEWING THE JUNCTION OF THE TERMINAL SOLID
| |
| PORTION OF THE MtJLLERIAN DUCT WITH THE WOLFFIAN DUCT. (After Balfour
| |
| | |
| and Sedgwick.)
| |
| | |
| In A the terminal portion of the duct is quite distinct ; in B it has united with the
| |
| walls of the Wolffian duct.
| |
| | |
| md. Mullerian duct ; Wd. Wolffian duct.
| |
| | |
| | |
| | |
| 72O AMNIOTA.
| |
| | |
| | |
| | |
| small rudiment, continuous with the hindermost of the three
| |
| peritoneal openings, and its solid extremity appears to unite
| |
| with the walls of the Wolffian duct.
| |
| | |
| After the atrophy of the pronephros, the Miillerian duct
| |
| commences to grow rapidly, and for the first part of its course it
| |
| appears to be split off as a solid rod from the outer or ventral
| |
| wall of the Wolffian duct (fig. 406). Into this rod the lumen,
| |
| present in its front part, subsequently extends. Its mode of
| |
| development in front is thus precisely similar to that of the
| |
| Miillerian duct in Elasmobranchii and Amphibia.
| |
| | |
| This mode of development only occurs however in the
| |
| anterior part of the duct. In the posterior part of its course its
| |
| growing point lies in a bay formed by the outer walls of the
| |
| Wolffian duct, but does not become definitely attached to that
| |
| duct. It seems however possible that, although not actually
| |
| split off from the walls of the Wolrfian duct, it may grow backwards from cells derived from that duct.
| |
| | |
| The Miillerian duct finally reaches the cloaca though it does
| |
| not in the female for a long time open into it, and in the male
| |
| never does so.
| |
| | |
| The mode of growth of the Miillerian duct in the posterior part of its
| |
| course will best be understood from the following description quoted from
| |
| the paper by Sedgwick and myself.
| |
| | |
| "A few sections before its termination the Miillerian duct appears as a
| |
| well-defined oval duct lying in contact with the wall of the Wolffian duct on
| |
| the one hand and the germinal epithelium on the other. Gradually, however,
| |
| as we pass backwards, the Miillerian duct dilates ; the external wall of the
| |
| Wolffian duct adjoining it becomes greatly thickened and pushed in in its
| |
| middle part, so as almost to touch the opposite wall of the duct, and so form
| |
| a bay in which the Miillerian duct lies. As soon as the Miillerian duct has
| |
| come to lie in this bay its walls lose their previous distinctness of outline,
| |
| and the cells composing them assume a curious vacuolated appearance. No
| |
| well-defined line of separation can any longer be traced between the walls of
| |
| the Wolffian duct and those of the Miillerian, but between the two is a
| |
| narrow clear space traversed by an irregular network of fibres, in some of
| |
| the meshes of which nuclei are present.
| |
| | |
| The Miillerian duct may be traced in this condition for a considerable
| |
| number of sections, the peculiar features above described becoming more
| |
| and more marked as its termination is approached. It continues to dilate
| |
| and attains a maximum size in the section or so before it disappears. A
| |
| lumen may be observed in it up to its very end, but is usually irregular in
| |
| outline and frequently traversed by strands of protoplasm. The Miillerian
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 721
| |
| | |
| duct finally terminates quite suddenly, and in the section immediately
| |
| behind its termination the Wolffian duct assumes its normal appearance,
| |
| and the part of its outer wall on the level of the Miillerian duct conies into
| |
| contact with the germinal epithelium."
| |
| | |
| Before describing the development of the Mullerian duct in other
| |
| Amniotic types it will be well to say a few words as to the identifications
| |
| above adopted. The identification of the duct, usually called the Wolffian
| |
| duct, with the segmental duct (exclusive of the pronephros) appears to be
| |
| morphologically justified for the following reasons : (i) that it gives rise to
| |
| part of the Mullerian duct as well as to the duct of the Wolffian body ;
| |
| behaving in this respect precisely as does the segmental duct of Elasmobranchii and Amphibia. (2) That it serves as the duct for the Wolffian
| |
| body, before the Mullerian duct originates from it. (3) That it develops in a
| |
| manner strikingly similar to that of the segmental duct of various lower
| |
| forms.
| |
| | |
| With reference to the pronephros it is obvious that the organ identified
| |
| as such is in many respects similar to the pronephros of the Amphibia.
| |
| Both consist of a somewhat convoluted longitudinal canal, with a certain
| |
| number of peritoneal openings ;
| |
| | |
| The main difficulties in the homology are :
| |
| | |
| (1) the fact that the pronephros in the Bird is not united with the
| |
| segmental duct ;
| |
| | |
| (2) the fact that it is situated behind the front end of the Wolffian body.
| |
| It is to be remembered in connection with the first of these difficulties
| |
| | |
| that in the formation of the Mullerian duct in Elasmobranchii the anterior
| |
| undivided extremity of the primitive segmental duct, with the peritoneal
| |
| opening, which probably represents the pronephros, is attached to the
| |
| Mullerian duct, and not to the Wolffian duct ; though in Amphibia the
| |
| reverse is the case. To explain the discontinuity of the pronephros with the
| |
| segmental duct it is only necessary to suppose that the segmental duct and
| |
| pronephros, which in the Ichthyopsida develop as a single formation,
| |
| develop in the Bird as two independent structures a far from extravagant
| |
| supposition, considering that the pronephros in the Bird is undoubtedly
| |
| quite functionless.
| |
| | |
| With reference to the posterior position of the pronephros it is only
| |
| necessary to remark that a change in position might easily take place after
| |
| the acquirement of an independent development, and that the shifting is
| |
| probably correlated with a shifting of the abdominal opening of the
| |
| Mullerian duct.
| |
| | |
| The pronephros has only been observed in Birds, and is very
| |
| possibly not developed in other Amniota. The Mullerian duct
| |
| is also usually stated to develop as a groove of the peritoneal
| |
| epithelium, shewn in the Lizard in fig. 354, md., which is continued backward as a primitively solid rod in the space between
| |
| B. ill. 46
| |
| | |
| | |
| | |
| 722
| |
| | |
| | |
| | |
| AM N IOTA.
| |
| | |
| | |
| | |
| the Wolffian duct and peritoneal epithelium, without becoming
| |
| attached to the Wolffian duct.
| |
| | |
| On the formation of the Miillerian duct, the duct of the
| |
| mesonephros becomes the true mesonephric or Wolffian duct.
| |
| | |
| After these changes have taken place a new organ of great
| |
| importance makes its appearance. This organ is the permanent
| |
| kidney, or metanephros.
| |
| | |
| Metanephros. The mode of development of the metanephros has as yet only been satisfactorily elucidated in the Chick
| |
| (Sedgwick, No. 549). The ureter and the collecting tubes of
| |
| the kidney are developed from a dorsal outgrowth of the hinder
| |
| part of the Wolffian duct. The outgrowth from the Wolffian
| |
| duct grows forwards, and extends along the outer side of a mass
| |
| of mesoblastic tissue which lies mainly behind, but somewhat
| |
| overlaps the dorsal aspect of the Wolffian body.
| |
| | |
| This mass of mesoblastic cells may be called the metanephric blastema. Sedgwick, of the accuracy of whose
| |
| account I have satisfied myself, has shewn that in the Chick it is
| |
| derived from the intermediate cell mass of the region of about
| |
| the thirty-first to the thirty-fourth somite. It is at first continuous with, and indistinguishable in structure from, the portion
| |
| of the intermediate cell mass of the region immediately in front
| |
| of it, which breaks up into Wolffian tubules. The metanephric
| |
| blastema remains however quite passive during the formation of
| |
| the Wolffian tubules in the adjoining blastema ; and on the
| |
| formation of the ureter breaks off from the Wolffian body in
| |
| front, and, growing forwards and dorsalwards, places itself on
| |
| the inner side of the ureter in the position just described.
| |
| | |
| In the subsequent development of the kidney collecting tubes
| |
| grow out from the ureter, and become continuous with masses of
| |
| cells of the metanephric blastema, which then differentiate themselves into the kidney tubules.
| |
| | |
| The process just described appears to me to prove that the
| |
| kidney of the A mniota is a specially differentiated posterior section
| |
| of the primitive mesonephros.
| |
| | |
| According to the view of Remak and Kolliker the outgrowths from the
| |
| ureter give rise to the whole of the tubuli uriniferi and the capsules of the
| |
| Malpighian bodies, the mesoblast around them forming blood-vessels, etc.
| |
| On the other hand some observers (Kupffer, Bornhaupt, Braun) maintain, in
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 723
| |
| | |
| | |
| | |
| accordance with the account given above, that the outgrowths of the ureter
| |
| form only the collecting tubes, and that the secreting tubuli, etc. are formed
| |
| in situ in the adjacent mesoblast.
| |
| | |
| Braun (No. 542) has arrived at the conclusion that in the Lacertilia the
| |
| tissue, out of which the tubuli of the metanephros are formed, is derived
| |
| from irregular solid ingrowths of the peritoneal epithelium, in a region
| |
| behind the Wolffian body, but in a position corresponding to that in which
| |
| the segmental tubes take their origin. These ingrowths, after separating
| |
| from the peritoneal epithelium, unite together to form a cord into which the
| |
| ureter sends the lateral outgrowths already described. These outgrowths
| |
| unite with secreting tubuli and Malpighian bodies, formed in situ. In
| |
| Lacertilia the blastema of the kidney extends into a postanal region.
| |
| Braun's account of the origin of the metanephric blastema does not appear
| |
| to me to be satisfactorily demonstrated.
| |
| | |
| The ureter does not long remain attached to the Wolffian
| |
| duct, but its opening is gradually carried back, till (in the Chick
| |
| between the 6th and 8th day) it opens independently into the
| |
| cloaca.
| |
| | |
| Of the further changes in the excretory system the most important is the atrophy of the greater part of the Wolffian body,
| |
| and the conversion of the Wolffian duct in the male sex into the
| |
| vas deferens, as in Amphibia and the Elasmobranchii.
| |
| | |
| The mode of connection of the testis with the Wolffian duct
| |
| is very remarkable, but may be derived from the primitive
| |
| arrangement characteristic of Elasmobranchii and Amphibia.
| |
| | |
| In the structures connecting the testis with the Wolffian body
| |
| two parts have to be distinguished, (i) that equivalent to the
| |
| testicular network of the lower types, (2) that derived from the
| |
| segmental tubes. The former is probably to be found in peculiar
| |
| outgrowths from the Malpighian bodies at the base of the testes.
| |
| | |
| These were first discovered by Braun in Reptilia, and consist
| |
| in this group of a series of outgrowths from the primary (?)
| |
| Malpighian bodies along the base of the testis : they unite to
| |
| form an interrupted cord in the substance of the testis, from
| |
| which the testicular tubuli (with the exception of the seminiferous cells) are subsequently differentiated. These outgrowths,
| |
| with the exception of the first two or three, become detached
| |
| from the Malpighian bodies. Outgrowths similar to those in
| |
| the male are found in the female, but subsequently atrophy.
| |
| | |
| Outgrowths homologous with those found by Braun have
| |
| | |
| 46 2
| |
| | |
| | |
| | |
| 724 AMNIOTA.
| |
| | |
| | |
| | |
| been detected by myself (No. 555) in Mammals. It is not
| |
| certain to what parts of the testicular tubuli they give rise, but
| |
| they probably form at any rate the vasa recta and rete vasculosum.
| |
| | |
| In Mammals they also occur in the female, and give rise to
| |
| cords of tissue in the ovary, which may persist through life.
| |
| | |
| The comparison of the tubuli, formed out of these structures,
| |
| with the Elasmobranch and Amphibian testicular network is
| |
| justified in that both originate as outgrowths from the primary
| |
| Malpighian bodies, and thence extend into the testis, and come
| |
| into connection with the true seminiferous stroma.
| |
| | |
| As in the lower types the semen is transported from the
| |
| testicular network to the Wolffian duct by parts of the glandular
| |
| tubes of the Wolffian body. In the case of Reptilia the anterior
| |
| two or three segmental tubes in the region of the testis probably
| |
| have this function. In the case of Mammalia the vasa efferentia,
| |
| i.e. the coni vasculosi, appear, according to the usually accepted
| |
| view, to be of this nature, though Banks and other investigators
| |
| believe that they are independently developed structures. Further
| |
| investigations on this point are required. In Birds a connection
| |
| between the Wolffian body and the testis appears to be established as in the other types. The Wolffian duct itself becomes,
| |
| in the males of all Amniota, the vas deferens and the convoluted
| |
| canal of the epididymis the latter structure (except the head)
| |
| being entirely derived from the Wolffian duct.
| |
| | |
| In the female the Wolffian duct atrophies more or less
| |
| completely.
| |
| | |
| In Snakes (Braun) the posterior part remains as a functionless canal,
| |
| commencing at the ovary, and opening into the cloaca. In the Gecko
| |
| (Braun) it remains as a small canal joining the ureter ; in Blindworms a
| |
| considerable part of the canal is left, and in Lacerta (Braun) only interrupted
| |
| portions.
| |
| | |
| In Mammalia the middle part of the duct, known as Gaertner's canal,
| |
| persists in the females of some monkeys, of the pig and of many ruminants.
| |
| | |
| The Wolffian body atrophies nearly completely in both
| |
| sexes ; though, as described above, part of it opposite the testis
| |
| persists as the head of the epididymis. The posterior part of
| |
| the gland from the level of the testis may be called the sexual
| |
| part of the gland, the anterior part forming the non-sexual part.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 725
| |
| | |
| The latter, i.e. the anterior part, is first absorbed ; and in some
| |
| Reptilia the posterior part, extending from the region of the genital
| |
| glands to the permanent kidney, persists till into the second year.
| |
| | |
| Various remnants of the Wolffian body are found in the adults of both
| |
| sexes in different types. The most constant of them is perhaps the part in
| |
| the female equivalent to the head of the epididymis and to parts also of the
| |
| coiled tube of the epididymis, which may be called, with Waldeyer, the
| |
| epoophoron 1 . This is found in Reptiles, Birds and Mammals ; though in a
| |
| very rudimentary form in the first-named group. Remnants of the anterior
| |
| non-sexual part of the Wolffian bodies have been called by Waldeyer
| |
| parepididymis in the male, and paroophoron in the female. Such remnants
| |
| are not (Braun) found in Reptilia, but are stated to be found in both male
| |
| and female Birds, as a small organ consisting of blindly ending tubes with
| |
| yellow pigment. In some male Mammals (including Man) a parepididymis
| |
| is found on the upper side of the testis. It is usually known as the organ of
| |
| Giraldes.
| |
| | |
| The Mlillerian duct forms, as has been stated, the oviduct in
| |
| the female. The two ducts originally open independently into
| |
| the cloaca, but in the Mammalia a subsequent modification of
| |
| this arrangement occurs, which is dealt with in a separate
| |
| section. In Birds the right oviduct atrophies, a vestige being
| |
| sometimes left. In the male the Miillerian ducts atrophy more
| |
| or less completely.
| |
| | |
| In most Reptiles and in Birds the atrophy of the Miillerian ducts is
| |
| complete in the male, but in Lacerta and Anguis a rudiment of the anterior
| |
| part has been detected by Leydig as a convoluted canal. In the Rabbit
| |
| (Kolliker) 2 and probably other Mammals the whole of the ducts probably
| |
| disappears, but in some Mammals, e.g. Man, the lower fused ends of the
| |
| Miillerian ducts give rise to a pocket opening into the urethra, known as the
| |
| uterus masculinus ; and in other cases, e.g. the Beaver and the Ass, the
| |
| rudiments are more considerable, and may be continued into horns homologous with the horns of the uterus (Weber).
| |
| | |
| The hydatid of Morgani in the male is supposed (Waldeyer) to represent
| |
| the abdominal opening of the Fallopian tube in the female, and therefore to
| |
| be a remnant of the Miillerian duct.
| |
| | |
| Changes in the lower parts of the urinogenital ducts in the Amniota.
| |
| | |
| The genital cord. In the Monodelphia the lower part of
| |
| the Wolffian ducts becomes enveloped in both sexes in a special
| |
| | |
| 1 This is also called parovarium (His), and Rosenmiiller's organ.
| |
| | |
| 2 Weber (No. 553) states that a uterus masculinus is present in the Rabbit, but
| |
| his account is by no means satisfactory, and its presence is distinctly denied by
| |
| Kolliker.
| |
| | |
| | |
| | |
| 726
| |
| | |
| | |
| | |
| AMNIOTA.
| |
| | |
| | |
| | |
| cord of tissue, known as -the genital cord (fig. 407, gc), within the
| |
| lower part of which the MUllerian ducts are also enclosed. In
| |
| the male the MUllerian ducts in this cord atrophy, except at
| |
| their distal end where they unite to form the uterus masculinus.
| |
| The Wolffian ducts, after becoming the vasa deferentia, remain
| |
| for some time enclosed in the common cord, but afterwards
| |
| separate from each other. The seminal vesicles are outgrowths
| |
| of the vasa deferentia.
| |
| | |
| In the female the Wolffian ducts within the genital cord
| |
| atrophy, though rudiments of them are for a long time visible or
| |
| even permanently persistent. The lower parts of the MUllerian
| |
| ducts unite to form the vagina and body of the uterus. The
| |
| junction commences in the middle and extends forwards and
| |
| backwards ; the stage with a median junction being retained
| |
| permanently in Marsupials.
| |
| | |
| The urinogenital sinus and external generative organs.
| |
| In all the Amniota, there open at first into the common cloaca
| |
| the alimentary canal dorsally, the allantois ventrally, and the
| |
| Wolffian and MUllerian ducts and ureters laterally. In Reptilia
| |
| and Aves the embryonic condition is retained. In both groups
| |
| the allantois serves as an embryonic urinary bladder, but while
| |
| it atrophies in Aves, its stalk dilates to form a permanent
| |
| urinary bladder in Reptilia. In Mammalia the dorsal part of
| |
| the cloaca with the alimentary tract becomes first of all partially
| |
| constricted off from the ventral, which then forms a urinogenital
| |
| sinus (fig. 407, ug). In the course of development the urinogenital sinus becomes, in all Mammalia but the Ornithodelphia,
| |
| completely separated from the intestinal cloaca, and the two
| |
| parts obtain separate external openings. The ureters (fig. 407,
| |
| 3) open higher up than the other ducts into the stalk of the
| |
| allantois which dilates to form the bladder (4). The stalk
| |
| connecting the bladder with the ventral wall of the body constitutes the urachus, and loses its lumen before the close of
| |
| embryonic life. The part of the stalk of the allantois below the
| |
| openings of the ureters narrows to form the urethra, which opens
| |
| together with the Wolffian and MUllerian ducts into the urinogenital cloaca.
| |
| | |
| In front of the urinogenital cloaca there is formed a genital
| |
| prominence (fig. 407, cp), with a groove continued from the
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 727
| |
| | |
| urinogenital opening ; and on each side a genital fold (&). In
| |
| the male the sides of the groove on the prominence coalesce
| |
| together, embracing between them the opening of the urinogenital cloaca ; and the prominence itself gives rise to the penis,
| |
| | |
| | |
| | |
| | |
| FIG. 407. DIAGRAM OF THE URINOGENITAL ORGANS OF A MAMMAL AT AN
| |
| EARLY STAGE. (After Allen Thomson ; from Quain's Anatomy.)
| |
| | |
| The parts are seen chiefly in profile, but the Miillerian and Wolffian ducts are
| |
| seen from the front.
| |
| | |
| 3. ureter; 4. urinary bladder ; 5. urachus; of. genital ridge (ovary or testis) ; W.
| |
| left Wolffian body ; x. part at apex from which coni vasculosi are afterwards
| |
| developed ; w. Wolffian duct ; m. Miillerian duct ; gc. genital cord consisting of
| |
| Wolffian and Mullerian ducts bound up in a common sheath ; i. rectum ; ug. urinogenital sinus ; cp. elevation which becomes the clitoris or penis ; Is. ridge from
| |
| which the labia majora or scrotum are developed.
| |
| | |
| along which the common urinogenital passage is continued.
| |
| The two genital folds unite from behind forwards to form the
| |
| scrotum.
| |
| | |
| In the female the groove on the genital prominence gradually
| |
| disappears, and the prominence remains as the clitoris, which is
| |
| therefore the homologue of the penis : the two genital folds form
| |
| the labia majora. The urethra and vagina open independently
| |
| into the common urinogenital sinus.
| |
| | |
| | |
| | |
| 728 GENERAL CONCLUSIONS.
| |
| | |
| General conclusions and Summary.
| |
| | |
| Pronephros. Sedgwick has pointed out that the pronephros
| |
| is always present in types with a larval development, and either
| |
| absent or imperfectly developed in those types which undergo
| |
| the greater part of their development within the egg. Thus it
| |
| is practically absent in the embryos of Elasmobranchii and the
| |
| Amniota, but present in the larvae of all other forms.
| |
| | |
| This coincidence, on the principles already laid down in a
| |
| previous chapter on larval forms, affords a strong presumption
| |
| that the pronephros is an ancestral organ ; and, coupled with
| |
| the fact that it is the first part of the excretory system to be
| |
| developed, and often the sole excretory organ for a considerable
| |
| period, points to the conclusion that the pronephros and its duct
| |
| the segmental duct are the most primitive parts of the
| |
| Vertebrate excretory system. This conclusion coincides with
| |
| that arrived at by Gegenbaur and Fiirbringer.
| |
| | |
| The duct of the pronephros is always developed prior to the
| |
| gland, and there are two types according to which its development may take place. It may either be formed by the closing
| |
| in of a continuous groove of the somatic peritoneal epithelium
| |
| (Amphibia, Teleostei, Lepidosteus), or as a solid knob or rod of
| |
| cells derived from the somatic mesoblast, which grows backwards
| |
| between the epiblast and the mesoblast (Petromyzon, Elasmobranchii, and the Amniota).
| |
| | |
| It is quite certain that the second of these processes is not a
| |
| true record of the evolution of 'the duct, and though it is more
| |
| possible that the process observable in Amphibia and the
| |
| Teleostei may afford some indications of the manner in which
| |
| the duct was established, this cannot be regarded as by any
| |
| means certain.
| |
| | |
| The mode of development of the pronephros itself is apparently partly dependent on that of its duct. In Petromyzon,
| |
| where the duct does not at first communicate with the body
| |
| cavity, the pronephros is formed as a series of outgrowths from
| |
| the duct, which meet the peritoneal epithelium and open into
| |
| the body cavity ; but in other instances it is derived from the
| |
| anterior open end of the groove which gives rise to the segmental
| |
| duct. The open end of this groove may either remain single
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 729
| |
| | |
| (Teleostci, Ganoidei) or be divided into two, three or more
| |
| apertures (Amphibia). The main part of the gland in either
| |
| case is formed by convolutions of the tube connected with the
| |
| peritoneal funnel or funnels. The peritoneal funnels of the
| |
| pronephros appear to be segmentally arranged.
| |
| | |
| The pronephros is distinguished from the mesonephros by
| |
| developmental as well as structural features. The most important of the former is the fact that the glandular tubules of
| |
| which it is formed are always outgrowths of the segmental duct ;
| |
| while in the mesonephros they are always or almost always 1
| |
| formed independently of the duct.
| |
| | |
| The chief structural peculiarity of the pronephros is the
| |
| absence from it of Malpighian bodies with the same relations as
| |
| those in the meso- and metanephros; unless the structures found
| |
| in Myxine are to be regarded as such. Functionally the place
| |
| of such Malpighian bodies is taken by the vascular peritoneal
| |
| ridge spoken of in the previous pages as the glomerulus.
| |
| | |
| That this body is really related functionally to the pronephros appears to
| |
| be indicated (i) by its constant occurrence with the pronephros and its
| |
| position opposite the peritoneal openings of this body ; (2) by its atrophy at
| |
| the same time as the pronephros ; (3) by its enclosure together with the
| |
| pronephridian stoma in a special compartment of the body-cavity in
| |
| Teleostei and Ganoids, and its partial enclosure in such a compartment in
| |
| Amphibia.
| |
| | |
| The pronephros atrophies more or less completely in most
| |
| types, though it probably persists for life in the Teleostei and
| |
| Ganoids, and in some members of the former group it perhaps
| |
| forms the sole adult organ of excretion.
| |
| | |
| The cause of its atrophy may perhaps be related to the fact that it is
| |
| situated in the pericardial region of the body-cavity, the dorsal part of which
| |
| is aborted on the formation of a closed pericardium ; and its preservation in
| |
| Teleostei and Ganoids may on this view be due to the fact that in these types
| |
| its peritoneal funnel and its glomerulus are early isolated in a special cavity.
| |
| | |
| Mesonephros. The mesonephros is in all instances composed of a series of tubules (segmental tubes) which are
| |
| developed independently of the segmental duct. Each tubule is
| |
| | |
| 1 According t.o Sedgwick some of the anterior segmental tubes of Aves form an
| |
| exception to the general rule that there is no outgrowth from the segmental or
| |
| metanephric duct to meet the segmental tubes.
| |
| | |
| | |
| | |
| 730 GENERAL CONCLUSIONS.
| |
| | |
| typically formed of (i) a peritoneal funnel opening into (2) a
| |
| Malpighian body, from which there proceeds (3) a coiled glandular tube, finally opening by (4) a collecting tube into the
| |
| segmental duct, which constitutes the primitive duct for the
| |
| mesonephros as well as for the pronephros.
| |
| | |
| The development of the mesonephridian tubules is subject to
| |
| considerable variations.
| |
| | |
| (1) They may be formed as differentiations of the intermediate cell mass, and be from the first provided with a lumen,
| |
| opening into the body-cavity, and directly derived from the
| |
| section of the body-cavity present in the intermediate cell
| |
| mass; the peritoneal funnels often persisting for life (Elasmobranchii).
| |
| | |
| (2) They may be formed as solid cords either attached to
| |
| or independent of the peritoneal epithelium, which after first
| |
| becoming independent of the peritoneal epithelium subsequently
| |
| send downwards a process, which unites with it and forms a
| |
| peritoneal funnel, which may or may not persist (Acipenser,
| |
| Amphibia).
| |
| | |
| (3) They may be formed as in the last case, but acquire no
| |
| secondary connection with the peritoneal epithelium (Teleostei,
| |
| Amniota). In connection with the original attachment to the
| |
| peritoneal epithelium, a true peritoneal funnel may however be
| |
| developed (Aves, Lacertilia).
| |
| | |
| Physiological considerations appear to shew that of these
| |
| three methods of development the first is the most primitive.
| |
| The development of the tubes as solid cords can hardly be
| |
| primary.
| |
| | |
| A question which has to be answered in reference to the segmental tubes
| |
| is that of the homology of the secondarily developed peritoneal openings of
| |
| Amphibia, with the primary openings of the Elasmobranchii. It is on the
| |
| one hand difficult to understand why, if the openings are homologous in the
| |
| two types, the original peritoneal attachment should be obliterated in
| |
| Amphibia, only to be shortly afterwards reacquired. On the other hand
| |
| it is still more difficult to understand what physiological gain there could be,
| |
| on the assumption of the non-homology of the openings, in the replacement
| |
| of the primary opening by a secondary opening exactly similar to it.
| |
| Considering the great variations in development which occur in undoubtedly
| |
| homologous parts I incline to the view that the openings in the two types
| |
| are homologous.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 731
| |
| | |
| | |
| | |
| In the majority of the lower Vertebrata the mesonephric
| |
| tubes have at first a segmental arrangement, and this is no
| |
| doubt the primitive condition. The coexistence of two, three, or
| |
| more of them in a single segment in Amphibia, Aves and
| |
| Mammalia has recently been shewn, by an interesting discovery
| |
| of Eisig, to have a parallel amongst Chaetopods, in the coexistence of several segmental organs in a single segment in
| |
| some of the Capitellidae.
| |
| | |
| In connection with the segmental features of the mesonephros it is perhaps worth recalling the fact that in Elasmobranchii as well as other types there are traces of segmental
| |
| tubes in some of the postanal segments. In the case of all the
| |
| segmental tubes a Malpighian body becomes established close
| |
| to the extremity of the tube adjoining the peritoneal opening, or
| |
| in an homologous position in tubes without such an opening.
| |
| The opposite extremity of the tube always becomes attached to
| |
| the segmental duct.
| |
| | |
| In many of the segments of the mesonephros, especially in
| |
| the hinder ones, secondary and tertiary tubes become developed
| |
| in certain types, which join the collecting canals of the primary
| |
| tubes, and are provided, like the primary tubes, with Malpighian
| |
| bodies at their blind extremities.
| |
| | |
| There can it appears to me be little or no doubt that the
| |
| secondary tubes in the different types are homodynamous if not
| |
| homologous. Under these circumstances it is surprising to find
| |
| in what different ways they take their origin. In Elasmobranchii a bud sprouts out from the Malpighian body of one
| |
| segment, and joins the collecting tube of the preceding segment,
| |
| and subsequently, becoming detached from the Malpighian body
| |
| from which it sprouted, forms a fresh secondary Malpighian
| |
| body at its blind extremity. Thus the secondary tubes of one
| |
| segment are formed as buds from the segment behind. In
| |
| Amphibia (Salamandra) and Aves the secondary tubes develop
| |
| independently in the mesoblast. These great differences in
| |
| development are important in reference to the homology of
| |
| the metanephros or permanent kidney, which is discussed
| |
| below.
| |
| | |
| Before leaving the mesonephros it may be worth while putting forward
| |
| some hypothetical suggestions as to its origin and relation to the pro
| |
| | |
| | |
| 732 GENERAL CONCLUSIONS.
| |
| | |
| nephros, leaving however the difficult questions as to the homology of the
| |
| segmental tubes with the segmental organs of Chastopods for subsequent
| |
| discussion.
| |
| | |
| It is a peculiarity in the development of the segmental tubes that they at
| |
| first end blindly, though they subsequently grow till they meet the segmental
| |
| duct with which they unite directly, without the latter sending out any
| |
| offshoot to meet them 1 . It is difficult to believe that peritoneal infundibula
| |
| ending blindly and unprovided with some external orifice can have had an
| |
| excretory function, and we are therefore rather driven to suppose that the
| |
| peritoneal infundibula which become the segmental tubes were either from
| |
| the first provided each with an orifice opening to the exterior, or were united
| |
| with the segmental duct. If they were from the first provided with external
| |
| openings we may suppose that they became secondarily attached to the duct
| |
| of the pronephros (segmental duct), and then lost their external openings, no
| |
| trace of these structures being left, even in the ontogeny of the system.
| |
| It would appear to me more probable that the pronephros, with its duct
| |
| opening into the cloaca, was the only excretory organ of the unsegmented
| |
| ancestors of the Chordata, and that, on the elongation of the trunk and its
| |
| subsequent segmentation, a series of metameric segmental tubes became
| |
| evolved opening into the segmental duct, each tube being in a sort of way
| |
| serially homologous with the primitive pronephros. With the segmentation
| |
| of the trunk the latter structure itself may have acquired the more or less
| |
| definite metameric arrangement of its parts.
| |
| | |
| Another possible view is that the segmental tubes may be modified
| |
| derivatives of posterior lateral branches of the pronephros, which may at
| |
| first have extended for the whole length of the body-cavity. If there is any
| |
| truth in this hypothesis it is necessary to suppose that, when the unsegmented ancestor of the Chordata became segmented, the posterior
| |
| branches of the primitive excretory organ became segmentally arranged,
| |
| and that, in accordance with the change thus gradually introduced in them,
| |
| the time of their development became deferred, so as to accord to a certain
| |
| extent with the time of formation of the segments to which they belonged.
| |
| The change in their mode of development which would be thereby introduced is certainly not greater than that which has taken place in the case of
| |
| segmental tubes, which, having originally developed on the Elasmobranch
| |
| type, have come to develop as they do in the posterior part of the mesonephros of Salamandra, Birds, etc.
| |
| | |
| Genital ducts. So far the origin and development of the
| |
| excretory organs have been considered without reference to the
| |
| modifications introduced by the excretory passages coming to
| |
| serve as generative ducts. Such an unmodified state of the
| |
| | |
| 1 As mentioned in the note on p. 729 Sedgwick maintains that the anterior
| |
| segmental tubes of the Chick form an exception to this general statement.
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 733
| |
| | |
| | |
| | |
| excretory organs is perhaps found permanently in Cyclostomata 1 and transitorily in the embryos of most forms.
| |
| | |
| At first the generative products seem to have been discharged
| |
| freely into the body-cavity, and transported to the exterior by
| |
| the abdominal pores (vide p. 626).
| |
| | |
| The secondary relations of the excretory ducts to the
| |
| generative organs seem to have been introduced by an opening
| |
| connected with the pronephridian extremity of the segmental
| |
| duct having acquired the function of admitting the generative
| |
| products into it, and of carrying them outwards ; so that
| |
| primitively the segmental duct must have served as efferent duct
| |
| both for the generative products and the pronepJiric secretion (just
| |
| as the Wolffian duct still does for the testicular products and
| |
| secretion of the Wolffian body in Elasmobranchii and Amphibia).
| |
| | |
| The opening by which the generative products entered the
| |
| segmental duct can hardly have been specially developed for
| |
| this purpose, but must almost certainly have been one of the
| |
| peritoneal openings of the pronephros. As a consequence (by a
| |
| process of natural selection) of the segmental duct having both a
| |
| generative and a urinary function, a further differentiation took
| |
| place, by which that duct became split into two a ventral
| |
| Mullerian duct and a dorsal Wolffian duct.
| |
| | |
| The Mullerian duct was probably continuous with one or
| |
| more of the abdominal openings of the pronephros which served
| |
| as generative pores. At first the segmental duct was probably
| |
| split longitudinally into two equal portions, and this mode
| |
| of splitting is exceptionally retained in some Elasmobranchii ;
| |
| but the generative function of the Mullerian duct gradually
| |
| impressed itself more and more upon the embryonic development, so that, in the course of time, the Mullerian duct
| |
| developed less and less at the expense of the Wolffian duct.
| |
| This process appears partly to have taken place in Elasmobranchii, and still more in Amphibia, the Amphibia offering in
| |
| this respect a less primitive condition than the Elasmobranchii ;
| |
| while in Aves it has been carried even further, and it seems
| |
| possible that in some Amniota the Mullerian and segmental
| |
| | |
| 1 It is by no means certain that the transportation outwards of the genital products
| |
| by the abdominal pores in the Cyclostomata may not be the result of degeneration.
| |
| | |
| | |
| | |
| 734 GENERAL CONCLUSIONS.
| |
| | |
| ducts may actually develop independently, as they do exceptionally in individual specimens of Salamandra (Fiirbringer). The
| |
| abdominal opening no doubt also became specialised. At first it
| |
| is quite possible that more than one pronephric abdominal
| |
| funnel may have served for the entrance of the generative
| |
| products ; this function being, no doubt, eventually restricted to
| |
| one of them.
| |
| | |
| Three different types of development of the abdominal
| |
| opening of the Mullerian duct have been observed.
| |
| | |
| In Amphibia (Salamandra) the permanent opening of the
| |
| Mullerian duct is formed independently, some way behind the
| |
| pronephros.
| |
| | |
| In Elasmobranchii the original opening of the segmental
| |
| duct forms the permanent opening of the Mullerian duct, and no
| |
| true pronephros appears to be formed.
| |
| | |
| In Birds the anterior of the three openings of the rudimentary
| |
| pronephros remains as the permanent opening of the Mullerian
| |
| duct.
| |
| | |
| These three modes of development very probably represent
| |
| specialisations of the primitive state along three different lines.
| |
| In Amphibia the specialisation of the opening appears to have
| |
| gone so far that it no longer has any relation to the pronephros.
| |
| It was probably originally one of the posterior openings of this
| |
| gland.
| |
| | |
| In Elasmobranchii, on the other hand, the functional opening
| |
| is formed at a period when we should expect the pronephros to
| |
| develop. This state is very possibly the result of a differentiation by which the pronephros gradually ceased to become
| |
| developed, but one of its peritoneal openings remained as the
| |
| abdominal aperture of the Mullerian duct. Aves, finally, appear
| |
| to have become differentiated along a third line ; since in their
| |
| ancestors the anterior (?) pore of the head-kidney appears to
| |
| have become specialised as the permanent opening of the
| |
| Mullerian duct.
| |
| | |
| The Mullerian duct is usually formed in a more or less complete manner in both sexes. In Ganoids, where the separation
| |
| between it and the Wolffian duct is not completed to the cloaca,
| |
| and in the Dipnoi, it probably serves to carry off the generative
| |
| products of both sexes. In other cases however only the female
| |
| | |
| | |
| | |
| EXCRETORY ORGANS.
| |
| | |
| | |
| | |
| 735
| |
| | |
| | |
| | |
| products pass out by it, and the partial or complete formation
| |
| of the Mullerian duct in the male in these cases needs to be
| |
| explained. This may be done either by supposing the Ganoid
| |
| arrangement to have been the primitive one in the ancestors of
| |
| the other forms, or, by supposing characters acquired primitively
| |
| by the female to have become inherited by both sexes.
| |
| | |
| It is a question whether the nature of the generative ducts of
| |
| Teleostei can be explained by comparison with those of Ganoids.
| |
| The fact that the Mullerian ducts of the Teleostean Ganoid
| |
| Lepidosteus attach themselves to the generative organs, and thus
| |
| acquire a resemblance to the generative ducts of Teleostei,
| |
| affords a powerful argument in favour of the view that the
| |
| generative ducts of both sexes in the Teleostei are modified
| |
| Mullerian ducts. Embryology can however alone definitely
| |
| settle this question.
| |
| | |
| In the Elasmobranchii, Amphibia, and Amniota the male
| |
| products are carried off by the Wolffian duct, and they are
| |
| transported to this duct, not by open peritoneal funnels of the
| |
| mesonephros, but by a network of ducts which sprout either
| |
| from a certain number of the Malpighian bodies opposite the
| |
| testis (Amphibia, Amniota), or from the stalks connecting the
| |
| Malpighian bodies with the open funnels (Elasmobranchii).
| |
| After traversing this network the semen passes (except in
| |
| certain Anura) through a variable number of the segmental
| |
| tubes directly to the Wolffian duct. The extent of the connection of the testis with the Wolffian body is subject to great
| |
| variations, but it is usually more or less in the anterior region.
| |
| Rudiments of the testicular network have in many cases become
| |
| inherited by the female.
| |
| | |
| The origin of the connection between the testis and Wolffian body is still
| |
| very obscure. It would be easy to understand how the testicular products,
| |
| after falling into the body-cavity, might be taken up by the open extremities
| |
| of some of the peritoneal funnels, and how such open funnels might have
| |
| groove-like prolongations along the mesorchium, which might eventually be
| |
| converted into ducts. Ontogeny does not however altogether favour this
| |
| view of the origin of the testicular network. It seems to me nevertheless the
| |
| most probable view which has yet been put forward.
| |
| | |
| The mode of transportation of the semen by means of the mesonephric
| |
| tubules is so peculiar as to render it highly improbable that it was twice
| |
| acquired, it becomes therefore necessary to suppose that the Amphibia and
| |
| | |
| | |
| | |
| 736 GENERAL CONCLUSIONS.
| |
| | |
| Amniota inherited this mode of transportation of the semen from the same
| |
| ancestors as the Elasmobranchii. It is remarkable therefore that in the
| |
| Ganoidei and Dipnoi this arrangement is not found.
| |
| | |
| Either (i) the arrangement (found in the Ganoidei and Dipnoi) of the
| |
| Miillerian duct serving for both sexes is the primitive arrangement, and the
| |
| Elasmobranch is secondary, or (2) the Ganoid arrangement is a secondary
| |
| condition, which has originated at a stage in the evolution of the Vertebrata
| |
| when some of the segmental tubes had begun to serve as the efferent ducts
| |
| of the testis, and has resulted in consequence of a degeneration of the latter
| |
| structures. Although the second alternative is the more easy to reconcile
| |
| with the affinities of the Ganoid and Elasmobranch types, as indicated by
| |
| the other features of their organization, I am still inclined to accept the
| |
| former ; and consider that the incomplete splitting of the segmental duct in
| |
| Ganoidei is a strong argument in favour of this view.
| |
| | |
| Metanephros. With the employment of the Wolffian duct
| |
| to transport the semen there seems to be correlated (i) a
| |
| tendency of the posterior segmental tubes to have a duct of
| |
| their own, in which the seminal and urinary fluids cannot become
| |
| mixed, and (2) a tendency on the part of the anterior segmental
| |
| tubes to lose their excretory function. The posterior segmental
| |
| tubes, when connected in this way with a more or less specialised
| |
| duct, have been regarded in the preceding pages as constituting
| |
| a metanephros.
| |
| | |
| This differentiation is hardly marked in the Anura, but is
| |
| well developed in the Urodela and in the Elasmobranchii ; and
| |
| in the latter group has become inherited by both sexes. In the
| |
| Amniota it culminates, according to the view independently
| |
| arrived at by Semper and myself, (i) in the formation of a
| |
| completely distinct metanephros in both sexes, formed however,
| |
| as shewn by Sedgwick, from the same blastema as the Wolffian
| |
| body, and (2) in the atrophy in the adult of the whole Wolffian
| |
| body, except the part uniting the testis and the Wolffian duct.
| |
| | |
| The homology between the posterior metanephridian section of the
| |
| Wolffian body, in Elasmobranchii and Urodela, and the kidney of the
| |
| Amniota, is only in my opinion a general one, i.e. in both cases a common
| |
| cause, viz. the Wolffian duct acting as vas deferens, has resulted in a more
| |
| or less similar differentiation of parts.
| |
| | |
| Fiirbringer has urged against Semper's and my view that no satisfactory proof of it has yet been offered. This proof has however, since
| |
| Fiirbringer wrote his paper, been supplied by Sedgwick's observations.
| |
| The development of the kidney in the Amniota is no doubt a direct as
| |
| opposed to a phylogenetic development ; and the substitution of a direct for
| |
| | |
| | |
| | |
| EXCRETORY ORGANS. 737
| |
| | |
| | |
| | |
| a phylogenetic development has most probably been rendered possible by
| |
| the fact that the anterior part of the mesonephros continued all the while
| |
| to be unaffected and to remain as the main excretory organ during foetal
| |
| life.
| |
| | |
| The most serious difficulty urged by Fiirbringer against the homology is
| |
| the fact that the ureter of the metanephros develops on a type of its own,
| |
| which is quite distinct from the mode of development of the ureters of the
| |
| metanephros of the Ichthyopsidan forms. It is however quite possible, though
| |
| far from certain, that the ureter of Amniota may be a special formation
| |
| confined to that group, and this fact would in no wise militate against the
| |
| homology I have been attempting to establish.
| |
| | |
| Comparison of the Excretory organs of the Chordata and
| |
| Invertebrata.
| |
| | |
| The structural characters and development of the various forms of
| |
| excretory organs described in the preceding pages do not appear to me to
| |
| be sufficiently distinctive to render it possible to establish homologies
| |
| between these organs on a satisfactory basis, except in closely related
| |
| groups.
| |
| | |
| The excretory organs of the Platyelminthes are in many respects similar
| |
| to the provisional excretory organ of the trochosphere of Polygordius
| |
| and the Gephyrea on the one hand, and to the Vertebrate pronephros
| |
| on the other ; and the Platyelminth excretory organ with an anterior
| |
| opening might be regarded as having given origin to the trochosphere organ,
| |
| while that with a posterior opening may have done so for the Vertebrate
| |
| pronephros 1 .
| |
| | |
| Hatschek has compared the provisional trochosphere excretory organ of
| |
| Polygordius to the Vertebrate pronephros, and the posterior Chastopod
| |
| segmental tubes to the mesonephric tubes ; the latter homology having
| |
| been already suggested independently by both Semper and myself. With
| |
| reference to the comparison of the pronephros with the provisional excretory
| |
| organ of Polygordius there are two serious difficulties :
| |
| | |
| (1) The pronephric (segmental) duct opens directly into the cloaca,
| |
| while the duct of the provisional trochosphere excretory organ opens anteriorly, and directly to the exterior.
| |
| | |
| (2) The pronephros is situated within the segmented region of the
| |
| trunk, and has a more or less distinct metameric arrangement of its parts ;
| |
| while the provisional trochosphere organ is placed in front of the segmented
| |
| region of the trunk, and is in no way segmented.
| |
| | |
| The comparison of the mesonephric tubules with the segmental excretory organs of the Chaetopoda, though not impossible, cannot be satisfactorily admitted till some light has been thrown upon the loss of the supposed
| |
| external openings of the tubes, and the origin of their secondary connection
| |
| with the segmental duct.
| |
| | |
| 1 This suggestion has I believe been made by Fiirbringer.
| |
| B. III. 47
| |
| | |
| | |
| | |
| 738 BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| Confining our attention to the Invertebrata it appears to me fairly clear
| |
| that Hatschek is justified in holding the provisional trochosphere excretory
| |
| organs of Polygordius, Echiurus and the Mollusca to be homologous. The
| |
| atrophy of all these larval organs may perhaps be due to the presence of a
| |
| well-developed trunk region in the adult (absent in the larva), in which
| |
| excretory organs, probably serially homologous with those present in the
| |
| anterior part of the larva, became developed. The excretory organs in the
| |
| trunk were probably more conveniently situated than those in the head,
| |
| and the atrophy of the latter in the adult state was therefore brought about,
| |
| while the trunk organs became sufficiently enlarged to serve as the sole
| |
| excretory organs.
| |
| | |
| BIBLIOGRAPHY OF THE EXCRETORY ORGANS.
| |
| Invertebrata.
| |
| | |
| (512) H. Eisig. " Die Segmentalorgane d. Capitelliden." Mitth. a. d. zool.
| |
| Stat. z. Neapel, Vol. I. 1879.
| |
| | |
| (513) J. Fraipont. " Recherches s. 1'appareil excreteur des Trematodes et d.
| |
| Cesto'ides." Archives de Biologic, Vol. I. 1880.
| |
| | |
| (514) B. Hatschek. "Studien lib. Entwick. d. Anneliden." Arbeit, a. d.
| |
| zool. Instit. Wien, Vol. I. 1878.
| |
| | |
| (515) B. Hatschek. "Ueber Entwick. von Echiurus," etc. Arbeit, a. d.
| |
| zool. Instit. Wien, Vol. in. 1880.
| |
| | |
| EXCRETORY ORGANS OF VERTEBRATA.
| |
| General.
| |
| | |
| (516) F. M. Balfour. "On the origin and history of the urinogenital organs of
| |
| Vertebrates." yournal of Anat. and Phys., Vol. X. 1876.
| |
| | |
| (517) Max. Furbringer 1 . "Zur vergleichenden Anat. u. Entwick. d. Excretionsorgane d. Vertebraten." Morphol. Jahrbuch, Vol. IV. 1878.
| |
| | |
| (518) H. Meek el. Zur Morphol. d. Hani- u. Geschlechtnverkz.d. Wirbelthiere,
| |
| etc. Halle, 1848.
| |
| | |
| (519) Joh. Miiller. Bildungsgeschichte d. Genitalien, etc. Diisseldorf, 1830.
| |
| | |
| (520) H. Rathke. " Beobachtungen u. Betrachtungen u. d. Entwicklung d.
| |
| Geschlechtswerkzeuge bei den Wirbelthieren." N. Schriften d. naturf. Gesell. in
| |
| Dantzig, Bd. I. 1825.
| |
| | |
| (521) C. Semper 1 . "Das Urogenitalsystem d. Plagiostomen u. seine Bedeutung f. d. iibrigen Wirbelthiere." Arb. a. d. zool.-zoot. Instit. Wurzburg, Vol. II.
| |
| 1875
| |
| (522) W. Waldeyer 1 . Eierstock u. Ei. Leipzig, 1870.
| |
| | |
| | |
| | |
| 1 The papers of Furbringer, Semper and Waldeyer contain full references to the
| |
| literature of the Vertebrate excretory organs.
| |
| | |
| | |
| | |
| BIBLIOGRAPHY. 739
| |
| | |
| | |
| | |
| ElasmobrancJdi.
| |
| | |
| (523) A. Schultz. "Zur Entwick. d. Selachiereies." Archiv f. mikr. Anat.,
| |
| Vol. XI. 1875.
| |
| | |
| Vide also Semper (No. 521) and Balfour (No. 292).
| |
| | |
| Cyclostomata.
| |
| | |
| (524) J. Miiller. " Untersuchungen ii. d. Eingeweide d. Fische." Abh. d. k.
| |
| Ak. Wiss. Berlin, 1845.
| |
| | |
| (525) W. Miiller. "Ueber d. Persistenz d. Urniere b. Myxine glutinosa."
| |
| Jenaische Zeitschrift, Vol. VII. 1873.
| |
| | |
| (526) W. Miiller. "Ueber d. Urogenitalsystem d. Amphioxus u. d. Cyclostomen." Jenaische Zeitschri/t, Vol. IX. 1875.
| |
| | |
| (527) A. Schneider. Beitrdge z. vergleich. Anat. u. Entwick. d. Wirbelthiere.
| |
| Berlin, 1879.
| |
| | |
| (528) W. B. Scott. "Beitrage z. Entwick. d. Petromyzonten." Morphol.
| |
| Jahrbuch, Vol. vn. 1881.
| |
| | |
| Teleostei.
| |
| | |
| (529) J. Hyrtl. "Das uropoetische System d. Knochenfische." Denkschr. d.
| |
| k. k. Akad. Wiss. Wien, Vol. n. 1850.
| |
| | |
| (530) A. Rosenberg. Untersuchungen iib. die Entivicklung d. Teleostierniere.
| |
| Dorpat, 1867.
| |
| | |
| Vide also Oellacher (No. 72).
| |
| | |
| Amphibia.
| |
| | |
| (531) F. H. Bidder. Vergleichend-anatomische u. histologische Untersitchungen
| |
| ii. die mdnnlichen Geschleehts- und Harnwerkzeuge d. nackten Amphibien. Dorpat,
| |
| 1846.
| |
| | |
| (532) C. L. Duvernoy. "Fragments s. les Organes genito-urinaires des
| |
| Reptiles," etc. Mem. Acad. Sciences. Paris. Vol. xi. 1851, pp. 17 95.
| |
| | |
| (533) M. Fiirbringer. Zur Entwicklung d. Amphibienniere. Heidelberg, 1877.
| |
| | |
| (534) F. Leydig. Anatomie d. Amphibien u. Reptilien. Berlin, 1853.
| |
| | |
| (535) F. Leydig. Lehrbuch d. Hisiologie. Hamm, 1857.
| |
| | |
| (536) F. Meyer. "Anat. d. Urogenitalsystems d. Selachier u. Amphibien."
| |
| Sitz. d. naturfor. Gesellsch. Leipzig, 1875.
| |
| | |
| (537) J. W. Spengel. "Das Urogenitalsystem d. Amphibien." Arb. a. d.
| |
| zool.- zoot. Instil. Wiirzburg. Vol. III. 1876.
| |
| | |
| (538) VonWittich. "Harn- u. Geschlechtswerkzeuge d. Amphibien." Zeit.
| |
| f. wiss. Zool., Vol. IV.
| |
| | |
| Vide also Gotte (No. 296).
| |
| | |
| Amniota.
| |
| | |
| (539) F. M. Balfour and A. Sedgwick. "On the existence of a head -kidney
| |
| in the embryo Chick," etc. Quart. J. of Micr. Science, Vol. xix. 1878.
| |
| | |
| (540 ) Banks. On the Wolffian bodies of the fatus and their remains in the adult.
| |
| Edinburgh, 1864.
| |
| | |
| 472
| |
| | |
| | |
| | |
| 74O BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| (541) Th. Bornhaupt. Untersuchungen iib. die Entwicklung d. Urogenitalsystems beim Hiihnchen. Inaug. Diss. Riga, 1867.
| |
| | |
| (542) Max Braun. "Das Urogenitalsystem d. einheimischen Reptilien."
| |
| Arbeiten a. d. zool.-zoot. Instit. Wiirzburg. Vol. iv. 1877.
| |
| | |
| (543) J. Dansky u. J. Kostenitsch. "Ueb. d. Entwick. d. Keimblatter u. d.
| |
| WolfFschen Ganges im Hiihnerei." Mini. Acad. Imp. Petersbourg, vn. Series, Vol.
| |
| xxvil. 1880.
| |
| | |
| (544) Th. Egli. Beitrage zur Anat. und Entwick. d. Geschlechtsorgane. Inaug.
| |
| Diss. Zurich, 1876.
| |
| | |
| (545) E. Gasser. Beitrage zur Entwicklungsgeschichte d. Allantois, der
| |
| Milllcr'schen Gange u. des Afters. Frankfurt, 1874.
| |
| | |
| (546) E. Gasser. "Beob. iib. d. Entstehung d. Wolff schen Ganges bei Embryonen von Hiihnern u. Gansen." Arch, fiir mikr. Anat., Vol. xiv. 1877.
| |
| | |
| (547) E. Gasser. "Beitrage z. Entwicklung d. Urogenitalsystems d. Hiihnerembryonen." Sitz. d. GeseU. zur Befdrderung d. gesam. Naturwiss. Marburg, 1879.
| |
| | |
| (548) C. Kupffer. " Untersuchting iiber die Entwicklung des Harn- und Geschlechtssystems." Archiv fiir mikr. Anat., Vol. II. 1866.
| |
| | |
| (549) A. Sedgwick. "Development of the kidney in its relation to the
| |
| Wolffian body in the Chick." Quart. J. of Micros. Science, Vol. xx. 1880.
| |
| | |
| (550) A. Sedgwick. "On the development of the structure known as the
| |
| glomerulus of the head-kidney in the Chick." Quart. J. of Micros. Science, Vol. xx.
| |
| 1880.
| |
| | |
| (551) A. Sedgwick. "Early development of the Wolffian duct and anterior
| |
| Wolffian tubules in the Chick ; with some remarks on the vertebrate excretory
| |
| system." Quart. J. of Micros. Science, Vol. xxi. 1881.
| |
| | |
| (552) M. Watson. "The homology of the sexual organs, illustrated by comparative anatomy and pathology." Journal of Anat. and Phys., Vol. xiv. 1879.
| |
| | |
| (553) E. H. Weber. Zusdtze z. Lehre von Baue u. d. Verrichtungen d. Geschlechtsorgane. Leipzig, 1846.
| |
| | |
| Vide also Remak (No. 302), Foster and Balfour (No. 295), His (No. 297),
| |
| Kolliker (No. 298).
| |
| | |
| | |
| | |
| CHAPTER XXIV.
| |
| GENERATIVE ORGANS AND GENITAL DUCTS.
| |
| | |
| GENERATIVE ORGANS.
| |
| | |
| THE structure and growth of the ovum and spermatozoon
| |
| were given in the first chapter of this work, but their derivation
| |
| from the germinal layers was not touched on, and it is this
| |
| subject with which we are here concerned. If there are any
| |
| structures whose identity throughout the Metazoa is not open
| |
| to doubt these structures are the ovum and spermatozoon ;
| |
| and the constancy of their relations to the germinal layers
| |
| would seem to be a crucial test as to whether the latter have
| |
| the morphological importance usually attributed to them.
| |
| | |
| The very fragmentary state of our knowledge of the origin of
| |
| the generative cells has however prevented this test being so far
| |
| very generally applied.
| |
| | |
| Porifera. In the Porifera the researches of Schulze have
| |
| clearly demonstrated that both the ova and the spermatozoa
| |
| take their origin from indifferent cells of the general parenchyma, which may be called mesoblastic. The primitive germinal cells of the two sexes are not distinguishable ; but a
| |
| germinal cell by enlarging and becoming spherical gives rise
| |
| to an ovum ; and by subdivision forms a sperm-morula, from
| |
| the constituent cells of which the spermatozoa are directly
| |
| developed.
| |
| | |
| Ccelenterata. The greatest confusion prevails as to the
| |
| germinal layer from which the male and female products are
| |
| derived in the Ccelenterata 1 .
| |
| | |
| 1 E. van Beneden (No. 556) was the first to discover a different origin for the
| |
| generative products of the two sexes in Hydractinia, and his observations have led to
| |
| numerous subsequent researches on the subject. For a summary of the observations
| |
| on the Hydroids vide Weismann (No. 560).
| |
| | |
| | |
| | |
| 742 CCELENTERATA.
| |
| | |
| | |
| | |
| The following apparent modes of origin of these products
| |
| have been observed.
| |
| | |
| (1) The generative products of both sexes originate in the
| |
| ectoderm (epiblast) : Hydra, Cordylophora, Tubularia, all (?) free
| |
| Gonophores of Hydromedusae, the Siphonophora, and probably
| |
| the Ctenophora.
| |
| | |
| (2) The generative products of both sexes originate in the
| |
| entoderm (hypoblast) : Plumularia and Sertularella, amongst
| |
| the Hydroids, and the. whole of the Acraspeda and Actinozoa.
| |
| | |
| (3) The male cells are formed in the ectoderm, and the
| |
| female in the entoderm : Gonothyraea, Campanularia, Hydractinia, Clava.
| |
| | |
| In view of the somewhat surprising results to which the
| |
| researches on the origin of the genital products amongst the
| |
| Ccelenterata have led, it would seem to be necessary either to
| |
| hold that there is no definite homology between the germinal
| |
| layers in the different forms of Ccelenterata, or to offer some
| |
| satisfactory explanation of the behaviour of the genital products, which would not involve the acceptance of the first
| |
| alternative.
| |
| | |
| Though it can hardly be said that such an explanation has
| |
| yet been offered, some observations of Kleinenberg (No. 557)
| |
| undoubtedly point to such an explanation being possible.
| |
| | |
| Kleinenberg has shewn that in Eudendrium the ova migrate
| |
| freely from the ectoderm into the endoderm, and vice versa ; but
| |
| he has given strong grounds for thinking that they originate in
| |
| the ectoderm. He has further shewn that the migration in this
| |
| type is by no means an isolated phenomenon.
| |
| | |
| Since it is usually only possible to recognise generative
| |
| elements after they have advanced considerably in development,
| |
| the mere position of a generative cell, when first observed, can
| |
| afford, after what Kleinenberg has shewn, no absolute proof
| |
| of its origin. Thus it is quite possible that there is really
| |
| only one type of origin for the generative cells in the Ccelenterata.
| |
| | |
| Kleinenberg has given reasons for thinking that the migration of the ova
| |
| into the entoderm may have a nutritive object. If this be so, and there are
| |
| numerous facts which shew that the position of generative cells is often
| |
| largely influenced by their nutritive requirements, it seems not impossible
| |
| | |
| | |
| | |
| GENERATIVE ORGANS. 743
| |
| | |
| that the endodermal position of the generative organs in the Actinozoa and
| |
| acraspedote Medusre may have arisen by a continuously earlier migration of
| |
| the generative cells from the ectoderm into the endoderm ; and that the
| |
| migration may now take place at so early a period of the development, that
| |
| we should be justified in formally holding the generative products to be
| |
| endodermal in origin.
| |
| | |
| \Ve might perhaps, on this view, formulate the origin of the generative
| |
| products in the Ccelenterata in the following way :
| |
| | |
| Both ova and spermatozoa primitively originated in the ectoderm, but in
| |
| order to secure a more complete nutrition the cells which give rise to them
| |
| exhibit in certain groups a tendency to migrate into the endoderm. This
| |
| migration, which may concern the generative cells of one or of both the
| |
| sexes, takes place in some cases after the generative cells have become
| |
| recognisable as such, and very probably in other cases at so early a period
| |
| that it is impossible to distinguish the generative cells from indifferent
| |
| embryonic cells.
| |
| | |
| Very little is known with reference to the origin of the
| |
| generative cells in the triploblastic Invertebrata.
| |
| | |
| Chaetopoda and Gephyrea. In the Chaetopoda and
| |
| Gephyrea, the germinal cells are always developed in the adult
| |
| from the epithelial lining of the body cavity ; so that their origin
| |
| from the mesoblast seems fairly established.
| |
| | |
| If we are justified in holding the body cavity of these forms
| |
| to be a derivative of the primitive archenteron (vide pp. 356 and
| |
| 357) the generative cells may fairly be held to originate from a
| |
| layer which corresponds to the endoderm of the Ccelenterata 1 .
| |
| | |
| Chaetognatha. In Sagitta the history of the generative
| |
| cells, which was first worked out by Kowalevsky and Biitschli,
| |
| has been recently treated with great detail by O. Hertwig 2 .
| |
| | |
| The generative cells appear during the gastrula stage, as two
| |
| large cells with conspicuous nuclei, which are placed in the
| |
| hypoblast lining the archenteron, at the pole opposite the
| |
| blastopore. These cells soon divide, and at the same time pass
| |
| out of the hypoblast, and enter the archenteric cavity (fig. 408
| |
| - A, ge). The division into four cells, which is not satisfactorily
| |
| represented ifl my diagram, takes place in such a way that two
| |
| | |
| 1 The Hertwigs (No. 271) state that in their opinion the generative cells arise
| |
| from the lining of the body cavity in all the forms whose body cavity is a product of
| |
| the archenteron. We do not know anything of the embryonic development of the
| |
| generative organs in the Echinodermata, but the adult position of the generative
| |
| organs in this group is very unfavourable to the Hertwigs' view.
| |
| | |
| 2 O. Hertwig, Die Chcetognathen. Jena, 1880
| |
| | |
| | |
| 744
| |
| | |
| | |
| | |
| CH^ETOGNATHA.
| |
| | |
| | |
| | |
| cells are placed nearer the median line, and two externally. The
| |
| two inner cells form the eventual testes, and the outer the
| |
| | |
| | |
| | |
| | |
| FIG. 408. THREK STAGES IN THE DEVELOPMENT OF SAGITTA. (A and C after
| |
| | |
| Biitschli, and B after Kowalevsky.)
| |
| The three embryos are represented in the same positions.
| |
| | |
| A. Represents the gastrula stage.
| |
| | |
| B. Represents a succeeding stage, in which the primitive archenteron is commencing to be divided into three.
| |
| | |
| C. Represents a later stage, in which the mouth involution (in) has become continuous with the alimentary tract, and the blastopore is closed.
| |
| | |
| ///. mouth ; al. alimentary canal ; ac. archenteron ; bl.p. blastopore ; pv. perivisceral cavity ; sp, splanchnic mesoblast ; so. somatic mesoblast ; ge. generative
| |
| organs.
| |
| | |
| ovaries, one half of each primitive cell thus forming an ovary, and
| |
| the other a testis.
| |
| | |
| | |
| | |
| | |
| FIG. 409. Two VIEWS OF A LATE EMBRYO OF SAGITTA. A, from the dorsal
| |
| | |
| surface. B, from the side. (After Biitschli.)
| |
| | |
| m. mouth ; al. alimentary canal ; v.g. ventral ganglion (thickening of epiblast) ;
| |
| <.'/. epiblast ; c.pv. cephalic section of body cavity ; so. somatopleure ; sp. splanchnopleure ; ge. generative organs.
| |
| | |
| | |
| | |
| GENERATIVE ORGANS.
| |
| | |
| | |
| | |
| 745
| |
| | |
| | |
| | |
| When the archenteric cavity is divided into a median
| |
| alimentary tract, and two lateral sections forming the body
| |
| cavity, the generative organs are placed in the common vestibule
| |
| into which both the body cavity and alimentary cavity at first
| |
| open (fig. 408).
| |
| | |
| The generative organs long retain their character as simple
| |
| cells. Eventually (fig. 409) the two ovaries travel forwards, and
| |
| apply themselves to the body walls, while the two testes also
| |
| become separated by a backward prolongation of the median
| |
| alimentary tract.
| |
| | |
| On the formation of the transverse septum dividing the tail
| |
| from the body, the ovarian cells lie immediately in front of this
| |
| septum, and the testicular cells in the region behind it.
| |
| | |
| Polyzoa. In Pedicellina amongst the entoproctous Polyzoa
| |
| Hatschek finds that the generative organs originate from a pair
| |
| of specially large mesoblast cells, situated in the space between
| |
| the stomach and the floor of the vestibule. The two cells
| |
| undergo changes, which have an obvious resemblance to those of
| |
| the generative cells of the Chsetognatha. They become surrounded by an investment of mesoblast cells, and divide so as to
| |
| form two masses. Each of these masses at a later period
| |
| separates into an anterior and a posterior part. The former
| |
| becomes the ovary, the latter the testis.
| |
| | |
| Nematoda. In the Nematoda the generative organs are
| |
| derived from the division of a single cell which would appear to
| |
| be mesoblastic 1 .
| |
| | |
| Insecta. The generative cells have been observed at a very
| |
| early embryonic stage in several insect forms (Vol. II. p. 404), but
| |
| the observations so far recorded with reference to them do not
| |
| enable us to determine with certainty from which of the germinal
| |
| layers they are derived.
| |
| | |
| Crustacea. In Moina, one of the Cladocera, Grobben 2 has
| |
| shewn that the generative organs are derived from a single cell,
| |
| which becomes differentiated during the segmentation. This
| |
| cell, which is in close contiguity with the cells from which both
| |
| the mesoblast and hypoblast originate, subsequently divides ;
| |
| | |
| 1 Fide Vol. n. p. 374; also Gotte, Zool. Anzeiger, No. 80, p. 189.
| |
| | |
| 2 C. Grobben. "Die Entwick. d. Moina rectirostris." Arbeit, a. d. zool. Instil.
| |
| Wien. Vol. II. 1879.
| |
| | |
| | |
| | |
| 746
| |
| | |
| | |
| | |
| CHORDATA.
| |
| | |
| | |
| | |
| sp.c
| |
| | |
| | |
| | |
| but at the gastrula stage, and after the mesoblast has become
| |
| formed, the cells it gives rise to are enclosed in the epiblast, and
| |
| do not migrate inwards till a later stage. The products of the
| |
| division of the generative cell subsequently divide into two
| |
| masses. It is not possible to assign the generative cell of Moina
| |
| to a definite germinal layer. Grobben, however, thinks that it
| |
| originates from the division of a cell, the remainder of which
| |
| gives rise to the hypoblast.
| |
| | |
| Chordata. In the Vertebrata, the primitive generative cells
| |
| (often known as primitive ova) are early distinguishable, being
| |
| imbedded amongst the cells of two linear streaks of peritoneal
| |
| epithelium, placed on the dorsal side of the body cavity, one on
| |
| each side of the mesentery (figs. 405
| |
| C and 4io,/0). They appear to be
| |
| derived from the epithelial cells
| |
| amongst which they lie ; and are
| |
| characterized by containing a large
| |
| granular nucleus, surrounded by a
| |
| considerable body of protoplasm.
| |
| The peritoneal epithelium in which
| |
| they are placed is known as the
| |
| germinal epithelium.
| |
| | |
| It is at first impossible to distinguish the germinal cells which will
| |
| become ova from those which will
| |
| become spermatozoa.
| |
| | |
| The former however remain within the peritoneal epithelium (fig. 41 1),
| |
| and become converted into ova in a
| |
| manner more particularly described
| |
| in Vol. II. pp. 54 59.
| |
| | |
| The history of the primitive
| |
| germinal cells in the male has not
| |
| been so adequately worked out as in
| |
| the female.
| |
| | |
| The fullest history of them is
| |
| that given by Semper (No. 559) for
| |
| the Elasmobranchii, the general accuracy of which I can fully support ;
| |
| | |
| | |
| | |
| | |
| FIG. 410. SECTION THROUGH
| |
| THE TRUNK OF A SCYLLIUM
| |
| EMBRYO SLIGHTLY YOUNGER
| |
| | |
| THAN 28 F.
| |
| | |
| sp.c. spinal cord ; W. white
| |
| matter of spinal cord ; pr. posterior nerve-roots ; ch. notochord ;
| |
| x. sub-notochordal rod ; ao. aorta ;
| |
| mp. muscle-plate ; mp'. inner layer
| |
| of- muscle-plate already converted
| |
| into muscles ; Vr. rudiment of
| |
| vertebral body ; st. segmental
| |
| tube; sd. segmental duct; sp.v.
| |
| spiral valve ; v. subintestinal vein ;
| |
| i>.o. primitive generative cells.
| |
| | |
| | |
| | |
| GENERATIVE ORGANS.
| |
| | |
| | |
| | |
| 747
| |
| | |
| | |
| | |
| though with reference to certain stages in the history further
| |
| researches are still required 1 .
| |
| | |
| In Elasmobranchii the male germinal cells, instead of remaining in the germinal epithelium, migrate into the adjacent stroma,
| |
| accompanied I believe by some of the indifferent epithelial cells.
| |
| Here they increase in number, and give rise to masses of variable
| |
| form, composed partly of true germinal cells, and partly of
| |
| smaller cells with deeply staining nuclei, which are, I believe,
| |
| derived from the germinal epithelium.
| |
| | |
| | |
| | |
| | |
| FIG. 411. TRANSVERSE SECTION THROUGH THE OVARY OF A YOUNG EMBRYO
| |
| OK SCYLLIUM CANICULA, TO SHEW THE PRIMITIVE GERMINAL CELLS (po) LYING
| |
| IN THE GERMINAL EPITHELIUM ON THE OUTER SIDE OF THE OVARIAN RIDGE.
| |
| | |
| These masses next break up into ampullae, mainly formed of
| |
| germinal cells, and each provided with a central lumen ; and
| |
| these ampullae attach themselves to tubes derived from the
| |
| smaller cells, which are in their turn continuous with the
| |
| testicular network. The spermatozoa are developed from the
| |
| cells forming the walls of the primitive ampulla;; but the
| |
| process of their formation does not concern us in this chapter.
| |
| | |
| In the Reptilia Braun has traced the passage of the primitive
| |
| germinal cells into the testicular tubes, and I am able to confirm
| |
| his observations on this point : he has not however traced their
| |
| further history.
| |
| | |
| 1 Balbiani (No. 554) has also recently dealt with this subject, but I cannot bring
| |
| my own observations into accord with his as to the structure of the Elasmobranch
| |
| testis.
| |
| | |
| | |
| | |
| MODE OF EXIT OF GENITAL PRODUCTS.
| |
| | |
| | |
| | |
| In Mammalia the evidence of the origin of the spermatospores from the germinal epithelium is not quite complete, but
| |
| there can be but little doubt of its occurrence 1 .
| |
| | |
| In Amphioxus Langerhans has shewn that the ova and
| |
| spermatozoa are derived from similar germinal cells, which may
| |
| be compared to the germinal epithelium of the Vertebrata.
| |
| These cells are however segmentally arranged as separate
| |
| masses (vide Vol. II. p. 54).
| |
| | |
| BIBLIOGRAPHY.
| |
| | |
| (554) G. Balbiani. Lemons s. la generation des Vcrlebrcs. Paris, 1879.
| |
| | |
| (555) F. M. Balfour. "On the structure and development of the Vertebrate
| |
| ovary." Quart, J. of Micr. Science, Vol. xvm.
| |
| | |
| (556) E. van Beneden. "De la distinction originelle dutecticule et clel'ovaire,
| |
| etc." Bull. Ac. roy. belgique, Vol. xxxvil. 1874.
| |
| | |
| (557) N. Kleinenberg. "Ueb. d. Entstehung d. Eier b. Eudendrium." Zcit.
| |
| f. -wiss. Zool., Vol. xxxv. 1881.
| |
| | |
| (558) H. Ludwig. "Ueb. d. Eibildung im Theirreiche." Arbeit, a. d. zool.zoot. Inslit. Wilrzburg, Vol. I. 1874.
| |
| | |
| (559) C. Semper. "Das Urogenilalsystem d. Plagiostomen, etc." Arbeit, a.
| |
| d. zooL-zoot. Ins tit. Witrzbiirg, Vol. II. 1875.
| |
| | |
| (560) A. Weismann. "Zur Frage nach dem Ursprung d. Geschlechtszellen bei
| |
| den Hydroiden." Zool. Anzeiger, No. 55, 1880.
| |
| | |
| Fitffcalso O. and R. Hertwig (No. 271), Kolliker (No. 298), etc.
| |
| | |
| GENITAL DUCTS.
| |
| | |
| The development and evolution of the generative ducts is as
| |
| yet very incompletely worked out, but even in the light of our
| |
| present knowledge a comparative review of this subject brings to
| |
| light features of considerable interest, and displays a fruitful
| |
| field for future research.
| |
| | |
| In the Ccelenterata there are no generative ducts.
| |
| | |
| In the Hydromedusae and Siphonophora the generative
| |
| products are liberated by being dehisced directly into the
| |
| surrounding medium ; while in the Acraspeda, the Actinozoa
| |
| and the Ctenophora, they are dehisced into parts of the gastrovascular system, and carried to the exterior through the mouth.
| |
| | |
| The arrangement in the latter forms indicates the origin of
| |
| | |
| 1 An entirely different view of the origin of the sperm cells has been adopted by
| |
| Balbiani, for which the reader is referred to his Memoir (No. 554).
| |
| | |
| | |
| | |
| GENITAL DUCTS.
| |
| | |
| | |
| | |
| 749
| |
| | |
| | |
| | |
| the methods of transportation of the genital products to the
| |
| exterior in many of the higher types.
| |
| | |
| It has been already pointed out that the body cavity in a
| |
| very large number of forms is probably derived from parts of a
| |
| gastrovascular system like that of the Actinozoa.
| |
| | |
| When the part of the gastrovascular system into which the
| |
| generative products were dehisced became, on giving rise to the
| |
| body cavity, shut off from the exterior, it would be essential that
| |
| some mode of transportation outwards of the generative products
| |
| should be constituted.
| |
| | |
| In some instances simple pores (probably already existing at
| |
| the time of the establishment of a closed body cavity) become
| |
| the generative ducts. Such seems probably to have been the
| |
| case in the Chaetognatha (Sagitta) and in the primitive
| |
| Chordata.
| |
| | |
| In the latter forms the generative products are sometimes dehisced into
| |
| the peritoneal cavity, and thence transported by the abdominal pores to the
| |
| exterior (Cyclostomata and some Teleostei, vide p. 626). In Amphioxus
| |
| they pass by dehiscence into the atrial cavity, and thence through the gill
| |
| slits and by the mouth, or by the abdominal pore (?) to the exterior. The
| |
| arrangement in Amphioxus and the Teleostei is probably secondary, as
| |
| possibly also is that in the Cyclostomata ; so that the primitive mode of
| |
| exit of the generative products in the Chordata is still uncertain. It is
| |
| highly improbable that the generative ducts of the Tunicata are primitive
| |
| structures.
| |
| | |
| A better established and more frequent mode of exit of the
| |
| generative products when dehisced into the body cavity is by
| |
| means of the excretory organs. The generative products pass
| |
| from the body cavity into the open peritoneal funnels of such
| |
| organs, and thence through their ducts to the exterior. This
| |
| mode of exit of the generative products is characteristic of the
| |
| Chaetopoda, the Gephyrea, the Brachiopoda and the Vertebrata,
| |
| and probably also of the Mollusca. It is moreover quite possible
| |
| that it occurs in the Polyzoa, some of the Arthropoda, the
| |
| Platyelminthes and some other types.
| |
| | |
| The simple segmental excretory organs of the Polychaeta,
| |
| the Gephyrea and the Brachiopoda serve as generative canals,
| |
| and in many instances they exhibit no modification, or but a
| |
| very slight one, in connection with their secondary generative
| |
| | |
| | |
| | |
| 750 DERIVATION FROM EXCRETORY ORGANS.
| |
| | |
| function ; while in other instances, e.g. Bonellia, such modification is very considerable.
| |
| | |
| The generative ducts of the Oligochaeta are probably derived from
| |
| excretory organs. In the Terricola ordinary excretory organs are present in
| |
| the generative segments in addition to the generative ducts, while in the
| |
| Limicola generative ducts alone are present in the adult, but before their
| |
| development excretory organs of the usual type are found, which undergo
| |
| atrophy on the appearance of the generative ducts (Vedjovsky).
| |
| | |
| From the analogy of the splitting of the segmental duct of the Vertebrata
| |
| into the Miillerian and Wolffian ducts, as a result of a combined generative
| |
| and excretory function (vide p. 728), it seems probable that in the generative segments of the Oligochasta the excretory organs had at first both an
| |
| excretory and a generative function, and that, as a secondary result of this
| |
| double function, each of them has become split into two parts, a generative
| |
| and an excretory. The generative part has undergone in all forms great
| |
| modifications. The excretory parts remain unmodified in the Earthworms
| |
| (Terricola), but completely abort on the development of the generative ducts
| |
| in the Limicola. An explanation may probably be given of the peculiar
| |
| arrangements of the generative ducts in Saccocirrus amongst the Polychaeta (vide Marion and Bobretzky), analogous to that just offered for the
| |
| Oligochaeta.
| |
| | |
| The very interesting modifications produced in the excretory
| |
| organs of the Vertebrata by their serving as generative ducts
| |
| were fully described in the last chapter ; and with reference to
| |
| this part of our subject it is only necessary to call attention to
| |
| the case of Lepidosteus and the Teleostei.
| |
| | |
| In Lepidosteus the Mullerian duct appears to have become
| |
| attached to the generative organs, so that the generative
| |
| products, instead of falling directly into the body cavity and
| |
| thence entering the open end of a peritoneal funnel of the
| |
| excretory organs, pass directly into the Mullerian duct without
| |
| entering the body cavity. In most Teleostei the modification is
| |
| more complete, in that the generative ducts in the adult have no
| |
| obvious connection with the excretory organs.
| |
| | |
| The transportation of the male products to the exterior in all
| |
| the higher Vertebrata, without passing into the body cavity, is
| |
| in principle similar to the arrangement in Lepidosteus.
| |
| | |
| The above instances of the peritoneal funnels of an excretory
| |
| organ becoming continuous with the generative glands, render it
| |
| highly probable that there may be similar instances amongst the
| |
| In vertebrata.
| |
| | |
| | |
| | |
| GENITAL DUCTS.
| |
| | |
| | |
| | |
| 751
| |
| | |
| | |
| | |
| As has been already pointed out by Gegenbaur there are
| |
| many features in the structure of the genital ducts in the more
| |
| primitive Mollusca, which point to their having been derived
| |
| from the excretory organs. In several Lamellibranchiata 1
| |
| (Spondylus, Lima, Pecten) the generative ducts open into the
| |
| excretory organs (organ of Bojanus), so that the generative
| |
| products have to pass through the excretory organ on their way
| |
| to the exterior. In other Lamellibranchiata the genital and
| |
| excretory organs open on a common papilla, and in the remaining types they are placed close together.
| |
| | |
| In the Cephalopoda again the peculiar relations of the
| |
| generative organs to their ducts point to the latter having
| |
| primitively had a different, probably an excretory, function.
| |
| The glands are not continuous with the ducts, but are placed in
| |
| special capsules from which the ducts proceed. The genital
| |
| products are dehisced into these capsules and thence pass into
| |
| the ducts.
| |
| | |
| In the Gasteropoda the genital gland is directly continuous
| |
| with its duct, and the latter, especially in the Pulmonata and
| |
| Opisthobranchiata, assumes such a complicated form that its
| |
| origin from the excretory organ would hardly have been
| |
| suspected. The fact however that its opening is placed near
| |
| that of the excretory organ points to its being homologous with
| |
| the generative ducts of the more primitive types.
| |
| | |
| In the Discophora, where the generative ducts are continuous
| |
| with the glands, the structure both of the generative glands and
| |
| ducts points to the latter having originated from excretory
| |
| organs.
| |
| | |
| It seems, as already mentioned, very possible that there are
| |
| other types in which the generative ducts are derived from the
| |
| excretory organs. In the Arthropoda for instance the generative
| |
| ducts, where provided with anteriorly placed openings, as in the
| |
| Crustacea, Arachnida and the Chilognathous Myriapoda, the
| |
| Pcecilopoda, etc., may possibly be of this nature, but the data
| |
| for deciding this point are so scanty that it is not at present
| |
| possible to do more than frame conjectures.
| |
| | |
| The ontogeny of the generative ducts of the Nematoda and
| |
| | |
| 1 For a summary of the facts on this subject vide Bronn, Klassen u. Ordnungen d.
| |
| Thierreichs, Vol. in. p. 404.
| |
| | |
| | |
| | |
| 752 DERIVATION FROM EXCRETORY ORGANS.
| |
| | |
| the Insecta appears to point to their having originated independently of the excretory organs.
| |
| | |
| In the Nematoda the generative organs of both sexes
| |
| originate from a single cell (Schneider, Vol. I. No. 390).
| |
| | |
| This cell elongates and its nuclei multiply. After assuming
| |
| a somewhat columnar form, it divides into (i) a superficial
| |
| investing layer, and (2) an axial portion.
| |
| | |
| In the female the superficial layer is only developed distinctly
| |
| in the median part of the column. In the course of the further
| |
| development the two ends of the column become the blind ends
| |
| of the ovary, and the axial tissue they contain forms the
| |
| germinal tissue of nucleated protoplasm. The superficial layer
| |
| gives rise to the epithelium of the uterus and oviduct. The
| |
| germinal tissue, which is originally continuous, is interrupted in
| |
| the middle part (where the superficial layer gives rise to the
| |
| uterus and oviduct), and is confined to the two blind extremities
| |
| of the tube.
| |
| | |
| In the male the superficial layer, which gives rise tc the
| |
| epithelium of the vas deferens, is only formed at the hinder ond
| |
| of the original column. In other respects the development takes
| |
| place as in the female.
| |
| | |
| In the Insecta again the evidence, though somewhat conflicting,
| |
| indicates that the generative ducts arise very much as in Nematodes, from the same primitive mass as the generative organs. In
| |
| both of these types it would seem probable that the generative
| |
| organs were primitively placed in the body cavity, and attached
| |
| to the epidermis, through a pore in which their products passed
| |
| out ; and that, acquiring a tubular form, the peripheral part of
| |
| the gland gave rise to a duct, the remainder constituting the true
| |
| generative gland. It is quite possible that the generative ducts
| |
| of such forms as the Platyelminthes may have had a similar
| |
| origin to those in Insecta and Nematoda, but from the analogy
| |
| of the Mollusca there is nearly as much to be said for regarding
| |
| them as modified excretory organs.
| |
| | |
| In the Echinodermata nothing is unfortunately known as to
| |
| the ontogeny of the generative organs and ducts. The structure
| |
| of these organs in the adult would however seem to indicate that
| |
| the most primitive type of echinoderm generative organ consists
| |
| of a blind sack, projecting into the body cavity, and opening by
| |
| | |
| | |
| | |
| GENITAL DUCTS. 753
| |
| | |
| | |
| | |
| a pore to the exterior. The sack is lined by an epithelium,
| |
| continuous with the epidermis, the cells of which give rise to the
| |
| ova or spermatozoa. The duct of these organs is obviously
| |
| hardly differentiated from the gland ; and the whole structure
| |
| might easily be derived from the type of generative organ
| |
| characteristic of the Hydromedusae, where the generative cells
| |
| are developed from special areas of the ectoderm, and, when ripe,
| |
| pass directly into the surrounding medium.
| |
| | |
| If this suggestion is correct we may suppose that the generative ducts of the Echinodermata have a different origin to those
| |
| of the majority of 1 the remaining triploblastica.
| |
| | |
| Their ducts have been evolved in forms in which the
| |
| generative products continued to be liberated directly to the
| |
| exterior, as in the Hydromedusae ; while those of other types
| |
| have been evolved in forms in which the generative products
| |
| were first transported, as in the Actinozoa, into the gastrovascular
| |
| canals 2 .
| |
| | |
| 1 It would be interesting to have further information about Balanoglossus.
| |
| | |
| 2 These views fit in very well with those already put forward in Chapter xm. on
| |
| the affinities of the Echinodermata.
| |
| | |
| | |
| | |
| B. III.
| |
| | |
| | |
| | |
| 48
| |
| | |
| | |
| | |
| CHAPTER XXV.
| |
| | |
| THE ALIMENTARY CANAL AND ITS APPENDAGES, IN
| |
| THE CHORDATA.
| |
| | |
| THE alimentary canal in the Chordata is always formed of
| |
| three sections, analogous to those so universally present in the
| |
| Invertebrata. These sections are (i) the mesenteron lined by
| |
| hypoblast ; (2) the stomodaeum or mouth lined by epiblast, and
| |
| (3) the proctodaeum or anal section lined like the stomodaeum by
| |
| epiblast.
| |
| | |
| Mesenteron.
| |
| | |
| The early development of the epithelial wall of the mesenteron
| |
| has already been described (Chapter XI.). It forms at first a
| |
| simple hypoblastic tube extending from near the front end of the
| |
| body, where it terminates blindly, to the hinder extremity where
| |
| it is united with the neural tube by the neurenteric canal (fig.
| |
| 420, ne). It often remains for a long time widely open in the
| |
| middle towards the yolk-sack.
| |
| | |
| It has already been shewn that from the dorsal wall of the
| |
| mesenteron the notochord is separated off nearly at the same
| |
| time as the lateral plates of mesoblast (pp. 292 300).
| |
| | |
| The subnotochordal rod. At a period slightly subsequent
| |
| to the formation of the notochord, and before any important
| |
| differentiations in the mesenteron have become apparent, a
| |
| remarkable rod-like body, which was first discovered by Gotte,
| |
| becomes split off from the dorsal wall of the alimentary tract in
| |
| all the Ichthyopsida. This body, which has a purely provisional
| |
| existence, is known as the subnotochordal rod.
| |
| | |
| | |
| | |
| MESENTERON.
| |
| | |
| | |
| | |
| 755
| |
| | |
| | |
| | |
| It develops in Elasmobranch embryos in two sections, one situated in
| |
| the head, and the other in the trunk.
| |
| | |
| The section in the trunk is the first to appear. The wall of the
| |
| alimentary canal becomes thickened along the median dorsal line (fig. 412,
| |
| r), or else produced into a ridge into which there penetrates a narrow
| |
| prolongation of the lumen of the alimentary canal. In either case the cells
| |
| at the extreme summit become gradually constricted off as a rod, which lies
| |
| immediately dorsal to the alimentary tract, and ventral to the notochord
| |
| (fig. 413, *).
| |
| | |
| | |
| | |
| | |
| | |
| FIG. 412. TRANSVERSE SECTION
| |
| THROUGH THE TAIL REGION OF A
| |
| PRISTIURUS EMBRYO OF THE SAME
| |
| AGE AS FIG. 28 E.
| |
| | |
| df. dorsal fin ; sp.c. spinal cord ;
| |
| //. body cavity ; sp. splanchnic layer
| |
| of mesoblast ; so. somatic layer of
| |
| mesoblast; mp'. portion of splanchnic
| |
| mesoblast commencing to be differentiated into muscles ; ch. notochord ; x.
| |
| subnotochordal rod arising as an outgrowth of the dorsal wall of the alimentary tract ; al. alimentary tract.
| |
| | |
| | |
| | |
| FIG. 413. TRANSVERSE SECTION THROUGH THE TRUNK OF AN
| |
| EMBRYO SLIGHTLY OLDER THAN
| |
| FIG. 28 E.
| |
| | |
| nc. neural canal ; pr. posterior
| |
| root of spinal nerve; x. subnotochordal rod; ao. aorta; sc. somatic
| |
| mesoblast; sp. splanchnic mesoblast; mp. muscle-plate; mp'. portion of muscle-plate converted into
| |
| muscle ; Vv. portion of the vertebral
| |
| plate which will give rise to the vertebral bodies ; al. alimentary tract.
| |
| | |
| | |
| | |
| In the hindermost part of the body its mode of formation differs somewhat from that above described. In this part the alimentary wall is' very
| |
| thick, and undergoes no special growth prior to the formation of the subnotochordal rod ; on the contrary, a small linear portion of the wall becomes
| |
| scooped out along the median dorsal line, and eventually separates from the
| |
| remainder as the rod in question. In the trunk the splitting off of the rod
| |
| takes place from before backwards, so that the anterior part of it is formed
| |
| before the posterior.
| |
| | |
| The section of the subnotochordal rod in the head would appear to
| |
| develop in the same way as that in the trunk, and the splitting off from the
| |
| throat proceeds from before backwards.
| |
| | |
| 482
| |
| | |
| | |
| | |
| 756 MESENTERY.
| |
| | |
| | |
| | |
| On the formation of the dorsal aorta, the subnotochordal rod becomes
| |
| separated from the wall of the gut and the aorta interposed between the two
| |
| (fig. 367, *).
| |
| | |
| When the subnotochordal rod attains its fullest development it terminates
| |
| anteriorly some way in front of the auditory vesicle, though a little behind
| |
| the end of the notochord ; posteriorly it extends very nearly to the extremity
| |
| of the tail and is almost co-extensive with the postanal section of the
| |
| alimentary tract, though it does not reach quite so far back as the caudal
| |
| vesicle (fig. 424, b x). Very shortly after it has attained its maximum size it
| |
| begins to atrophy in front. We may therefore conclude that its atrophy,
| |
| like its development, takes place from before backwards. During the later
| |
| embryonic stages not a trace of it is to be seen. It has also been met with
| |
| in Acipenser, Lepidosteus, the Teleostei, Petromyzon, and the Amphibia, in
| |
| all of which it appears to develop in fundamentally the same way as in
| |
| Elasmobranchii. In Acipenser it appears to persist in the adult as the
| |
| subvertebral ligament (Bridge, Salensky). It has not yet been found in a
| |
| fully developed form in any amniotic Vertebrate, though a thickening of the
| |
| hypoblast, which may perhaps be a rudiment of it, has been found by
| |
| Marshall and myself in the Chick (fig. 1 10, x).
| |
| | |
| Eisig has instituted an interesting comparison between it and an organ
| |
| which he has found in a family of Chaetopods, the Capitellidas. In these
| |
| forms there is a tube underlying the alimentary tract for nearly its whole
| |
| length, and opening into it in front, and probably behind. A remnant of
| |
| such a tube might easily form a rudiment like the subnotochordal rod of the
| |
| Ichthyopsida, and as Eisig points out the prolongation into the latter during
| |
| its formation of the lumen of the alimentary tract distinctly favours such a
| |
| view of its original nature. We can however hardly suppose that there is
| |
| any direct genetic connection between Eisig's organ in the Capitellidas and
| |
| the subnotochordal rod of the Chordata.
| |
| | |
| | |
| | |
| Splanchnic mesoblast and mesentery- The mesentcron
| |
| consists at first of a simple hypoblastic tube, which however
| |
| becomes enveloped by a layer of splanchnic mesoblast. This
| |
| layer, which is not at first continued over the dorsal side of the
| |
| mesenteron, gradually grows in, and interposes itself between the
| |
| hypoblast of the mesenteron, and the organs above. At the same
| |
| time it becomes differentiated into two layers, viz. an outer
| |
| cpithelioid layer which gives rise to part of the peritoneal
| |
| epithelium, and an inner layer of undifferentiated cells which in
| |
| time becomes converted into the connective tissue and muscular
| |
| walls of the mesenteron. The connective tissue layers become
| |
| first formed, while of the muscular layers the circular is the first
| |
| to make its appearance.
| |
| | |
| | |
| | |
| ALIMENTARY CANAL. 757
| |
| | |
| Coincidently with their differentiation the connective tissuestratum of the peritoneum becomes established.
| |
| | |
| The Mesentery. Prior to the splanchnic mesoblast growing
| |
| round the alimentary tube above, the attachment of the latter
| |
| structure to the dorsal wall of the body is very wide. On the
| |
| completion of this investment the layer of mesoblast suspending
| |
| the alimentary tract becomes thinner, and at the same time the
| |
| alimentary canal appears to be drawn downwards and away from
| |
| the vertebral column.
| |
| | |
| In what may be regarded as the thoracic division of the general
| |
| pleuroperitoneal space, along that part of the alimentary canal
| |
| which will form the oesophagus, this withdrawal is very slight, but
| |
| it is very marked in the abdominal region. In the latter the at
| |
| first straight digestive canal comes to be suspended from the body
| |
| above by a narrow flattened band of mesoblastic tissue. This
| |
| flattened band is the mesentery, shewn commencing in fig. 117,
| |
| and much more advanced in fig. 1 19, M. It is covered on either
| |
| side by a layer of flat cells, which form part of the general
| |
| peritoneal epithelioid lining, while its interior is composed of
| |
| indifferent tissue.
| |
| | |
| The primitive simplicity in the arrangement of the mesentery
| |
| is usually afterwards replaced by a more complicated disposition,
| |
| owing to the subsequent elongation and consequent convolution
| |
| of the intestine and stomach.
| |
| | |
| The layer of peritoneal epithelium on the ventral side of the
| |
| stomach is continued over the liver, and after embracing the liver,
| |
| becomes attached to the ventral abdominal wall (fig. 380). Thus
| |
| in the region of the liver the body cavity is divided into two
| |
| halves by a membrane, the two sides of which are covered by the
| |
| peritoneal epithelium, and which encloses the stomach dorsally
| |
| and the liver ventrally. The part of the membrane between the
| |
| stomach and liver is narrow, and constitutes a kind of mesentery
| |
| suspending the liver from the stomach : it is known to human
| |
| anatomists as the lesser omentum.
| |
| | |
| The part of the membrane connecting the liver with the
| |
| anterior abdominal wall constitutes the fa lei form or suspensory ligament of the liver. It arises by a secondary fusion, and
| |
| is not a remnant of a primitive ventral mesentery (vide pp. 624
| |
| and 625).
| |
| | |
| | |
| | |
| 758 MESENTERY.
| |
| | |
| | |
| | |
| The mesentery of the stomach, or mesogastrium, enlarges in
| |
| Mammalia to form a peculiar sack known as the greater
| |
| omentum.
| |
| | |
| The mesenteron exhibits very early a trifold division. An
| |
| anterior portion, extending as far as the stomach, becomes
| |
| separated off as the respiratory division. On the formation
| |
| of the anal invagination the portion of the mesenteron behind
| |
| the anus becomes marked off as the postanal division, and
| |
| between the postanal section and the respiratory division is a
| |
| middle portion forming an intestinal and cloacal division.
| |
| | |
| The respiratory division of the mesenteron.
| |
| | |
| This section of the alimentary canal is distinguished by the
| |
| fact that its walls send out a series of paired diverticula, which
| |
| meet the skin, and after a perforation has been effected at the
| |
| regions of contact, form the branchial or visceral clefts.
| |
| | |
| In Amphioxus the respiratory region extends close up to the
| |
| opening of the hepatic diverticulum, and therefore to a position
| |
| corresponding with the commencement of the intestine in higher
| |
| types. In the craniate Vertebrata the number of visceral clefts
| |
| has become reduced, but from the extension of the visceral clefts
| |
| in Amphioxus, combined with the fact that in the higher Vertebrata the vagus nerve, which is essentially the nerve of the
| |
| branchial pouches, supplies in addition the walls of the oesophagus
| |
| and stomach, it may reasonably be concluded, as has been pointed
| |
| out by Gegenbaur, that the true respiratory region primitively
| |
| included the region which in the higher types forms the
| |
| oesophagus and stomach.
| |
| | |
| In Ascidians the respiratory sack is homologous with the
| |
| respiratory tract of Amphioxus.
| |
| | |
| The details of the development of the branchial clefts in the
| |
| different groups of Vertebrata have already been described in
| |
| the systematic part of this work.
| |
| | |
| In all the Ichthyopsida the walls of a certain number of
| |
| clefts become folded ; and in the mesoblast within these folds a
| |
| rich capillary network, receiving its blood from the branchial
| |
| arteries, becomes established. These folds constitute the true
| |
| internal gills.
| |
| | |
| | |
| | |
| ALIMENTARY CANAL.
| |
| | |
| | |
| | |
| 759
| |
| | |
| | |
| | |
| In addition to internal gills external branchial processes covered
| |
| by epiblast are placed on certain of the visceral arches in the
| |
| larva of Polypterus, Protopterus and many Amphibia. The
| |
| external gills have probably no genetic connection with the
| |
| internal gills.
| |
| | |
| The so-called external gills of the embryos of Elasmobranchii
| |
| are merely internal gills prolonged outwards through the gill
| |
| clefts.
| |
| | |
| The posterior part of the primitive respiratory division of the
| |
| mesenteron becomes, in all the higher Vertebrata, the oesophagus
| |
| and stomach. With reference to the development of these parts
| |
| the only point worth especially noting is the fact that in
| |
| Elasmobranchii and Teleostei their lumen, though present in
| |
| very young embryos, becomes at a later stage completely filled
| |
| up, and thus the alimentary tract in the regions of the
| |
| oesophagus and stomach becomes a solid cord of cells (fig. 23
| |
| A, ces)\ as already suggested (p. 61) it seems not impossible that
| |
| this feature may be connected with the fact that the cesophageal
| |
| region of the throat was at one time perforated by gill clefts.
| |
| | |
| In addition to the gills two important organs, viz. the
| |
| thyroid body and the lungs, take their origin from the respiratory region of the alimentary tract.
| |
| | |
| Thyroid body. In the Ascidians the origin of a groovelike diverticulum of the ventral wall of the branchial sack,
| |
| bounded by two lateral folds, and known as the endostyle or
| |
| hypopharyngeal groove, has already been described (p. 18).
| |
| This groove remains permanently open to the pharyngeal sack,
| |
| | |
| | |
| | |
| | |
| FIG. 414. DIAGRAMMATIC VERTICAL SECTION OF A JUST-HATCHED LARVA
| |
| | |
| OF PETROMYZON. (From Gegenbaur ; after Calberla.)
| |
| | |
| o. mouth ; 6. olfactory pit ; v. septum between stomodteum and mesenteron ;
| |
| h. thyroid involution ; n. spinal cord ; ch. notochord; c. heart ; a. auditory vesicle.
| |
| | |
| | |
| | |
| 760
| |
| | |
| | |
| | |
| THE THYROID BODY.
| |
| | |
| | |
| | |
| | |
| and would seem to serve as a glandular organ secreting mucus.
| |
| As was first pointed out by W. Miiller there is present in
| |
| Amphioxus a very similar and probably homologous organ,
| |
| known as the hypopharyngeal groove.
| |
| | |
| In the higher Vertebrata this organ never retains its primitive condition in the adult state. In the larva of Petromyzon
| |
| there is, however, present a ventral groove-like diverticulum of
| |
| the throat, extending from about the second to the fourth
| |
| visceral cleft. This organ is shewn in longitudinal section in
| |
| fig. 414, h, and in transverse section in fig. 415, and has been
| |
| identified by W. Muller (Nos. 565 and 566) with the hypopharyngeal groove of Amphioxus and Ascidians. It does
| |
| not, however, long retain its
| |
| primitive condition, but its opening becomes gradually reduced
| |
| to a pore, placed between the
| |
| third and fourth of the permanent clefts (fig. 416, tli). This
| |
| opening is retained throughout
| |
| the Ammoccete condition, but
| |
| the organ becomes highly complicated, with paired anterior
| |
| and posterior horns and a
| |
| median spiral portion. In the adult the connection with the
| |
| pharynx is obliterated, and the organ is partly absorbed and
| |
| partly divided up into a series of glandular follicles, and eventually forms the thyroid body.
| |
| | |
| From the consideration of the above facts W. Muller was led
| |
| to the conclusion tJiat the tJiyroid body of the Craniata was
| |
| derived from the endostyle or Jiypopharyngeal groove. In all the
| |
| higher Vertebrata the thyroid body arises as a diverticulum of
| |
| the ventral wall of the throat in the region either of the mandibular or hyoid arches (fig. 417, Tk}, which after being segmented
| |
| off becomes divided up into follicles.
| |
| | |
| In Elasmobranch embryos it appears fairly early as a diverticulum from
| |
| the ventral surface of the throat in the region of the niandibular arc/i,
| |
| extending from the border of the mouth to the point where the ventral aorta
| |
| divides into the two aortic branches of the mandibular arch (fig. 417, Th}.
| |
| | |
| | |
| | |
| FIG. 415. DIAGRAMMATIC TRANSVERSE SECTIONS THROUGH THE BRANCHIAL REGION OF YOUNG LARV.K OF
| |
| PETROMYZON. (From Gegenbaur ; after
| |
| Calberla.)
| |
| | |
| d. branchial region of throat.
| |
| | |
| | |
| | |
| ALIMENTARY CANAL.
| |
| | |
| | |
| | |
| 761
| |
| | |
| | |
| | |
| Somewhat later it becomes in Scyllium and Torpedo solid, though still
| |
| retaining its attachment to the wall of the oesophagus. It continues to grow
| |
| in length, and becomes divided up into a number of solid branched lobules
| |
| separated by connective tissue septa. Eventually its connection with the
| |
| throat becomes lost, and the lobules develop a lumen. In Acanthias the
| |
| lumen of the gland is retained (W. Miiller) till after its detachment from the
| |
| | |
| | |
| | |
| -- "
| |
| | |
| | |
| Pti
| |
| | |
| | |
| | |
| | |
| FIG. 416. DIAGRAMMATIC VERTICAL SECTION THROUGH THE HEAD OF A
| |
| LARVA OF PETROMYZON.
| |
| | |
| The larva had been hatched three days, and was 4 '8 mm. in length. The optic
| |
| and auditory vesicles are supposed to be seen through the tissues. The letter tv
| |
| pointing to the base of the velum is where Scott believes the hyomandibular cleft to
| |
| be situated.
| |
| | |
| c.h. cerebral hemisphere ; th. optic thalamus; in. infundibulum ; pn. pineal gland ;
| |
| mb. mid-brain ; cb, cerebellum ; md. medulla oblongata ; au.v. auditory vesicle ; op.
| |
| optic vesicle; ol. olfactory pit; m. mouth; br.c. branchial pouches; th. thyroid
| |
| involution; v.ao. ventral aorta; ht. ventricle of heart ; ch. notochord.
| |
| | |
| throat. It preserves its embryonic position through life. In Amphibia it
| |
| originates, as in Elasmobranchii, from the region of the mandibular arch ;
| |
| but when first visible it forms a double epithelial wall connecting the throat
| |
| with the nervous layer of the epidermis. It subsequently becomes detached
| |
| from the epidermis, and then has the usual form of a diverticulum from the
| |
| throat. In most Amphibians it becomes divided into two lobes, and so
| |
| forms a paired body. The peculiar connection between the thyroid diverticulum and the epidermis in Amphibia has been noted by Gotte in
| |
| Bombinator, and by Scott and Osborn in Triton. It is not very easy to see
| |
| what meaning this connection can have.
| |
| | |
| In the Fowl (W. Miiller) the thyroid body arises at the end of the second
| |
| or beginning of the third day as an outgrowth from the hypoblast of the
| |
| throat, opposite the point of origin of the anterior arterial arch. This
| |
| outgrowth becomes by the fourth day a solid mass of cells, and by the fifth
| |
| ceases to be connected with the epithelium of the throat, becoming at the
| |
| same time bilobed. By the seventh day it has travelled somewhat backwards, and the two lobes have completely separated from each other. By
| |
| | |
| | |
| | |
| 762
| |
| | |
| | |
| | |
| THE THYROID BODY.
| |
| | |
| | |
| | |
| the ninth day the whole is invested by a
| |
| capsule of connective tissue, which sends
| |
| in septa dividing it into a number of lobes
| |
| or solid masses of cells, and by the sixteenth day it is a paired body composed of
| |
| a number of hollow branched follicles, each
| |
| with a ' membrana propria,' and separated
| |
| from each other by septa of connective
| |
| tissue. It finally travels back to the point
| |
| of origin of the carotids.
| |
| | |
| Amongst Mammalia the thyroid arises
| |
| in the Rabbit (Kolliker) and Man (His) as
| |
| a hollow diverticulum of the throat at the
| |
| bifurcation of the foremost pair of aortic
| |
| arches. It soon however becomes solid,
| |
| and is eventually detached from the throat
| |
| and comes to lie on the ventral side of the
| |
| larynx or windpipe. The changes it undergoes are in the main similar to those in the
| |
| lower Vertebrata. It becomes partially
| |
| constricted into two lobes, which remain
| |
| however united by an isthmus 1 . The fact
| |
| that the thyroid sometimes arises in the
| |
| region of the first and sometimes in that of
| |
| the second cleft is probably to be explained
| |
| | |
| | |
| | |
| | |
| Tli
| |
| | |
| | |
| | |
| FIG. 417. SECTION THROUGH
| |
| THE HEAD OF AN ELASMOBRANCH
| |
| EMBRYO, AT THE LEVEL OF THE
| |
| AUDITORY INVOLUTION.
| |
| | |
| Th. rudiment of thyroid body ;
| |
| aup. auditory pit ; aim. ganglion
| |
| of auditory nerve ; iv. v. roof of
| |
| fourth ventricle ; a.c.v. anterior
| |
| cardinal vein ; aa. aorta ; f.aa
| |
| aortic trunk of mandibular arch ;
| |
| //. head cavity of mandibular
| |
| arch ; Ivc. alimentary pouch which
| |
| will form the first visceral cleft.
| |
| | |
| | |
| | |
| by its rudimentary character.
| |
| | |
| The Thymus gland. The thymus gland may conveniently be
| |
| dealt with here, although its origin is nearly as obscure as its function. It
| |
| has usually been held to be connected with the lymphatic system. Kolliker
| |
| was the first to shew that this view was probably erroneous, and he
| |
| attempted to prove that it was derived in the Rabbit from the walls of one
| |
| of the visceral clefts, mainly on the ground of its presenting in the embryo
| |
| an epithelial character.
| |
| | |
| 1 Wolfler (No. 571) states that in the Pig and Calf the thyroid body is formed as a
| |
| pair of epithelial vesicles, which are developed as outgrowths of the walls of the first
| |
| pair of visceral clefts. He attempts to explain the contradictory observations of other
| |
| embryologists by supposing that they have mistaken the ventral ends of visceral
| |
| pouches for an unpaired outgrowth of the throat. Stieda (No. 569) also states that in
| |
| the Pig and Sheep the thyroid arises as a paired body from the epithelium of a pair
| |
| of visceral clefts, at a much later period than would appear from the observations of
| |
| His and Kolliker. In view of the comparative development of this organ it is
| |
| difficult to accept either Wolfler's or Stieda's account. Wolfler's attempt to explain
| |
| the supposed errors of his predecessors is certainly not capable of being applied in
| |
| the case of Elasmobranch Fishes, or of Petromyzon ; and I am inclined to think that
| |
| the method of investigation by transverse sections, which has been usually employed,
| |
| is less liable to error than that by longitudinal sections which he has adopted.
| |
| | |
| | |
| | |
| ALIMENTARY CANAL. 763
| |
| | |
| | |
| | |
| Stieda (No. 569) has recently verified Kolliker's statements. He finds
| |
| that in the Pig and the Sheep the thymus arises as a paired outgrowth from
| |
| the epithelial remnants of a pair of visceral clefts. Its two lobes may at first
| |
| be either hollow (Sheep) or solid (Pig), but eventually become solid, and
| |
| unite in the median line. Stieda and His hold that in the adult gland, the
| |
| so-called corpuscles of Hassall are the remnants of the embryonic epithelial
| |
| part of the gland, and that the lymphatic part of it is of mesoblastic origin ;
| |
| but Kolliker believes the lymphatic cells to be direct products of the
| |
| embryonic epithelial cells.
| |
| | |
| The posterior visceral clefts in the course of their atrophy give rise to
| |
| various more or less conspicuous bodies of a pseudo-glandular nature, which
| |
| have been chiefly studied by Remak 1 .
| |
| | |
| Swimming bladder and lungs. A swimming bladder is
| |
| present in all Ganoids and in the vast majority of Teleostei.
| |
| Its development however is only imperfectly known.
| |
| | |
| In the Salmon and Carp it arises, as was first shewn by Von
| |
| Baer, as an outgrowth of the alimentary tract, shortly in front of
| |
| the liver. In these forms it is at first placed on the dorsal side
| |
| and slightly to the right, and grows backwards on the dorsal
| |
| side of the gut, between the two folds of the mesentery.
| |
| | |
| The absence of a pneumatic duct in the Physoclisti would
| |
| appear to be due to a post-larval atrophy.
| |
| | |
| In Lepidosteus the air-bladder appears to arise, as in the
| |
| Teleostei, as an invagination of the dorsal wall of the oesophagus.
| |
| | |
| In advanced embryos of Galeus, Mustelus and Acanthias, MikluchoMaclay detected a small diverticulum opening on the dorsal side of the
| |
| oesophagus, which he regards as a rudiment of a swimming bladder. This
| |
| interpretation must however be regarded as somewhat doubtful.
| |
| | |
| The lungs. The lungs originate in a nearly identical way in
| |
| all the Vertebrate forms in which their development has been
| |
| observed. They are essentially buds or processes of the ventral
| |
| wall of the primitive oesophagus.
| |
| | |
| At a point immediately behind the region of the visceral
| |
| clefts the cavity of the alimentary canal becomes compressed
| |
| laterally, and at the same time constricted in the middle, so that
| |
| its transverse section (fig. 418 i) is somewhat hourglass-shaped,
| |
| and shews an upper or dorsal chamber d, joining on to a lower
| |
| or ventral chamber / by a short narrow neck.
| |
| | |
| 1 For details on these organs vide Kolliker, Entwicklungsgeschichte, p. 88 1.
| |
| | |
| | |
| | |
| 764
| |
| | |
| | |
| | |
| THE LUNGS.
| |
| | |
| | |
| | |
| | |
| The hinder end of the lower tube enlarges (fig. 418 2), and
| |
| then becomes partially divided into two lobes (fig. 418 3). All
| |
| these parts at first freely communicate, but the two lobes,
| |
| partly by their own growth,
| |
| and partly by a process of constriction, soon become isolated
| |
| posteriorly; while in front they
| |
| open into the lower chamber
| |
| of the oesophagus (fig. 422).
| |
| | |
| By a continuation forwards
| |
| of the process of constriction
| |
| the lower chamber of the oesophagus, carrying with it the
| |
| two lobes above mentioned,
| |
| becomes gradually transformed
| |
| into an independent tube,
| |
| opening in front by a narrow
| |
| slit-like aperture into the oesophagus. The single tube in
| |
| front is the rudiment of the
| |
| trachea and larynx, while the
| |
| two diverticula behind become
| |
| (fig. 419, Ig) the bronchial tubes
| |
| and lungs.
| |
| | |
| While the above changes
| |
| are taking place in the hypoblastic walls of the alimentary
| |
| tract, the splanchnic mesoblast
| |
| surrounding these structures
| |
| becomes very much thickened ; but otherwise bears no marks of
| |
| the internal changes which are going on, so that the above
| |
| formation of the lungs and trachea cannot be seen from the
| |
| surface. As the paired diverticula of the lungs grow backwards,
| |
| the mesoblast around them takes however the form of two lobes,
| |
| into which they gradually bore their way.
| |
| | |
| There do not seem to be any essential differences in the mode of
| |
| formation of the above structures in the types so far observed, viz. Amphibia,
| |
| Aves and Mammalia. Writers differ as to whether the lungs first arise as
| |
| | |
| | |
| | |
| FlG. 418. FOUR DIAGRAMS ILLUSTRATING THE FORMATION OF THE LUNGS.
| |
| | |
| (After Gotte.)
| |
| | |
| a. mesoblast; b. hypoblast; d. cavity
| |
| of digestive canal ; /. cavity of the pulmonary diverticulum.
| |
| | |
| In (i) the digestive canal has commenced to be constricted into an upper
| |
| and lower canal ; the former the true
| |
| alimentary canal, the latter the pulmonary tube; the two tubes communicate
| |
| with each other in the centre.
| |
| | |
| In (2) the lower (pulmonary) tube has
| |
| become expanded.
| |
| | |
| In (3) the expanded portion of the
| |
| tube has become constricted into two
| |
| tubes, still communicating with each other
| |
| and with the digestive canal.
| |
| | |
| In (4) these are completely separated
| |
| from each other and from the digestive
| |
| canal, and the mesoblast has also begun
| |
| to exhibit externally changes corresponding to the internal changes which have
| |
| been going on.
| |
| | |
| | |
| | |
| ALIMENTARY CANAL.
| |
| | |
| | |
| | |
| 765
| |
| | |
| | |
| | |
| re
| |
| | |
| | |
| | |
| paired diverticula, or as a single diverticulum ; and as to whether the
| |
| rudiments of the lungs are established
| |
| before those of the trachea. If the above
| |
| account is correct it would appear that
| |
| any of these positions might be maintained. Phylogenetically interpreted the
| |
| ontogeny of the lungs appears however
| |
| to imply that this organ was first an
| |
| unpaired structure and has become
| |
| secondarily paired, and that the trachea
| |
| was relatively late in appearing.
| |
| | |
| The further development of the
| |
| lungs is at first, in the higher types
| |
| at any rate, essentially similar to
| |
| that of a racemose gland. From
| |
| each primitive diverticulum numerous branches are given off
| |
| In Aves and Mammalia (fig. 355)
| |
| they are mainly confined to the
| |
| dorsal and lateral parts. These
| |
| branches penetrate into the surrounding mesoblast and continue
| |
| to give rise to secondary and
| |
| tertiary branches. In the meso
| |
| | |
| | |
| | |
| At
| |
| | |
| | |
| | |
| FIG. 419. SECTION THROUGH
| |
| THE CARDIAC REGION OF AN EMBRYO
| |
| OF LACERTA MURALIS OF 9 MM. TO
| |
| SHEW THE MODE OF FORMATION OF
| |
| THE PERICARDIAL CAVITY.
| |
| | |
| ht. heart ; pc . pericardial cavity ;
| |
| al. alimentary tract; Ig. lung; /.
| |
| liver; pp. body cavity; md. open
| |
| end of Mullerian duct; wd. Wolffian
| |
| duct ; vc. vena cava inferior ; ao.
| |
| aorta; ch. notochord; me, medullary
| |
| cord.
| |
| | |
| | |
| | |
| blast around them numerous capillaries make their appearance, and the further growth of the
| |
| bronchial tubes is supposed by Boll to be due to the mutual
| |
| interaction of the hitherto passive mesoblast and of the hypoblast.
| |
| | |
| The further changes in the lungs vary somewhat in the different forms.
| |
| | |
| The air sacks are the most characteristic structures of the avian lung.
| |
| They are essentially the dilated ends of the primitive diverticula or of their
| |
| main branches.
| |
| | |
| In Mammalia (Kolliker, No. 298) the ends of the bronchial tubes become
| |
| dilated into vesicles, which may be called the primary air-cells. At first,
| |
| owing to their development at the ends of the bronchial branches, these are
| |
| confined to the surface of the lungs. At a later period the primary air-cells
| |
| divide each into two or three parts, and give rise to secondary air-cells, while
| |
| at the same time the smallest bronchial tubes, which continue all the while
| |
| to divide, give rise at all points to fresh air-cells. Finally the bronchial
| |
| tubes cease to become more branched, and the air-cells belonging to each
| |
| minute lobe come in their further growth to open into a common chamber.
| |
| | |
| | |
| | |
| 766 THE CLOACA.
| |
| | |
| | |
| | |
| Before the lungs assume their function the embryonic air-cells undergo a
| |
| considerable dilatation.
| |
| | |
| The trachea and larynx. The development of the trachea and larynx
| |
| does not require any detailed description. The larynx is formed as a simple
| |
| dilatation of the trachea. The cartilaginous structures of the larynx are of
| |
| the same nature as those of the trachea.
| |
| | |
| It follows from the above account that the whole pulmonary
| |
| structure is the result of the growth by budding of a system of
| |
| branched hypoblastic tubes in the midst of a mass of mesoblastic
| |
| tissue, the hypoblastic elements giving rise to the epithelium of
| |
| the tubes, and the mesoblast providing the elastic, muscular,
| |
| cartilaginous, vascular, and other connective tissues of the
| |
| tracheal and bronchial walls.
| |
| | |
| There can be no doubt that the lungs and air-bladder are
| |
| homologous structures, and the very interesting memoir of Eisig
| |
| on the air-bladder of the Chaetopoda 1 shews it to be highly
| |
| probable that they are the divergent modifications of a primitive
| |
| organ, which served as a reservoir for gas secreted in the
| |
| alimentary tract, the gas in question being probably employed
| |
| for respiration when, for any reason, ordinary respiration by the
| |
| gills was insufficient.
| |
| | |
| Such an organ might easily become either purely respiratory,
| |
| receiving its air from the exterior, and so form a true lung ; or
| |
| mainly hydrostatic, forming an air-bladder, as in Ganoidei and
| |
| Teleostei.
| |
| | |
| It is probable that in the Elasmobranchii the air-bladder has
| |
| become aborted, and the organ discovered by Micklucho-Maclay
| |
| may perhaps be a last remnant of it.
| |
| | |
| The middle division of the mesenteron. The middle
| |
| division of the mesenteron, forming the intestinal and cloacal
| |
| region, is primitively a straight tube, the intestinal region of
| |
| which in most Vertebrate embryos is open below to the yolksack.
| |
| | |
| Cloaca. In the Elasmobranchii, the embryos of which
| |
| probably retain a very primitive condition of the mesenteron,
| |
| this region is not at first sharply separated from the postanal
| |
| section behind. Opposite the point where the anus will even
| |
| 1 H. Eisig, " Ueb. d. Vorkommen eines schwimmblasenahnlichen Organs bei
| |
| Anneliden." Mittheil. a. d. zool. Station z. Neafel, Vol. II. 1881.
| |
| | |
| | |
| | |
| ALIMENTARY CANAL.
| |
| | |
| | |
| | |
| 767
| |
| | |
| | |
| | |
| tually appear a dilatation of the mesenteron arises, which comes
| |
| in contact with the external skin (fig. 28 E, an}. This dilatation
| |
| becomes the hypoblastic section of the cloaca. It communicates
| |
| behind with the postanal gut (fig. 424 D), and in front with the
| |
| intestine ; and may be defined as the dilated portion of the alimentary tract which receives the genital and urinary ducts and opens
| |
| externally by the proctodczum.
| |
| | |
| In Acipenser and Amphibia the cloacal region is indicated
| |
| as a ventral diverticulum of the mesenteron even before the
| |
| closure of the blastopore. It is shewn in the Amphibia at an
| |
| early stage in fig. 73, and at a later period, when in contact with
| |
| the skin at the point where the anal invagination is about to
| |
| appear, in fig. 420.
| |
| | |
| | |
| | |
| | |
| FIG. 420. LONGITUDINAL SECTION THROUGH AN ADVANCED EMBRYO OF
| |
| | |
| BOMBINATOR. (After Gotte.)
| |
| | |
| m. mouth ; an. anus ; /. liver ; ne. neurenteric canal ; me. medullary canal ; ch.
| |
| notochord ; pn. pineal gland.
| |
| | |
| In the Sauropsida and Mammalia the cloaca appears as a
| |
| dilatation of the mesenteron, which receives the opening of the
| |
| allantois almost as soon as the posterior part of the mesenteron
| |
| is established.
| |
| | |
| The eventual changes which it undergoes have been already
| |
| dealt with in connection with the urinogenital organs.
| |
| | |
| Intestine. The region in front of the cloaca forms the
| |
| intestine. In certain Vertebrata it nearly retains its primitive
| |
| character as a straight tube ; and in these types its anterior
| |
| part is characterised by the presence of a peculiar fold, which in
| |
| a highly specialised condition is known as the spiral valve.
| |
| This structure appears in its simplest form in Ammocoetes. It
| |
| | |
| | |
| | |
| 768 THE INTESTINE.
| |
| | |
| | |
| | |
| there consists of a fold in the wall of the intestine, giving to the
| |
| lumen of this canal a semilunar form in section, and taking a
| |
| half spiral.
| |
| | |
| In Elasmobranchii a similar fold to that in Ammoccetes first
| |
| makes its appearance in the embryo. This fold is from the
| |
| first not quite straight, but winds in a long spiral round the
| |
| intestine. In the course of development it becomes converted
| |
| into a strong ridge projecting into the lumen of the intestine
| |
| (fig. 388, /). The spiral it makes becomes much closer, and it
| |
| thus acquires the form of the adult spiral valve. A spiral valve
| |
| is also found in Chimaera and Ganoids. No rudiment of such
| |
| an organ is found in the Teleostei, the Amphibia, or the higher
| |
| Vertebrata.
| |
| | |
| The presence of this peculiar organ appears to be a very
| |
| primitive Vertebrate character. The intestine of Ascidians
| |
| exhibits exactly the same peculiarity as that of Ammoccetes,
| |
| and we may probably conclude from embryology that the
| |
| ancestral Chordata were provided with a straight intestine
| |
| having a fold projecting into its lumen, to increase the area of
| |
| the intestinal epithelium.
| |
| | |
| In all forms in which there is not a spiral valve, with the
| |
| exception of a few Teleostei, the intestine becomes considerably
| |
| longer than the cavity which contains it, and therefore necessarily more or less convoluted.
| |
| | |
| The posterior part usually becomes considerably enlarged to
| |
| form the rectum or in Mammalia the large intestine.
| |
| | |
| In Elasmobranchii there is a peculiar gland opening into the
| |
| dorsal side of the rectum, and in many other forms there is a
| |
| caecum at the commencement of the rectum or of the large
| |
| intestine.
| |
| | |
| In Teleostei, the Sturgeon and Lepidosteus there opens into
| |
| the front end of the intestine a number of caecal pouches known
| |
| as the pancreatic caeca. In the adult Sturgeon these pouches
| |
| unite to form a compact gland, but in the embryo they arise as
| |
| a series of isolated outgrowths of the duodenum.
| |
| | |
| Connected with the anterior portion of the middle region of
| |
| the alimentary canal, which may be called the duodenum, are
| |
| two very important and constant glandular organs, the liver and
| |
| the pancreas.
| |
| | |
| | |
| | |
| ALIMENTARY CANAL.
| |
| | |
| | |
| | |
| 769
| |
| | |
| | |
| | |
| ITlf
| |
| | |
| | |
| | |
| | |
| The liver. The liver is the earliest formed and largest
| |
| glandular organ in the embryo.
| |
| | |
| It appears in its simplest
| |
| form in Amphioxus as a single
| |
| unbranched diverticulum of the
| |
| alimentary tract, immediately
| |
| behind the respiratory region,
| |
| which is directed forwards and
| |
| placed on the left side of the
| |
| body.
| |
| | |
| In all true Vertebrata the
| |
| gland has a much more complicated structure. It arises as a
| |
| ventral outgrowth of the duodenum (fig. 420, /). This outgrowth may be at first single,
| |
| and then grow out into two
| |
| lobes, as in Elasmobranchii (fig.
| |
| 421) and Amphibia, or have from
| |
| the first the form of two somewhat unequal diverticula, as in
| |
| Birds (fig. 422), or again as in
| |
| the Rabbit (Kolliker) one diverticulum may be first formed, and a second one appear
| |
| somewhat later. The hepatic diverticula, whatever may be
| |
| their primitive form, grow into a special thickening of the
| |
| splanchnic mesoblast.
| |
| | |
| From the primitive diverticula there are soon given off a
| |
| number of hollow buds (fig. 421) which rapidly increase in
| |
| length and number, and form the so-called hepatic cylinders.
| |
| They soon anastomose and unite together, and so constitute an
| |
| irregular network. Coincidently with the formation of the
| |
| hepatic network the united vitelline and visceral vein or veins
| |
| (u.v\ in their passage through the liver, give off numerous
| |
| branches, and gradually break up into a plexus of channels
| |
| which form a secondary network amongst the hepatic cylinders.
| |
| In Amphibia these channels are stated by Gotte to be lacunar,
| |
| but in Elasmobranchii, and probably Vertebrata generally, they
| |
| arc from the first provided with distinct though delicate walls.
| |
| B. in. 49
| |
| | |
| | |
| | |
| FIG. 421. SECTION THROUGH THE
| |
| VENTRAL PART OF THE TRUNK OF A
| |
| YOUNG EMBRYO OF SCYLLIUM AT THE
| |
| LEVEL OF THE UMBILICAL CORD.
| |
| | |
| b. pectoral fin ; ao. dorsal aorta ;
| |
| cav. cardinal vein; ua. vitelline artery ; nv. vitelline vein united with
| |
| subintestinal vein ; al. duodenum ;
| |
| /. liver ; sd. opening of segmental
| |
| duct into the body-cavity ; mp. muscle-plate ; urn. umbilical canal.
| |
| | |
| | |
| | |
| 770
| |
| | |
| | |
| | |
| THE LIVER.
| |
| | |
| | |
| | |
| It is still doubtful whether the hepatic cylinders are as a rule hollow or
| |
| solid. In Elasmobranchii they are at first provided with a large lumen,
| |
| which though it becomes gradually smaller never entirely vanishes. The
| |
| same seems to hold good for Amphibia and some Mammalia. In Aves
| |
| the lumen of the cylinders is even from the first much more difficult
| |
| to see, and the cylinders are stated by Remak to be solid, and he has
| |
| been followed in this matter by Kolliker. In the Rabbit also Kolliker finds
| |
| the cylinders to be solid.
| |
| | |
| The embryonic hepatic network gives rise to the parenchyma
| |
| of the adult liver, with which in
| |
| its general arrangement it closely
| |
| agrees. The blood-channels are
| |
| at first very large, and have a
| |
| very irregular arrangement ; and
| |
| it is not till comparatively late
| |
| that the hepatic lobules with their
| |
| characteristic vascular structures
| |
| become established.
| |
| | |
| The biliary ducts are formed
| |
| either from some of the primitive hepatic cylinders, or, as
| |
| would seem to be the case in
| |
| Elasmobranchii and Birds (fig.
| |
| 422), from the larger diverticula of the two primitive outgrowths.
| |
| | |
| The gall-bladder is so inconstant, and the arrangement of
| |
| the ducts opening into the intestine so variable, that no general statements can be made about
| |
| them. In Elasmobranchii the primitive median diverticulum
| |
| (fig. 421) gives rise to the ductus choledochus. Its anterior end
| |
| dilates to form a gall-bladder.
| |
| | |
| In the Rabbit a ductus choledochus is formed by a diverticulum from the intestine at the point of insertion of the two
| |
| primitive lobes. The gall-bladder arises as a diverticulum of
| |
| the right primitive lobe.
| |
| | |
| The liver is relatively very large during embryonic life and
| |
| has, no doubt, important functions in connection with the circulation.
| |
| | |
| | |
| | |
| | |
| r
| |
| | |
| | |
| | |
| FIG. 422. DIAGRAM OF THE DIGESTIVE TRACT OF A CHICK UPON THE
| |
| FOURTH DAY. (After Gotte.)
| |
| | |
| The black line indicates the hypoblast. The shaded part around it is
| |
| the splanchnic mesoblast.
| |
| | |
| Ig. lung ; st. stomach ; p. pancreas ;
| |
| /. liver.
| |
| | |
| | |
| | |
| ALIMENTARY CANAL.
| |
| | |
| | |
| | |
| 771
| |
| | |
| | |
| | |
| The pancreas. So far as is known the development of the
| |
| pancreas takes place on a very constant type throughout the
| |
| series of craniate Vertebrata, though absent in some of the
| |
| Teleostean fishes and Cyclostomata, and very much reduced in
| |
| most Teleostei and in Petromyzon.
| |
| | |
| It arises nearly at the same time as the liver in the form of a
| |
| hollow outgrowth from the dorsal side of the intestine nearly
| |
| opposite but slightly behind the hepatic outgrowth (fig. 422, /).
| |
| It soon assumes, in Elasmobranchii and Mammalia, somewhat
| |
| the form of an inverted funnel, and from the expanded dorsal
| |
| part of the funnel there grow out numerous hollow diverticula
| |
| into the passive splanchnic mesoblast.
| |
| | |
| As the ductules grow longer and become branched, vascular
| |
| processes grow in between them, and the whole forms a compact
| |
| glandular body in the mesentery on the dorsal side of the
| |
| alimentary tract. The funnel-shaped receptacle loses its origi nal form, and elongating, assumes the character of a duct.
| |
| | |
| From the above mode of development it is clear that the
| |
| glandular cells of the pancreas are derived from the hypoblast.
| |
| | |
| Into the origin of the varying arrangements of the pancreatic
| |
| ducts it is not possible to enter in detail. In some cases,
| |
| e.g. the Rabbit (Kolliker), the two lobes and ducts arise from a
| |
| division of the primitive gland and duct. In other cases, e.g. the
| |
| Bird, a second diverticulum springs from the alimentary tract.
| |
| In a large number of instances the primitive condition with a
| |
| single duct is retained.
| |
| | |
| Postanal section of the mesenteron. In the embryos of
| |
| all the Chordata there is a section of the mesenteron placed
| |
| behind the anus. This section invariably atrophies at a comparatively early period of embryonic life ; but it is much better
| |
| developed in the lower forms than in the higher. At its
| |
| posterior extremity it is primitively continuous with the neural
| |
| tube (fig. 420), as was first shewn by Kowalevsky.
| |
| | |
| The canal connecting the neural and alimentary canals has
| |
| already been described as the neurenteric canal, and represents
| |
| the remains of the blastopore.
| |
| | |
| In the Tunicata the section of the mesenteron, which in all probability
| |
| corresponds to the postanal gut of the Vertebrata, is that immediately
| |
| | |
| 492
| |
| | |
| | |
| | |
| | |
| 772 POSTANAL SECTION OF THE MESENTERON.
| |
| | |
| following the dilated portion which gives rise to the branchial cavity
| |
| | |
| and permanent intestine. It has already
| |
| | |
| been shewn that from the dorsal and
| |
| | |
| lateral portions of this section of the
| |
| | |
| primitive alimentary tract the notochord
| |
| | |
| and muscles of the Ascidian tadpole are
| |
| | |
| derived. The remaining part of its walls
| |
| | |
| forms a solid cord of cells (fig. 423, al'},
| |
| | |
| which either atrophies, or, according to
| |
| | |
| Kowalevsky, gives rise to blood-vessels.
| |
| | |
| In Amphioxus the postanal gut, FIG. 423. TRANSVERSE OPTICAL
| |
| | |
| .hough distinctly developed, is no, very %
| |
| long, and atrophies at a comparatively (After Kowalevsky.)
| |
| early period. The sect i on ; s f rom an embryo of
| |
| | |
| In Elasmobranchii this section of the the same age as fig. 8 iv.
| |
| | |
| alimentary tract is very well developed, ch - notochord ; nc neural 1 canal ;
| |
| | |
| . , , me. mesoblast ; of. hypoblast of
| |
| and persists for a considerable period of ta ji <
| |
| | |
| embryonic life. The following is a
| |
| history of its development in the genus Scyllium.
| |
| | |
| Shortly after the stage when the anus has become marked out by the
| |
| alimentary tract sending down a papilliform process towards the skin, the
| |
| postanal gut begins to develop a terminal dilatation or vesicle, connected
| |
| with the remainder of the canal by a narrower stalk.
| |
| | |
| The walls both of the vesicle and stalk are formed of a fairly columnar
| |
| epithelium. The vesicle communicates in front by a narrow passage with
| |
| the neural canal, and behind is continued into two horns corresponding
| |
| with the two caudal swellings previously spoken of (p. 55). Where the
| |
| canal is continued into these two horns, its walls lose their distinctness of
| |
| outline, and become continuous with the adjacent mesoblast.
| |
| | |
| In the succeeding stages, as the tail grows longer and longer, the postanal section of the alimentary tract grows with it, without however undergoing alteration in any of its essential characters. At the period of the
| |
| maximum development, it has a length of about -J of that of the whole
| |
| alimentary tract.
| |
| | |
| Its features at a stage shortly before the external gills have become
| |
| prominent are illustrated by a series of transverse sections through the
| |
| tail (fig. 424). The four sections have been selected for illustration out of a
| |
| fairly-complete series of about one hundred and twenty.
| |
| | |
| Posteriorly (A) there is present a terminal vesicle (alv) '25 mm. in
| |
| diameter, which communicates dorsally by a narrow opening with the
| |
| neural canal (nc) ; to this is attached a stalk in the form of a tube, also
| |
| lined by columnar epithelium, and extending through about thirty sections
| |
| (B al}. Its average diameter is about '084 mm., and its walls are very thick.
| |
| Overlying its front end is the subnotochordal rod (x), but this does not
| |
| extend as far back as the terminal vesicle.
| |
| | |
| The thick-walled stalk of the vesicle is connected with the cloacal section
| |
| | |
| | |
| | |
| ALIMENTARY CANAL.
| |
| | |
| | |
| | |
| 773
| |
| | |
| | |
| | |
| of the alimentary tract by a very narrow thin-walled tube (C of). This for
| |
| the most part has a fairly uniform calibre, and a diameter of not more than
| |
| 035 mm. Its walls are formed of flattened epithelial cells. At a point not
| |
| far from the cloaca it becomes smaller, and its diameter falls to -03 mm. In
| |
| | |
| | |
| | |
| | |
| cl.al
| |
| | |
| | |
| | |
| FIG. 424. FOUR SECTIONS THROUGH THE POSTANAL PART OF THE TAIL
| |
| OF AN EMBRYO OF THE SAME AGE AS FIG. 28 F.
| |
| | |
| A. is the posterior section.
| |
| | |
| nc . neural canal ; al. postanal gut ; alv. caudal vesicle of postanal gut ; x.
| |
| subnotochordal rod; mp. muscle-plate; ch. notochord; cl.al. cloaca; ao. aorta;
| |
| v.cau, caudal vein.
| |
| | |
| front of this point it rapidly dilates again, and, after becoming fairly wide,
| |
| opens on the dorsal side of the cloacal section of the alimentary canal just
| |
| behind the anus (D al}.
| |
| | |
| Very shortly after the stage to which the above figures belong, at a
| |
| point a little behind the anus, where the postanal section of the canal
| |
| was thinnest in the previous stage, it becomes solid, and a rupture here
| |
| occurs in it at a slightly later period.
| |
| | |
| The atrophy of this part of the alimentary tract having once commenced
| |
| proceeds rapidly. The posterior part first becomes reduced to a small
| |
| rudiment near the end of the tail. There is no longer a terminal vesicle,
| |
| nor a neurenteric canal. The portion of the postanal section of the
| |
| alimentary tract, just behind the cloaca, is for a short time represented
| |
| by a small rudiment of the dilated part which at an earlier period opened
| |
| into the cloaca.
| |
| | |
| In Teleostei the vesicle at the end of the tail, discovered by Kupffer,
| |
| | |
| | |
| | |
| 774 THE STOMOD/EUM.
| |
| | |
| | |
| | |
| (fig- 34> hyv) is probably the equivalent of the vesicle at the end of the
| |
| postanal gut in Elasmobranchii.
| |
| | |
| In Petromyzon and in Amphibia there is a well-developed postanal
| |
| gut connected with a neurenteric canal which gradually atrophies. It is
| |
| shewh in the embryo of Bombinator in fig. 420.
| |
| | |
| Amongst the amniotic Vertebrata the postanal gut is less developed
| |
| than in the Ichthyopsida. A neurenteric canal is present for a short period
| |
| | |
| | |
| | |
| | |
| FIG. 425. DIAGRAMMATIC LONGITUDINAL SECTION THROUGH THE POSTERIOR
| |
| END OF AN EMBRYO BlRD AT THE TIME OF THE FORMATION OF THE ALLANTOIS.
| |
| | |
| ep. epiblast ; Sp.c. spinal canal ; ch. notochord ; n.e. neurenteric canal ; hy. hypoblast ; p.a.g, postanal gut ; pr. remains of primitive streak folded in on the ventral
| |
| side ; al. allantois ; me. splanchnic mesoblast ; an. point where anus will be formed ;
| |
| p.c. perivisceral cavity ; am. amnion ; so. somatopleure ; sp. splanchnopleure.
| |
| | |
| in various Birds (Gasser, etc.) and in the Lizard, but disappears very early.
| |
| There is however, as has been pointed out by Kolliker, a well-marked
| |
| postanal gut continued as a narrow tube from behind the cloaca into
| |
| the tail both in the Bird (fig. 425, p.a.g.} and Mammals (the Rabbit), but
| |
| especially in the latter. It atrophies early as in lower forms.
| |
| | |
| The morphological significance of the postanal gut and of the neurenteric canal has already been spoken of in Chapter xii., p. 323.
| |
| | |
| | |
| | |
| The anterior section of the permanent alimentary tract is
| |
| formed by an invagination of epiblast, constituting a more or
| |
| less considerable pit, with its inner wall in contact with the
| |
| blind anterior extremity of the alimentary tract.
| |
| | |
| In Ascidians this pit is placed on the dorsal surface (fig. 9, o),
| |
| and becomes the permanent oral cavity of these forms. In the
| |
| larva of Amphioxus it is stated to be formed unsymmetrically
| |
| | |
| | |
| | |
| THE STOMOD/EUM.
| |
| | |
| | |
| | |
| 775
| |
| | |
| | |
| | |
| | |
| (vide p. 5), but further observations on its development are
| |
| required.
| |
| | |
| In the true Vertebrata it is always formed on the ventral
| |
| surface of the head, immediately behind the level of the forebrain (fig. 426), and is deeper in Petromyzon (fig. 416, ;) than
| |
| in any other known form.
| |
| | |
| From the primary buccal cavity or stomodaeum there grows
| |
| out the pituitary pit (fig. 426, pt\ the
| |
| development of which has already
| |
| been described (p. 435).
| |
| | |
| The wall separating the stomodaeum from the mesenteron always
| |
| becomes perforated, usually at an
| |
| early stage of development, and
| |
| though in Petromyzon the boundary
| |
| between the two cavities remains
| |
| indicated by the velum, yet in the
| |
| higher Vertebrata all trace of this
| |
| boundary is lost, and the original
| |
| limits of the primitive buccal cavity
| |
| become obliterated ; while a secondary buccal cavity, partly lined by
| |
| hypoblast and partly by epiblast,
| |
| becomes established.
| |
| | |
| This cavity, apart from the organs which belong to it,
| |
| presents important variations in structure. In most Pisces it
| |
| retains a fairly simple character, but in the Dipnoi its outer
| |
| boundary becomes extended so as to enclose the ventral opening of the nasal sack, which thenceforward constitutes the
| |
| posterior nares.
| |
| | |
| In Amphibia and Amniota the posterior nares also open well
| |
| within the boundary of the buccal cavity.
| |
| | |
| In the Amniota further important changes take place.
| |
| | |
| In the first place a plate grows inwards from each of the
| |
| superior maxillary processes (fig. 427, /), and the two plates,
| |
| meeting in the middle line, form a horizontal septum dividing
| |
| the front part of the primitive buccal cavity into a dorsal
| |
| respiratory section (), containing the opening of the posterior
| |
| nares, and a ventral cavity, forming the permanent mouth. The
| |
| | |
| | |
| | |
| FIG. 426. LONGITUDINAL
| |
| SECTION THROUGH THE BRAIN OF
| |
| A YOUNG PRISTIURUS EMBRYO.
| |
| | |
| r.unpaired rudimentofthecerebral hemispheres \pn. pineal gland ;
| |
| /w.infundibulum ; //.ingrowth from
| |
| mouth to form the pituitary body ;
| |
| mb. mid-brain ; cb. cerebellum ; ch.
| |
| notochord; al. alimentary tract;
| |
| Zaa. artery of mandibular arch.
| |
| | |
| | |
| | |
| THE TEETH.
| |
| | |
| | |
| | |
| | |
| two divisions thus formed open into a common cavity behind.
| |
| The horizontal septum, on the development within it of an
| |
| osseous plate, constitutes the hard palate.
| |
| | |
| An internasal septum (fig. 427, e) may more or less completely divide the dorsal cavity into two canals, continuous
| |
| respectively with the two nasal cavities.
| |
| | |
| In Mammalia a posterior prolongation of the palate, in which
| |
| an osseous plate is not formed, constitutes the soft palate.
| |
| | |
| The second change in the Amniota, which also takes place in
| |
| some Amphibia, is caused by the section of the mesenteron into
| |
| which the branchial pouches open,
| |
| becoming, on the atrophy of these
| |
| structures, converted into the posterior part of the buccal cavity.
| |
| | |
| The organs derived from the
| |
| buccal cavity are the tongue, the
| |
| various salivary glands, and the
| |
| teeth ; but the latter alone will engage our attention here.
| |
| | |
| The teeth. The teeth are to be
| |
| regarded as a special product of the
| |
| oral mucous membrane. It has been
| |
| shewn by Gegenbaur and Hertwig
| |
| that in their mode of development
| |
| they essentially resemble the placoid
| |
| scales of Elasmobranchii, and that the latter structures extend
| |
| in Elasmobranchii for a certain distance into the cavity of the
| |
| mouth.
| |
| | |
| As pointed out by Gegenbaur, the teeth are therefore to be
| |
| regarded as more or less specialised placoid scales, whose
| |
| presence in the mouth is to be explained by the fact that the
| |
| latter structure is lined by an invagination of the epidermis.
| |
| The most important developmental point of difference between
| |
| teeth and placoid scales consists in the fact, that in the case
| |
| of the former there is a special ingrowth of epiblast to
| |
| meet a connective tissue papilla which is not found in the
| |
| latter.
| |
| | |
| | |
| | |
| FIG. 427. DIAGRAM SHEWING THE DIVISION OF THE PRIMITIVE BUCCAL CAVITY INTO THE
| |
| RESPIRATORY SECTION ABOVE
| |
| AND THE TRUE MOUTH BELOW.
| |
| (From Gegenbaur.)
| |
| | |
| p. palatine plate of superior
| |
| maxillary process; m. permanent
| |
| mouth ; n. posterior part of nasal
| |
| passage; e. internasal septum.
| |
| | |
| | |
| | |
| Although the teeth are to be regarded as primitively epiblastic structures, they are nevertheless found in Teleostei and Ganoidei on the hyoid
| |
| | |
| | |
| | |
| THE STOMOD/KUM.
| |
| | |
| | |
| | |
| 777
| |
| | |
| | |
| | |
| and branchial arches ; and very possibly the teeth on some other parts of
| |
| the mouth are developed in a true hypoblastic region.
| |
| | |
| The teeth are formed from two distinct organs, viz. an epithelial cap and
| |
| a connective tissue papilla.
| |
| | |
| The general mode of development, as has been more especially shewn
| |
| by the extended researches of Tomes, is practically the same for all Vertebrata, and it will be convenient to describe it as it takes place in Mammalia.
| |
| | |
| Along the line where the teeth are about to develop, there is formed
| |
| an epithelial ridge projecting into the subjacent connective tissue, and
| |
| derived from the innermost columnar layer of the oral epithelium. At the
| |
| points where a tooth is about to be formed this ridge undergoes special
| |
| changes. It becomes in the first place somewhat thickened by the development of a number of rounded cells in its interior ; so that it becomes
| |
| constituted of (i) an external layer of columnar cells, and (2) a central core
| |
| of rounded cells ; both of an epithelial nature. In the second place the
| |
| organ gradually assumes a dome-shaped form (fig. 428, e), and covers over a
| |
| papilla of the subepithelial connective tissue (p] which has in the meantime
| |
| been developed.
| |
| | |
| From the above epithelial structure, which may be called the enamel
| |
| organ, and from the papilla it covers, which
| |
| maybe spoken of as the dental papilla,
| |
| the whole tooth is developed. After these
| |
| parts have become established there is formed
| |
| round the rudiment of each tooth a special
| |
| connective tissue capsule ; known as the
| |
| dental capsule.
| |
| | |
| Before the dental capsule has become
| |
| definitely formed the enamel organ and the
| |
| dental papilla undergo important changes.
| |
| The rounded epithelial cells forming the core
| |
| of the enamel organ undergo a peculiar transformation into a tissue closely resembling
| |
| ordinary embryonic connective tissue, while
| |
| at the same time the epithelium adjoining
| |
| the dental papilla and covering the inner
| |
| surface of the enamel organ, acquires a somewhat different structure to the epithelium
| |
| on the outer side of the organ. Its cells
| |
| become very markedly columnar, and form
| |
| a very regular cylindrical epithelium. This
| |
| layer alone is concerned in forming the
| |
| enamel. The cells of the outer epithelial
| |
| layer of the enamel organ become somewhat
| |
| flattened, and the surface of the layer is raised into a series of short papilla?
| |
| which project into the highly vascular tissue of the dental sheath. Between
| |
| | |
| | |
| | |
| | |
| FIG. 428. DIAGRAM SHEWING THE DEVELOPMENT OF THE
| |
| TEETH. (From Gegenbaur.)
| |
| | |
| p. dental papilla ; e. enamel
| |
| organ.
| |
| | |
| | |
| | |
| 778 THE PROCTOD/EUM.
| |
| | |
| the epithelium of the enamel organ and the adjoining connective tissue
| |
| there is everywhere present a delicate membrane known as the membrana
| |
| praeformativa.
| |
| | |
| The dental papilla is formed of a highly vascular core and a non-vascular
| |
| superficial layer adjoining the inner epithelium of the enamel organ. The
| |
| cells of the superficial layer are arranged so as almost to resemble an
| |
| epithelium.
| |
| | |
| The first formation of the hard structures of the tooth commences at
| |
| the apex of the dental papilla. A calcification of the outermost layer of
| |
| the papilla sets in, and results in the formation of a thin layer of dentine.
| |
| Nearly simultaneously a thin layer of enamel is deposited over this,
| |
| from the inner epithelial layer of the enamel organ (fig. 428). Both
| |
| enamel and dentine continue to be deposited till the crown of the tooth has
| |
| reached its final form, and in the course of this process the enamel
| |
| organ is reduced to a thin layer, and the whole of the outer layer of the
| |
| dental papilla is transformed into dentine while the inner portion remains
| |
| as the pulp.
| |
| | |
| The root of the tooth is formed later than the crown, but the enamel
| |
| organ is not prolonged over this part, so that it is only formed of dentine.
| |
| | |
| By the formation of the root the crown of the tooth becomes pushed
| |
| outwards, and breaking through its sack projects freely on the surface.
| |
| | |
| The part of the sack which surrounds the root of the tooth gives rise
| |
| to the cement, and becomes itself converted into the periosteum of the
| |
| dental alveolus.
| |
| | |
| The general development of the enamel organs and dental papillae is
| |
| shewn in the diagram (fig. 428). From the epithelial ridge three enamel
| |
| organs are represented as being developed. Such an arrangement may
| |
| occur when teeth are successively replaced. The lowest and youngest
| |
| enamel organ (e) has assumed a cap-like form enveloping a dental papilla,
| |
| but no calcification has yet taken place.
| |
| | |
| In the next stage a cap of dentine has become formed, while in the
| |
| still older tooth this has become covered by a layer of enamel. As may be
| |
| gathered from this diagram, the primitive epithelial ridge from which the
| |
| enamel organ is formed is not necessarily absorbed on the formation of a
| |
| tooth, but is capable of giving rise to fresh enamel organs. When the
| |
| enamel organ has reached a certain stage of development, its connection
| |
| with the epithelial ridge is ruptured (fig. 428).
| |
| | |
| The arrangement represented in fig. 428, in which successive enamel
| |
| organs are formed from the same epithelial ridge, is found in most Vertebrata except the Teleostei. In the Teleostei, however (Tomes), a fresh
| |
| enamel organ grows inwards from the epithelium for each successively
| |
| formed tooth.
| |
| | |
| The Proctodceuni.
| |
| | |
| In all Vertebrata the cloacal section of the alimentary tract
| |
| which receives the urinogenital ducts is placed in communication
| |
| | |
| | |
| | |
| THE PROCTOD/EUM.
| |
| | |
| | |
| | |
| 779
| |
| | |
| | |
| | |
| with the exterior by means of an epiblastic invagination, constituting a proctodseum.
| |
| | |
| This invagination is not usually very deep, and in most
| |
| instances the boundary wall between it and the hypoblastic
| |
| cloaca is not perforated till considerably after the perforation of the
| |
| stomodseum ; in Petromyzon, however, its perforation is effected
| |
| before the mouth and pharynx are placed in communication.
| |
| | |
| The mode of formation of the proctodaeum, which is in
| |
| general extremely simple, is illustrated by fig. 420 an.
| |
| | |
| In most forms the original boundary between the cpiblast of
| |
| the proctodaeum and the hypoblast of the primitive cloaca
| |
| becomes obliterated after the two have become placed in free
| |
| communication.
| |
| | |
| | |
| | |
| | |
| FIG. 429. DIAGRAMMATIC LONGITUDINAL SECTION THROUGH THE POSTERIOR
| |
| END OF AN EMBRYO BlRD AT THE TIME OF THE FORMATION OF THE ALLANTOIS.
| |
| | |
| ep. epiblast ; Sp.c. spinal canal ; ch. notochord ; n.e. neurenteric canal ; hy, hypoblast ; p.a.g. postanal gut ; pr. remains of primitive streak folded in on the ventral
| |
| side ; al. allantois ; me. mesoblast ; an. point where anus will be formed ; p.c. perivisceral cavity ; am. amnion ; so. somatopleure ; sp. splanchnopleure.
| |
| | |
| In Birds the formation of the proctodseum is somewhat more complicated than in other types, owing to the outgrowth from it of the bursa
| |
| Fabricii.
| |
| | |
| The proctodseum first appears when the folding off of the tail end of
| |
| the embryo commences (fig. 429, an} and is placed near the front (originally
| |
| the apparent hind) end of the primitive streak. Its position marks out the
| |
| front border of the postanal section of the gut.
| |
| | |
| The bursa Fabricii first appears on the seventh day (in the chick), as a
| |
| dorsal outgrowth of the proctodaeum. The actual perforation of the septum between the proctodeeum and the cloacal section of the alimentary tract
| |
| is not effected till about the fifteenth day of fcetal life, and the approxi
| |
| | |
| | |
| 780 BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| mation of the epithelial layers of the two organs, preparatory to their
| |
| absorption, is partly effected by the tunneling of the mesoblastic tissue
| |
| between them by numerous spaces.
| |
| | |
| The hypoblastic section of the cloaca of birds, which receives the openings of the urinogenital ducts, is permanently marked off by a fold from
| |
| the epiblastic section or true proctodaeum, with which the bursa Fabricii
| |
| communicates.
| |
| | |
| BIBLIOGRAPHY.
| |
| Alimentary Canal and its appendages.
| |
| | |
| (561) B. Afanassiew. "Ueber Bau u. Entwicklung d. Thymus d. Saugeth."
| |
| Archivf. mikr. Anat. Bd. xiv. 1877.
| |
| | |
| (562) Fr. Boll. Das Princip d. Wachsthums. Berlin, 1876.
| |
| | |
| (563) E. Gasser. "Die Entstehung d. Cloakenoffnung bei Hiihnerembryonen."
| |
| Archivf. Anat. u. Physiol., Anat. Abth. 1880.
| |
| | |
| (564) A. Gotte. Beilrdge zur Entivicklungsgeschichle d. Darmkanah im
| |
| Hiihnchen. 1867.
| |
| | |
| (565) W. Millie r. "Ueber die Entwickelung der Schilddriise." Jenaische
| |
| Zeitschrift, Vol. vi. 1871.
| |
| | |
| (566) W. Miiller. "Die Hypobranchialrinne d. Tunicaten." Jenaische Zeitschrift, Vol. VII. 1872.
| |
| | |
| (567) S. L. Schenk. "Die Bauchspeicheldriise d. Embryo." Anatomischphysiologische Untcrsuchungen. 1872.
| |
| | |
| (568) E. Selenka. " Beitrag zur Entwicklungsgeschichte d. Luftsacke d.
| |
| Huhns." Zeit.f. wiss. Zool. 1866.
| |
| | |
| (569) L. Stieda. Untersuch. iib. d. Entwick. d. Glandula Thymus, Glandula
| |
| thyroidea,u. Glandula car otica. Leipzig, 1881.
| |
| | |
| (570) C. Fr. Wolff. " De formatione intestinorum." Nov. Comment. Akad.
| |
| Petrop. 1766.
| |
| | |
| (571) H. Wolfler. Ueb. d. Entwick. u. d. Bau d. Schilddriise. Berlin, 1880.
| |
| Vide also Kolliker (298), Gotte (296), His (232 and 297), Foster and Balfour (295),
| |
| | |
| Balfour (292), Remak (302), Schenk (303), etc.
| |
| | |
| Teeth.
| |
| | |
| (572) T. H. Huxley. "On the enamel and dentine of teeth." Quart. J. of
| |
| Micros. Science, Vol. in. 1855.
| |
| | |
| (573) R. Owen. Odontography . London, 1840 1845.
| |
| | |
| (574) Ch. S. Tomes. Manual of dental anatomy, human and comparative.
| |
| London, 1876.
| |
| | |
| (575) Ch. S. Tomes. " On the development of teeth." Quart. J. of Micros.
| |
| Science, Vol. xvi. 1876.
| |
| | |
| (576) W. Waldeyer. " Structure and development of teeth." Strieker's Histology. 1870.
| |
| | |
| Vide also Kolliker (298), Gegenbaur (294), Hertwig (306), etc.
| |
| | |
| | |
| | |
| INDEX TO VOLUME III.
| |
| | |
| | |
| | |
| Abdominal muscles, 675
| |
| | |
| Abdominal pore, 626, 749
| |
| | |
| Acipenser, development of, 102; affinities
| |
| of, 1 1 8 ; comparison of gastrula of, 279 ;
| |
| pericardial cavity of, 627
| |
| | |
| Actinotrocha, 373
| |
| | |
| Air-bladder of Teleostei, 77; Lepidosteus,
| |
| 117; blood supply of, 645 ; general account of, 763 ; homologies of, 766
| |
| | |
| Alciope, eye of, 480
| |
| | |
| Alisphenoid region of skull, 569
| |
| | |
| Alimentary canal and appendages, development of, 754
| |
| | |
| Alimentary tract ofAscidia, 18; Molgula,
| |
| 22; Pyrosoma, 24; Salpa, 31 ; Elasmobranchii, 52; Teleostei, 75; Petromyzon, 93, 97; Acipenser, no; Amphibia, 129, 136; Chick, 167; respiratory
| |
| region of, 754; temporary closure of
| |
| oesophageal region of, 759
| |
| | |
| Allantois, development of in Chick, 191,
| |
| 198; blood-vessels of in Chick, 193;
| |
| Lacerta, 205, 209; early development of
| |
| in Rabbit, 229, of Guinea-pig, 264;
| |
| origin of, 309. See also ' Placenta ' and
| |
| 'Bladder''
| |
| | |
| Alternation of generations in Ascidians,
| |
| origin of, 35 ; in Botryllus, 35 ; Pyrosoma, 36; Salpa, 36; Doliolum, 36
| |
| | |
| Alytes, branchial chamber of, 136; yolksack of, 139; branchiae, 141 ; Miillerian
| |
| duct of, 710
| |
| | |
| Amblystoma, ovum of, 120; larva of, 142,
| |
| | |
| H3
| |
| | |
| Amia, ribs of, 561
| |
| | |
| Ammocoetes, 95; metamorphosis of, 97;
| |
| | |
| eye of, 498
| |
| Amnion, early development of in Chick,
| |
| | |
| 185; later history of in Chick, 196;
| |
| | |
| Lacerta, 204, 210; Rabbit, 229; origin
| |
| | |
| of, 3.07. 39
| |
| | |
| Amphibia, development of, 120; viviparous, 121; gastrula of, 277; suctorial
| |
| mouth of, 317; cerebellum of, 426; infundibulum of, 431; pineal gland of,
| |
| 433; cerebrum of, 439; olfactory lobes
| |
| of, 444; nares of, 553; notochord and
| |
| its sheath, 548; vertebral column of,
| |
| 554; ribs of, 561 ; branchial arches of,
| |
| 574; mandibular and hyoid arches of,
| |
| 582 ; columella of, 582 ; pectoral girdle
| |
| of, 605; pelvic girdle of, 607; limbs of,
| |
| 619; heart of, 638; arterial system of,
| |
| f>45 ; venous system of, 655 ; excretory
| |
| | |
| | |
| | |
| system of, 707 ; vasa efierentia of, 711;
| |
| liver of, 769; postanal gut of, 774;
| |
| stomodaeum of, 778
| |
| | |
| Amphiblastula larva of Porifera, 344
| |
| | |
| Amphioxus, development of, i ; gastrula
| |
| of, 275 ; formation of mesoblast of, 292 ;
| |
| development of notochord of, 293; head
| |
| of, 314; spinal nerves of, 461; olfactory organ of, 462 ; venous system
| |
| of, 651; transverse abdominal muscle
| |
| f> 673; generative cells of, 748; liver
| |
| of, 769; postanal gut of, 772; stomodaeum of, 777
| |
| | |
| Amphistylic skulls, 578
| |
| | |
| Angular bone, 594
| |
| | |
| Anterior abdominal vein, 653
| |
| | |
| Anura, development of, 121; epiblast of,
| |
| 125; mesoblast of, 128; notochord of,
| |
| 128; hypoblast of, 129; general growth
| |
| of embryo of, 131; larva of, 134; vertebral column of, 556 ; mandibular arch
| |
| of, 584
| |
| | |
| Anus of Amphioxus, 7 ; Ascidia, 18; Pyrosoma, 28 ; Salpa, 31 ; Elasmobranchii,
| |
| 57; Amphibia, 130, 132; Chick, 167;
| |
| primitive, 324
| |
| | |
| Appendicularia, development of, 34
| |
| | |
| Aqueductus vestibuli, 519
| |
| | |
| Aqueous humour, 497
| |
| | |
| Arachnida, nervous system of, 409; eye
| |
| of, 481
| |
| | |
| Area, embryonic, of Rabbit, 218; epiblast
| |
| | |
| of, 219; origin of embryo from, 228
| |
| | |
| area opaca of Chick, 150; epiblast,
| |
| | |
| hypoblast, and mesoblast of, 159
| |
| area pellucida of Chick, 150; of Lacerta, 202
| |
| | |
| area vasculosa of Chick, 194; mesoblast of, 1 60; of Lizard, 209; Rabbit,
| |
| 228, 229
| |
| | |
| Arteria centralis retinas, 503
| |
| | |
| Arterial system of Petromyzon, 97; constitution of in embryo, 643 ; of Fishes,
| |
| 644; of Amphibia, 645; of Amniota, 647
| |
| | |
| Arthropoda, head of, 313 ; nervous system
| |
| of, 409 ; eye of, 480 ; excretory organs
| |
| of, 688
| |
| | |
| Articular bone of Teleostei, 581 ; of Sauropsida, 588
| |
| | |
| Ascidia, development of, 9
| |
| | |
| Ascidians. See 'Tunicata'
| |
| | |
| Ascidiozooids, 25
| |
| | |
| Atrial cavity of Amphioxus, 7; Ascidia,
| |
| 18; Pyrosoma, 24
| |
| | |
| | |
| | |
| 7 82
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| Atrial pore of Amphioxus, 7; Ascidia, 20;
| |
| Pyrosoma, 28 ; Salpa, 32
| |
| | |
| Auditory capsules, ossifications in, 595,
| |
| 59.6
| |
| | |
| Auditory involution of Elasmobranchii,
| |
| 57; Teleostei, 73; Petromyzon, 89,
| |
| 92; Acipenser, 106; Lepidosteus, 114;
| |
| Amphibia, 127; Chick, 170
| |
| | |
| Auditory nerve, development of, 459
| |
| | |
| Auditory organs, of Ascidia, 15; of Salpa,
| |
| 31; of Ammocoetes, 98; Ganoidei, 108,
| |
| 114; of Amphibia, 127; of Aves, 170;
| |
| general development of, 512; of aquatic
| |
| forms, 512; of land forms, 513; of
| |
| Ccelenterata, 513; of Mollusca, 515;
| |
| of Crustacea, 516; of Vertebrata, 517;
| |
| of Cyclostomata, 89, 92, 518; of Teleostei, Lepidosteus and Amphibia,
| |
| 518; of Mammalia, 519; accessory
| |
| structures of, 527; ofTunicata, 528
| |
| | |
| Auriculo-ventricular valves, 642
| |
| | |
| Autostylic skulls, 579
| |
| | |
| Aves, development of, 145; cerebellum
| |
| of, 426; midbrain of, 427; infundibulum of, 431; pineal gland of, 434;
| |
| pituitary body of, 436; cerebrum of,
| |
| 439 ; olfactory lobes of, 444 ; spinal
| |
| nerves of, 449 ; cranial nerves of, 455 ;
| |
| vagus of, 458; glossopharyngeal of,
| |
| 458; vertebral column of, 557; ossification of vertebral column of, 558;
| |
| branchial arches of, 572, 573; pectoral
| |
| girdle of, 603; pelvic girdle of, 608;
| |
| heart of, 637 ; arterial system of, 647 ;
| |
| venous system of, 658; muscle-plates
| |
| of, 670; excretory organs of, 714; mesonephros of, 715; pronephros of, 718;
| |
| Miillerian duct of, 718, 720; nature of
| |
| pronephros of, 721 ; connection of Miillerian duct with Wolffian in, 720 ;
| |
| kidney of, 722; lungs of, 764; liver of,
| |
| 769; postanal gut of, 774
| |
| | |
| Axolotl, 142, 143; ovum of, 120; midbrain of, 427; mandibular arch of, 583
| |
| | |
| Basilar membrane, 524
| |
| | |
| Basilar plate, 565
| |
| | |
| Basipterygium, 612
| |
| | |
| Basisphenoid region of skull, 569
| |
| | |
| Bilateral symmetry, origin of, 373-376
| |
| | |
| Bile duct, 770
| |
| | |
| Bladder, Amphibia, 131 ; of Amniota, 726
| |
| | |
| Blastodermic vesicle, of Rabbit, first development of, 217; of 7th day, 222;
| |
| Guinea-pig, 263; meaning of, 291
| |
| | |
| Blastoderm of Pyrosoma, 24; Elasmobranchii, 41; Chick, 150; Lacerta 202
| |
| | |
| Blastopore, of Amphioxus, 2; of Ascidia,
| |
| II ; Elasmobranchii, 42, 54, 62 ; Petromyzon, 87; Acipenser, 104 ; Amphibia,
| |
| 125, 130; Chick, 153; Rabbit, 216;
| |
| true Mammalian, 226; comparative
| |
| history of closure of, 284, 288; summary of fate of, 340; relation of to
| |
| primitive anus, 324
| |
| | |
| | |
| | |
| Blood-vessels, development of, 633
| |
| | |
| Body cavity, of Ascidia, 2 1 ; Molgula, 2 1 ;
| |
| Salpa, 31; Elasmobranchii, 47 ; of Teleostei, 75 ; Petromyzon, 94 ; Chick,
| |
| 169; development of in Chordata, 325;
| |
| views on origin of, 356 360, 377; of
| |
| Invertebrata, 623; of Chordata, 624;
| |
| of head, 676
| |
| | |
| Bombinator, branchial chamber of, 136;
| |
| vertebral column of, 556
| |
| | |
| Bonellia, excretory organs of, 687
| |
| | |
| Bones, origin of cartilage bones, 542 ;
| |
| origin of membrane bones, 543; development of, 543; homologies of membrane bones, 542 ; homologies of cartilage bones, 545
| |
| | |
| Brachiopoda, excretory organs of, 683 ;
| |
| generative ducts of, 749
| |
| | |
| Brain, of Ascidia, IT, 15; Elasmobranchii, 56, 59, 60; Teleostei, 77; Petromyzon, 89, 92 ; Acipenser, 105 ; Lepidosteus, 113; early development of in
| |
| Chick, 170; flexure of in Chick, 175;
| |
| later development of in Chick, 176;
| |
| Rabbit, 229, general account of development of, 419; flexureof, 420; histogeny of, 422
| |
| | |
| Branchial arches, prseoral, 570; disappearance of posterior, 573; dental plates
| |
| of in Teleostei, 574; relation of to
| |
| head cavities, 571 ; see ' Visceral arches'
| |
| | |
| Branchial chamber of Amphibia, 136
| |
| | |
| Branchial clefts, of Amphioxus, 7 ; of
| |
| Ascidia, 18, 20; Molgula, 23; Salpa,
| |
| 32; of Elasmobranchii, 57, 59 01;
| |
| Teleostei, 77; Petromyzon, 91, 96;
| |
| Acipenser, 105; Lepidosteus, 114, 116;
| |
| Amphibia, 132, 133; Chick, 178;
| |
| Rabbit, 231; praeoral, 312, 318; of
| |
| Invertebrata, 326; origin of, 326
| |
| | |
| Branchial rays, 574
| |
| | |
| Branchial skeleton, development of, 572,
| |
| 592; of Petromyzon, 96, 312, 571; of
| |
| Ichthyopsida, 572; dental plates of in
| |
| Teleostei, 574; relation of to head
| |
| cavities, 572
| |
| | |
| Branchiae, external of Elasmobranchii, 6r,
| |
| 62; of Teleostei, 77; Acipenser, 107;
| |
| Amphibia, 127, 133, 135
| |
| | |
| Brood-pouch, of Salpa, 29 ; Teleostei, 68,
| |
| Amphibia, 12 1
| |
| | |
| Brown tubes of Gephyrea, 686
| |
| | |
| Bulbus arteriosus, of Pishes, 638 ; Amphibia, 639
| |
| | |
| Bursa Fabricii, 167, 779
| |
| | |
| Canalis auricularis, 639
| |
| Canalis reuniens, 521
| |
| Capitellidre, excretory organs of, 683
| |
| Carcharias, placenta of, 66
| |
| Cardinal vein, 652
| |
| Carnivora, placenta of, 250
| |
| Carpus, development of, 620
| |
| Cartilage bones of skull, 595 ; homologies
| |
| of, 595
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| 783
| |
| | |
| | |
| | |
| Cat, placenta of, 250
| |
| | |
| Caudal swellings of Elasmobranchii, 46,
| |
| | |
| 55; Teleostei, 72; Chick, 162, 170
| |
| Cephalic plate of Elasmobranchii, 55
| |
| Cephalochorda, development of, i
| |
| Cephalopoda, eyes of, 473 477
| |
| Cerebellum, Petromyzon, 93; Chick, 176;
| |
| | |
| general account of development of, 424,
| |
| | |
| 425
| |
| | |
| Cerebrum of Petromyzon, 93, 97; Chick,
| |
| 175 ; general development of, 429, 438;
| |
| transverse fissure of, 443
| |
| Cestoda, excretory organs of, 68 1
| |
| Cetacea, placenta, 255
| |
| Chtetognatha, nervous system of, 349;
| |
| eye of, 479 ; generative organs of, 743 ;
| |
| generative ducts of, 749
| |
| Chcetopoda, head of, 313; eyes of, 479;
| |
| excretory organs of, 683; generative
| |
| organs of, 743 ; generative ducts of, 749
| |
| Charybdnea, eye of, 472
| |
| Cheiroptera, placenta of, 244
| |
| Cheiropterygium, 618; relation of to ich
| |
| thyopterygium, 621
| |
| | |
| Chelonia, development of, 210; pectoral
| |
| girdle of, 603 ; arterial system of, 649
| |
| Chick, development of, 145 ; general
| |
| growth of embryo of, 1 70 ; rotation of
| |
| embryo of, 173; fcetal membranes of,
| |
| 185; epiblast of, 150, 166; optic nerve
| |
| and choroid fissure of, 500
| |
| | |
| Chilognatha, eye of, 481
| |
| | |
| Chilopoda, eye of, 481
| |
| | |
| Chimasra, lateral line of, 539 ; vertebral
| |
| column of, 548; nares of, 533
| |
| | |
| Chiromantis, oviposition of, 121
| |
| | |
| Chorda tympani, development of, 460
| |
| | |
| Chordata, ancestor of, 311; branchial
| |
| system of, 312; evidence from Ammocuetes, 312; head of, 312; mouth of,
| |
| 318; table of phylogeny of, 327
| |
| | |
| Chorion, 237; villi of, 237, 257
| |
| | |
| Choroid coat, Ammoccetes, 99; general
| |
| account of, 487
| |
| | |
| Choroid fissure, of Vertebrate eye, 486,
| |
| 493 ; of Ammocoetes, 498 ; comparative
| |
| development of, 500; of Chick, 501;
| |
| of Lizards, 501 ; of Elasmobranchii,
| |
| 502 ; of Teleostei, 503 ; Amphibia, 503 ;
| |
| Mammals, 503, 504
| |
| | |
| Choroid gland, 320
| |
| | |
| Choroid pigment, 489
| |
| | |
| Choroid plexus, of fourth ventricle, 425 ;
| |
| of third ventricle, 432 ; of lateral ventricle, 442
| |
| | |
| Ciliated sack of Ascidia, 18; Pyrosoma,
| |
| 26; Salpa, 31
| |
| | |
| Ciliary ganglion, 461
| |
| | |
| Ciliary muscle, 490
| |
| | |
| Ciliary processes, 488; comparative development of, 506
| |
| | |
| Clavicle, 600
| |
| | |
| Clitoris, development of, 727
| |
| | |
| Clinoid ridge, 569
| |
| | |
| Cloaca, 766
| |
| | |
| | |
| | |
| Coccygeo-mesenteric vein, 66 1
| |
| | |
| Cochlear canal, 519
| |
| | |
| Coecilia, development of, 143; pronephros
| |
| of, 707; mesonephros of, 709; Mill
| |
| lerian duct of, 710
| |
| | |
| Coelenterata, larvae of, 367 ; eyes of, 47 1 ;
| |
| auditory organs of, 513; generative
| |
| organs of, 741
| |
| | |
| Columella auris, 529; of Amphibia, 582 ;
| |
| of Sauropsida, 588
| |
| | |
| Commissures, of spinal cord, 417; of
| |
| brain, 431, 432, 439, 443
| |
| | |
| Coni vasculosi, 724
| |
| | |
| Conus arteriosus, of Fishes, 638; of Amphibia, 638
| |
| | |
| Coracoid bone, 599
| |
| | |
| Cornea, of Ammocretes, 99 ; general development of, 495 ; corpuscles of, 496 ;
| |
| comparative development of, 499; of
| |
| Mammals, 499
| |
| | |
| Coronoid bone, 595
| |
| | |
| Corpora geniculata interna, 428
| |
| | |
| Corpora quadrigemina, 428
| |
| | |
| Corpora striata, development of, 437
| |
| | |
| Corpus callosum, development of, 443
| |
| | |
| Corti, organ of, 522; structure of, 525;
| |
| fibres of, 525 ; development of, 526
| |
| | |
| Cranial flexure, of Elasmobranchii, 58,
| |
| 60; of Teleostei, 77; Petromyzon, 93,
| |
| 94; of Amphibia, 131, 132; Chick,
| |
| 174; Rabbit, 231; characters of, 321;
| |
| significance of, 322
| |
| | |
| Cranial nerves, development of, 455;
| |
| relation of to head cavities, 461 ; anterior roots of, 462 464; view on
| |
| position of roots of, 466
| |
| | |
| Crocodilia, arterial system of, 649
| |
| | |
| Crura cerebri, 429
| |
| | |
| Crustacea, nervous system of, 41 1 ; eye of,
| |
| 481; auditory organs of, 515; generative cells of, 745 ; generative ducts of,
| |
| | |
| 75
| |
| | |
| Cupola, 524
| |
| | |
| Cutaneous muscles, 676
| |
| | |
| Cyathozooid, 25
| |
| | |
| Cyclostomata, auditory organs of, 517;
| |
| olfactory organ of, 532; notochord and
| |
| vertebral column of, 546, 549; abdominal pores of, 626 ; segmental duct of,
| |
| 700 ; pronephros of, 700 ; mesonephros
| |
| of, 700 ; generative ducts of, 733, 749 ;
| |
| venous system of, 651 ; excretory organs
| |
| of, 700
| |
| | |
| Cystignathus, oviposition of, 122
| |
| | |
| Dactylethra, branchial chamber of, 136;
| |
| | |
| branchise of, 136; tadpole of, 140
| |
| Decidua reflexa, of Rat, 242 ; of Insecti
| |
| vora, 243; of Man, 245
| |
| Deiter's cells, 526
| |
| Dental papilla, 777
| |
| Dental capsule, 777
| |
| Dentary bone, 595
| |
| Dentine, 780
| |
| Descemet's membrane, 496
| |
| | |
| | |
| | |
| 784
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| Diaphragm, 631 ; muscle of, 676
| |
| | |
| Dipnoi, nares of, 534; vertebral column
| |
| of, 548; membrane bones of skull of,
| |
| 592 ; heart of, 638 ; arterial system of,
| |
| 645 ; excretory system of, 707 ; stomodseum of, 777
| |
| | |
| Diptera, eye of, 481
| |
| | |
| Discophora, excretory organs of, 687
| |
| | |
| Dog, placenta of, 248
| |
| | |
| Dohni, on relations of Cyclostomata, 84 ;
| |
| on ancestor of Chordata, 311, 319
| |
| | |
| Doliolum, development of, 28
| |
| | |
| Ductus arteriosus, 649
| |
| | |
| Ductus Botalli, 648
| |
| | |
| Ductus Cuvieri, 654
| |
| | |
| Ductus venosus Arantii, 663
| |
| | |
| Dugong, heart of, 642
| |
| | |
| Dysticus, eye of, 481
| |
| | |
| Ear, see ' Auditory organ '
| |
| | |
| Echinodermata, secondary symmetry of
| |
| larva of, 380; excretory organs of, 689 ;
| |
| generative ducts of, 752
| |
| | |
| Echinorhinus, lateral line of, 539; vertebral column of, 548
| |
| | |
| Echiurus, excretory organs of, 686
| |
| | |
| Ectostosis, 543
| |
| | |
| Edentata, placenta of, 248, 250, 256
| |
| | |
| Eel, generative ducts of, 703
| |
| | |
| Egg-shell of Elasmobranchii, 40 ; Chick,
| |
| 146
| |
| | |
| Elasmobranchii, development of, 40; viviparous, 40; general features of development of, 55 ; gastrulaof, 281 ; development of mesoblast of, 294 ; notochord of, 294 ; meaning of formation of
| |
| mesoblast of, 295; restiform tracts of,
| |
| 425 ; optic lobes of, 427 ; cerebellum of,
| |
| 425 ; pineal gland of, 432 ; pituitary
| |
| body of, 435 ; cerebrum of, 438 ; olfactory lobes of, 444 ; spinal nerves, 449 ;
| |
| cranial nerves of, 457; sympathetic
| |
| nervous system of, 466; nares of, 533;
| |
| lateral line of, 539; vertebral column of,
| |
| 549 ; ribs of, 560 ; parachordals of, 567 ;
| |
| mandibular and hyoid arches of, 576 ;
| |
| pectoral girdle of, 600 ; pelvic girdle of,
| |
| 607; limbs of, 609; pericardial cavity
| |
| of, 627; arterial system of, 644 ; venous
| |
| system of, 65 1 ; muscle-plates of, 668 ;
| |
| excretory organs of, 690 ; constitution
| |
| of excretory organs in adult of, 697;
| |
| spermatozoa of, 747 ; swimming-bladder of, 763 ; intestines of, 767 ; liver of,
| |
| 769; postanal gut of, 772
| |
| | |
| Elrcoblast of Pyrosoma, 28; Salpa, 30
| |
| | |
| Elephant, placenta of, 249
| |
| | |
| Embolic formation of gastrula, 333
| |
| | |
| Enamel organ, 777
| |
| | |
| Endolymph of ear, 522
| |
| | |
| Endostosis, 543
| |
| | |
| Endostyle of Ascidia, 18, 759; Pyrosoma,
| |
| 25; Salpa, 32
| |
| | |
| Epiblast, of Elasmobranchii, 47 ; Teleostei, 71, 75; Petromyzon, 86; Lcpid
| |
| | |
| | |
| osteus, 112; Amphibia, 122, 125;
| |
| Chick, 149, 166; Lacerta, 203; Rabbit,
| |
| 216, 219; origin of in Rabbit, 221 ;
| |
| comparative account of development
| |
| of, 300
| |
| | |
| Epibolic formation of gastrula, 334
| |
| | |
| Epichordal formation of vertebral column,
| |
| 556
| |
| | |
| Epicrium glutinosum, 143
| |
| | |
| Epidermis, in Ccelenterata, 393; protective structures of, 394
| |
| | |
| Epididymis, 724
| |
| | |
| Epigastric vein, 653
| |
| | |
| Episkeletal muscles, 676
| |
| | |
| Episternum, 602
| |
| | |
| Epoophoron, 725
| |
| | |
| Ethmoid bone, 597
| |
| | |
| Ethmoid region of skull, 570
| |
| | |
| Ethmopalatine ligament of Elasmobranchs, 576
| |
| | |
| Euphausia, eye of, 483
| |
| | |
| Eustachian tube, of Amphibia, 135;
| |
| Chick, 1 80; Rabbit, 232; general
| |
| development of, 528
| |
| | |
| Excretory organs, general constitution of,
| |
| 680; of Platyelminthes, 680; of Mollusca, 681; of Polyzoa, 682; of Brachiopoda, 683 ; of Choetopoda, 683 ; of
| |
| Gephyrea, 686 ; of Discophora, 687 ; of
| |
| Arthropoda, 688; of Nematoda, 689;
| |
| of Echinodermata, 689 ; constitution of
| |
| in Craniata, 689; of Elasmobranchii,
| |
| 690; constitution of in adult Elasmobranch, 697; of Petromyzon, 700; of
| |
| Myxine, 701 ; of Teleostei, 701 ; of
| |
| Ganoidei, 704; of Dipnoi, 707; of
| |
| Amphibia, 707; of Amniota, 713;
| |
| comparison of Vertebrate and Invertebrate, 737
| |
| | |
| Excretory system, of Elasmobranchii, 49 ;
| |
| Teleostei, 78; Petromyzon, 95, 98;
| |
| Acipenser, 99; Amphibia, 133
| |
| | |
| Exoccipital bone, 595
| |
| | |
| Exoskeleton, dermal, 393 395 ; epidermal, 393396
| |
| | |
| External generative organs, 726
| |
| | |
| Extra-branchial skeleton, 572
| |
| | |
| Eye, of Ascidia, 16; Salpa, 31; Elasmobranchii, 56, 57, 58; Teleostei, 73;
| |
| Petromyzon, 92, 98; Aves, i/o; Rabbit, 229; general development of, 470;
| |
| evolution of, 470, 471; simple, 480;
| |
| compound, 481 ; aconous, 482; pseudoconous, 482 ; of Invertebrata, 471; of
| |
| Vertebrata, 483 ; comparative development of Vertebrate, 497 ; of Ammoccetes, 497 ; of Tunicata, 507 ; of Chordata, general views on, 508 ; accessory
| |
| eyes of Fishes, 509; muscles of, 677
| |
| | |
| Eyelids, development of, 506
| |
| | |
| Falciform ligament, 757
| |
| | |
| Falx cerebri, 439
| |
| | |
| Fasciculi terctes, of Elasmobranchii. 426
| |
| | |
| Feathers, development of, 396
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| 785
| |
| | |
| | |
| | |
| Fenestra rotunda and ovalis, 529
| |
| | |
| Fertilization, of Amphioxus, 2 ; of Urochorda, 9; Salpa, 29; Elasmobranchii,
| |
| 46; of Teleostei, 68; Petromyzon, 84 ;
| |
| Amphibia, 120; Chick, 145 ; Reptilia,
| |
| 202 ; meaning of, 331
| |
| | |
| Fifth nerve, development of, 460
| |
| | |
| Fifth ventricle, 443
| |
| | |
| Fins, of Elasmobranchii, 62 ; Teleostei,
| |
| 78; Petromyzon, 94, 95; Acipenser,
| |
| 109; Lepidosteus, 118; relation of
| |
| paired to unpaired, 611, 612 ; development of pelvic, 614; development of
| |
| pectoral, 615; views on nature of paired
| |
| fins, 616
| |
| | |
| Fissures of spinal cord, 417
| |
| | |
| Foetal development, 360 ; secondary variations in, 361
| |
| | |
| Foot, 618
| |
| | |
| Foramen of Munro, 430, 438
| |
| | |
| Foramen ovale, 642
| |
| | |
| Forebrain, of Elasmobranchii, 55, 59, 60;
| |
| Petromyzon, 93 ; general development
| |
| of, 428
| |
| | |
| Formative cells, of Chick, 154
| |
| | |
| Fornix, development of, 443
| |
| | |
| Fornix of Gottsche, 428
| |
| | |
| Fourth nerve, 464
| |
| | |
| Frontals, 592
| |
| | |
| Fronto-nasal process of Chick, 179
| |
| | |
| Gaertner's canals, 724
| |
| | |
| Gall-bladder, 770
| |
| | |
| Ganoidei, development of, 102; relations
| |
| of, 118; nares of, 534; notochord of,
| |
| 546 ; vertebral column of, 546, 553 ;
| |
| ribs of, 561 ; pelvic girdle of, 606; arterial system of, 645 ; excretory organs
| |
| of, 704; generative ducts of, 734
| |
| | |
| Gastropoda, eye of, 472
| |
| | |
| Gastrula, of Amphioxus, 2; of Ascidia, lo;
| |
| Elasmobranchii, 43, 44 ; Petromyzon,
| |
| 86; Acipenser, 103; Amphibia, 123;
| |
| comparative development of, in Invertebrata, 275 ; comparison of Mammalian, 291 ; phylogenetic meaning of, 333 ;
| |
| ontogeny of (general), 333 ; phylogeny
| |
| of, 338 343 ; secondary types of, 34!
| |
| | |
| Geckos, vertebral column of, 557
| |
| | |
| Generative cells, development of, 74! ;
| |
| origin of in Ccelenterata, 741 ; of Invertebrata, 743 ; of Vertebrata, 746
| |
| | |
| Generative ducts, of Teleostei, 704, 735 ;
| |
| of Ganoids, 704; of Cyclostomata, 733;
| |
| origin of, 733 ; of Lepidosteus, 735,
| |
| 750 ; development and evolution of,
| |
| 748 ; of Ccelenterata, 748 ; of Sagitta,
| |
| 749 ; of Tunicata, 749 ; Cheetopoda,
| |
| Gephyrea, etc., 749; of Mollusca, 751;
| |
| of Discophora, 751 ; of Echinodermata,
| |
| | |
| 75*
| |
| | |
| Generative system of Elasmobranchii, 51
| |
| Gephyrea, nervous system of, 412; excretory organs of, 686 ; generative cells of,
| |
| 743 ; generative ducts of, 749
| |
| | |
| B. III.
| |
| | |
| | |
| | |
| Germinal disc, of Elasmobranchii, 40;
| |
| Teleostei, 68 ; Chick, 147
| |
| | |
| Germinal epithelium, 746
| |
| | |
| Germinal layers, summary of organs <lrrived from, in Vertebrata, 304 ; historical account of views of, 332 ; homologies of in the Metazoa, 345
| |
| | |
| Germinal wall of Chick, 152, 159; structure and changes of, 160
| |
| | |
| Geryonia, auditory organ of, 5 r 5
| |
| | |
| Gill of Salpa, 31
| |
| | |
| Giraldes, organ of, 725
| |
| | |
| Glands, epidermic, development of, 397
| |
| | |
| Glomerulus, external, of Chick, 716
| |
| | |
| Glossopharyngeal nerve, development of,
| |
| | |
| 45 6 > 457
| |
| Grey matter of spinal cord, 417; of brain,
| |
| | |
| 423
| |
| Growth in length of Vertebrate embryo,
| |
| | |
| 306
| |
| Guinea-pig, primitive streak of, 223;
| |
| | |
| notochord of, 226 ; placenta of, 242 ;
| |
| | |
| development of, 262
| |
| Gymnophiona, see ' Ccecilia '
| |
| | |
| Habenula perforata, 525
| |
| | |
| Hairs, development of, 396
| |
| | |
| Halichrerus, placenta of, 250
| |
| | |
| Hand, 619
| |
| | |
| Head, comparative account of, 313; segmentation of, 314
| |
| | |
| Head cavities, of Elasmobranchii, 50 ;
| |
| Petromyzon, 90, 96; Amphibia, 127;
| |
| general development of, 676
| |
| | |
| Head-fold of Chick, 157, 167
| |
| | |
| Head kidney, see ' Pronephros '
| |
| | |
| Heart, of Pyrosoma, 25; Elasmobranchii,
| |
| 50, 58 ; Petromyzon, 94, 97 ; Acipenser, 106; Chick, 170 ; first appearance
| |
| of in Rabbit, 230; general development
| |
| of, 633 ; of Fishes, 635, 637 ; of Mammalia, 638; of Birds, 637, 639; meaning of development of, 637 ; of Amphibia, 638 ; of Amniota, 639 ; change of
| |
| position of, 643
| |
| | |
| Hind-brain, Elasmobranchii, 55, 59, 60 ;
| |
| Petromyzon, 93 ; general account of,
| |
| 424
| |
| | |
| Hippocampus major, development of, 442
| |
| | |
| Hirudo, development of blood-vessels of,
| |
| 633 ; excretory organs of, 688
| |
| | |
| Horse, placenta of, 253
| |
| | |
| Hyaloid membrane, 492
| |
| | |
| Hylodes, oviposition of, 1 21 ; metamorphosis of, -1 37
| |
| | |
| Hyobranchial cleft, 572
| |
| | |
| Hyoid arch, of Chick, 179; general account of, 572, 575 ; modifications of,
| |
| e !73> 577 > f Elasmobranchii, 576; of
| |
| Teleostei, 577 ; of Amphibia, 582 ;
| |
| of Sauropsida, 588; of Mammalia,
| |
| | |
| 589
| |
| | |
| Hyomandibular bar of Elasmobranchii,
| |
| 576, 577 ; of Teleostei, 579 ; of Amphibia, 582
| |
| | |
| 50
| |
| | |
| | |
| | |
| ;86
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| Hyomandibular cleft, of Fetromyzon, 91 ;
| |
| Chick, 179 ; general account of, 572
| |
| | |
| Hyostylic skulls, 582
| |
| | |
| Hypoblast of Elasmobranchii, 5! ; Teleostei, 71, 75; Petromyzon, 86; Acipenser, 104; Lepidosteus, 113; Amphibia,
| |
| 122, 129; Chick, 151, 167 ; Lacerta,
| |
| 203; Rabbit, 215, 216, 219 ; origin of
| |
| in Rabbit, 220
| |
| | |
| Hyposkeletal muscles, 675
| |
| | |
| Ilyrax, placenta of, 249
| |
| | |
| Incus, 529, 590
| |
| | |
| Infraclavicle, 600
| |
| | |
| Infundibulum of Petromyzon, 92 ; Chick,
| |
| 175 ; general development of, 430
| |
| | |
| Insectivora, placenta of, 243
| |
| | |
| Insects, nervous system of, 410 ; eye of,
| |
| 481; generative organs of, 745; generative ducts of, 751
| |
| | |
| Intercalated pieces of vertebral column,
| |
| | |
| 55 1
| |
| | |
| Interclavicle, homologies of, 602
| |
| | |
| Intermediate cell-mass of Chick, 183
| |
| | |
| Intermuscular septa, 672
| |
| | |
| Interorbital septum, 570
| |
| | |
| Interrenal bodies, 665
| |
| | |
| Iris, 489 ; comparative development of,
| |
| | |
| 506
| |
| | |
| Iris of Ammoccetes, 98
| |
| Island of Reil, 444
| |
| | |
| Jacobson's organ, 537
| |
| Jugal bone, 594
| |
| | |
| Kidney, see ' Metanephros '
| |
| | |
| Labia majora, development of, 727
| |
| | |
| Labial cartilages, 597
| |
| | |
| Labium tympanicum, 525 ; vestibulare,
| |
| | |
| 5 2 5
| |
| | |
| Lacertilia, general development of, 202 ;
| |
| nares of, 537 ; pectoral girdle of, 603 ;
| |
| pelvic girdle of, 607 ; arterial system
| |
| of, 649
| |
| | |
| Lacrymal bone, 593
| |
| | |
| Lacrymal duct, 506
| |
| | |
| Lacrymal glands, 506
| |
| | |
| Lremargus, vertebral column of, 548
| |
| | |
| Lagena, 524
| |
| | |
| Lamina spiralis, 524
| |
| | |
| Lamina terminalis, 438
| |
| | |
| Larva of Amphioxus, 2 ; of Ascidia, 1 5
| |
| it ; Teleostei, 81 ; Petromyzon, 89, 95;
| |
| Lepidosteus, 117, 318; Amphibia, 134,
| |
| 142; types of, in the Invertebrata, 363
| |
| | |
| Larvre, nature, origin, and affinities of,
| |
| 360 386; secondary variations of less
| |
| likely to be retained, 362 ; ancestral
| |
| history more fully recorded in, 362 ;
| |
| secondary variations in development of,
| |
| 363 ; ontogenetic record of secondary
| |
| variations in, 361; of freshwater and
| |
| land animals, 362; types of, 36.2; phosphorescence of, 364; of Coelenterata,
| |
| | |
| | |
| | |
| 367 ; table of, 365 ; of Invertebrata,
| |
| 367 et seq.
| |
| | |
| Larynx, 766
| |
| | |
| Lateral line sense organs, 538 ; comparison of, with invertebrate, 538 ; development of, in Teleostei, 538 ; development of, in Elasmobranchii, 539
| |
| | |
| Lateral ventricle, 438 ; anterior cornu of,
| |
| 440 ; descending cornu of, 440 ; choroicl
| |
| plexus of, 443
| |
| | |
| Layers, formation of, in Elasmobrancliii,
| |
| 41, 56 ; Teleostei, 71 ; Petromyzon,
| |
| 85 ; Acipenser, 103 ; Lepidosteus, 1 1 1 ;
| |
| Amphibia, 121; Chick, 150, 152;
| |
| Lacerta, 202; Rabbit, 215 227; comparison of Mammalia with lower forms,
| |
| 226, 289; comparison of formation of
| |
| in Vertebrata, 275; origin and homologies of, in the Metazoa, 331
| |
| | |
| Leech, see ' Hirudo '
| |
| | |
| Lemuridre, placenta, 256
| |
| | |
| Lens, of Elasmobranchii, 57, 58 ; Petromyzon, 94, 99; Acipenser, 106 ;
| |
| Lepidosteus, 115 ; Amphibia, 127 ;
| |
| Chick, 177 ; of Vertebrate eyes, 485 ;
| |
| general account of, 493 ; capsule of, 493 ;
| |
| comparative development of, 499 ; of
| |
| Amphibia, Teleostei, Lepidosteus, 499
| |
| | |
| Lepidosteus, development of, 1 1 1 ; larva
| |
| of, 117; relations of, 119; spinal nerves
| |
| of, 455; ribs of, 561 ; generative ducts
| |
| of, 704, 735 ; swimming-bladder of,
| |
| | |
| 763
| |
| | |
| Ligamentum pectinatum, 490
| |
| | |
| Ligamentum suspensorium, 557, 558
| |
| | |
| Ligamentum vesicse medium, 239
| |
| | |
| Limbs, of Elasmobranchii, 59 ; Teleostei,
| |
| 80 ; first appearance of in Chick,
| |
| 184 ; Rabbit, 232 ; muscles of, 673 ; of
| |
| Fishes, 609; relation of, to unpaired fins
| |
| of Fishes, 611, 612; of Amphibia, 61 8
| |
| | |
| Liver of Teleostei, 78 ; Petromyzon, 95,
| |
| 96; Acipenser, no; Amphibia 130;
| |
| general account of, 769
| |
| | |
| Lizard, development of, 202; general
| |
| growth of embryo of, 208 ; Mullerian
| |
| duct of, 721
| |
| | |
| Lizzia, eye of, 471
| |
| | |
| Lobi inferiores, 431
| |
| | |
| Lungs of Amphibia, 137 ; development
| |
| of, 763 ; homology of, 766
| |
| | |
| Lymphatic system, 664
| |
| | |
| Malleus, 529, 591 ; views on, 591
| |
| Malpighian bodies, development of accessory in Elasmobranchs, 695
| |
| Mammalia, development of, 214; comparison of gastrula of, 291 ; cerebellum
| |
| of, 427 ; infundibulum of, 431 ; pineal
| |
| gland of, 434; pituitary body of, 436;
| |
| cerebrum of, 439 ; spinal nerves of, 449 ;
| |
| sympathetic of, 466; vertebral column
| |
| of, 558; branchial arches of, 573, 574;
| |
| mandibular and hyoid arches of, 589 ;
| |
| pectoral girdle of, 604; pelvic girdle of,
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| 787
| |
| | |
| | |
| | |
| 608 ; heart of, 636 ; arterial system of,
| |
| 647; venous system of, 661 ; muscleplates of, 671 ; mesonephros of, 714;
| |
| testicular network of, 724 ; urinogenital
| |
| sinus of, 727 ; spermatozoa of, 747 ;
| |
| lungs of, 765 ; intestines of, 768 ; liver
| |
| of> 769; postanal gut of, 774; stomodseum of, 775
| |
| | |
| Mammary gland, development of, 398
| |
| Man, placenta of, 244 ; general account of
| |
| development of, 265 ; characters of embryo of, 270
| |
| | |
| Mandibular arch of Elasmobranchii, 62,
| |
| 576; Petromyzon, 91 ; Acipenser, 106,
| |
| 116; Chick, 179; general account of,
| |
| | |
| 572, 575; modification of to form jaws,
| |
| | |
| 573, 575; of Teleostei, 580; of Amphibia, 582; Sauropsida, 588; Mammalia, 589
| |
| | |
| Mandibular bar, evolution of, 311, 321
| |
| | |
| Manis, placenta of, 256
| |
| | |
| Marsupial bones, 608
| |
| | |
| Marsupialia, foetal membranes of, 240 ; cerebellum of, 426 ; corpus callosum of,
| |
| ' 443 ; uterus of, 726
| |
| | |
| Maxilla, 594
| |
| | |
| Meatus auditorius externus, of Chick, 181;
| |
| development of, 527
| |
| | |
| Meckelian cartilage, of Elasmobranchii,
| |
| 576; of Teleostei, 581 ; of Amphibia,
| |
| 584, 585; of Sauropsida, 588 ; of Mammalia, 590
| |
| | |
| Mediastinum anterior and posterior, 630
| |
| | |
| Medulla oblongata, of Chick, 176 ; general development of, 425
| |
| | |
| Medullary plate of Amphioxus, 4, 5 ; of
| |
| Ascidia, n; Elasmobranchii, 44, 47,
| |
| 55; Teleostei, 72; Petromyzon, 88;
| |
| Acipenser, 104; Lepidosteus, 1 1 1 ; Amphibia, 126, 127, 131; Chick, 159;
| |
| Lacerta, 204; Rabbit, 223, 227, 228;
| |
| primitive bilobed character of, 303, 317
| |
| | |
| Medusae, auditory organs of, 513
| |
| | |
| Membrana capsulo-pupillaris, 494, 504,
| |
| | |
| 507
| |
| | |
| Membrana elastica externa, 546
| |
| | |
| Membrana limitans of retina, 491
| |
| | |
| Membrana tectoria, 522, 525
| |
| | |
| Membrane bones, of Amphibia, 582 ; of
| |
| Sauropsida, 588; of Mammalia, 590;
| |
| of mandibular arch, 593 ; of pectoral
| |
| girdle, 599, 602 ; origin of, 592 ; homologies of, 593
| |
| | |
| Membranous labyrinth, development of
| |
| in Man, 519
| |
| | |
| Menobranchus, branchial arches of, 142
| |
| | |
| Mesenteron of Elasmobranchii, 43 ; Teleostei, 75 ; Petromyzon, 85 ; Acipenser,
| |
| 104; Amphibia, 123, 124, 129; Chick,
| |
| 167; general account of, 754
| |
| | |
| Mesentery, 626, 756
| |
| | |
| Mesoblast, of Amphioxus, 6 ; Ascidia,
| |
| 17, 20; Pyrosoma, 24; Salpa, 30;
| |
| Elasmobranchii, 44, 47; Teleostei, 75;
| |
| Petromyzon, 86; Acipenser, 105; Lepi
| |
| | |
| | |
| dosteus, 113; Amphibia, 125, 128, 129;
| |
| of Chick, 154, 167; double origin of in
| |
| Chick, 154, 158, 159; origin of from
| |
| lips of blastopore in Chick, 158; of
| |
| area vasculosa of Chick, iOo; Lacerta,
| |
| 203; origin of in Rabbit, 218, 223; of
| |
| area vasculosa in Rabbit, 227; comparative account of formation of, 292 ;
| |
| discussion of development of in Vertebrata, 297 ; meaning of development
| |
| of in Amniota, 298; phylogenetic origin
| |
| of, 346 ; summary of ontogeny of, 349
| |
| 352 ; views on ontogeny of, 352 360
| |
| | |
| Mesoblastic somites, of Amphioxus, 6 ;
| |
| Elasmobranchii, 48, 55 ; Petromyzon,
| |
| 88 ; Acipenser, 105 ; Lepidosteus,
| |
| 114; Amphibia, 129, 131; Chick,
| |
| 161, 1 80; Rabbit, 228; development
| |
| of in Chordata, 325; meaning of development of, 331; of head, 676
| |
| | |
| Mesogastrium, 758
| |
| | |
| Mesonephros, of Teleostei, 78, 702; Petromyzon, 95, 98, 700; Acipenser, 1 10,
| |
| 705; Amphibia, 134, 708; Chick, 184,
| |
| 714; general account of, 690 ; development of in Elasmobranchs, 691 ; of
| |
| Cyclostomata, 700 ; Ganoidei, 705 ;
| |
| sexual and non-sexual part of in Amphibia, 710; of Amniota, 713, 724;
| |
| summary and general conclusions as
| |
| to, 729; relation of to pronephros, 731
| |
| | |
| Mesopterygium, 616
| |
| | |
| Metagenesis of Ascidians, 34
| |
| | |
| Metamorphosis of Amphibia, 137, 140
| |
| | |
| Metanephros, 690; development of in
| |
| Elasmobranchii, 697; of Amphibia,
| |
| 712; of Amniota, 713; of Chick, 722;
| |
| of Lacertilia, 723; phylogeny of, 736
| |
| | |
| Metapterygium, 616
| |
| | |
| Metapterygoid, of Elasmobranchii, 576;
| |
| of Teleostei, 581
| |
| | |
| Metazoa, evolution of, 339, 342 ; ancestral
| |
| form of, 333, 345
| |
| | |
| Mid-brain, of Elasmobranchii, 55, 58,
| |
| 59; Petromyzon, 92; general account
| |
| of development of, 427
| |
| | |
| Moina, generative organs of, 745
| |
| | |
| Molgula, development of, 22
| |
| | |
| Mollusca, nervous system of, 414 ; eyes of,
| |
| 472; auditory organs of, 515; excretory organs of, 68 1
| |
| | |
| Monotremata, foetal membranes of, 240 ;
| |
| cerebellum of, 426; corpus callosum
| |
| of, 443 ; cerebrum of, 443 ; urinogenital sinus of, 726
| |
| | |
| Mormyrus, generative ducts of, 704
| |
| | |
| Mouth, of Amphioxus, 7; of Ascidia, 18;
| |
| Pyrosoma, 27; Salpa, 31; Elasmobranchii, 57, 60, 61, 62; Petromyzon,
| |
| 92, 94, 95, 99; Acipenser, 107; Lepidosteus, 118; Amphibia, 129, 132,
| |
| "134; Rabbit, 231 ; origin of, 317
| |
| | |
| Mouth, suctorial, of Petromyzon, 99;
| |
| Acipenser, 107; Lepidosteus, 116, 317;
| |
| Amphibia, 133, 141, 317
| |
| | |
| | |
| | |
| ;88
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| Mullerian duct, 690; of Elasmobranchs,
| |
| 693 ; of Ganoids, 704 ; of Amphibia,
| |
| 710; of Aves, 717,720; opening of into cloaca, 727; origin of, 733; summary of development of, 733; relation
| |
| of to pronephros, 733
| |
| | |
| Muscle-plates, of Amphioxus, 6; Elasmobranchii, 49, 668 ; Teleostei, 670 ;
| |
| Petromyzon, 94; Chick, 183, 670; general development of, 669 ; of Amphibia,
| |
| 670; Aves, 670; of Mammalia, 671;
| |
| origin of muscles from, 672
| |
| | |
| Muscles, of Ascidia, II, 17; development
| |
| of from muscle-plates, 672; of limbs,
| |
| 673 ; of head, 676 ; of branchial arches,
| |
| 678; of eye, 678
| |
| | |
| Muscular fibres, epithelial origin of, 667
| |
| | |
| Muscular system, development of, 667;
| |
| of Chordata, 668
| |
| | |
| Mustelus, placenta of, 66
| |
| | |
| Myoepithelial cells, 667
| |
| | |
| Mysis, auditory organ of, 517
| |
| | |
| Myxine, ovum of, loo; olfactory organ
| |
| of, 533 ; portal sinus of, 652 ; excretory
| |
| system of, 701
| |
| | |
| Nails, development of, 397
| |
| | |
| Nares, of Acipenser, 108; of Ichthyopsida, 534; development of in Chick,
| |
| 535; development of in Lacertilia, 537;
| |
| development of in Amphibia, 537
| |
| | |
| Nasal bones, 592
| |
| | |
| Nasal pits, Acipenser, 108; Chick, 176;
| |
| general development of, 531
| |
| | |
| Nematoda, excretory organs of, 689 ;
| |
| generative organs of, 745 ; generative
| |
| ducts of, 752
| |
| | |
| Nemertines, nervous system of, 311 ; excretory organs of, 68 1
| |
| | |
| Nerve cord, origin of ventral, 378
| |
| | |
| Nerves, spinal, 449 ; cranial, 455 466
| |
| | |
| Nervous system, central, general account
| |
| of development of in Vertebrata, 415 ;
| |
| conclusions as to, 445; sympathetic,
| |
| 466
| |
| | |
| Nervous system, of Amphioxus, 4; Ascidia, 15, 16; Molgula, 22; Pyrosoma,
| |
| 24, 25; Salpa, 30, 31; Elasmobranchii,
| |
| 44; Teleostei, 77 ; Petromyzon, 89, 93;
| |
| Acipenser, 105; Amphibia, 126; comparative account of formation of central,
| |
| 301; of Sagitta, 349; origin of in
| |
| Ccelenterata, 349; of pneoral lobe,
| |
| 377, 380; evolution of, 400405; development of in Invertebrates, 406;
| |
| of Arthropoda, 408; of Gephyrea, 412;
| |
| Mollusca, 414
| |
| | |
| Neural canal, of Ascidia, 10; Teleostei,
| |
| 72; Petromyzon, 88; Acipenser, 105;
| |
| Lepidosteus, 114; Amphibia, 126, 131 ;
| |
| Chick, 1 66, 171 ; Lacerta, 208; closure
| |
| of in Frog and Amphioxus, 279; closure
| |
| of in Elasmobranchii, 284; phylogcuctic origin of, 316
| |
| | |
| Neural crest, 449, 456, 457
| |
| | |
| | |
| | |
| Neurenteric canal, of Amphioxus, 4, 5 ;
| |
| Ascidia, lo; Elasmobranchii, 54; Petromyzon, 88 ; Acipenser, 105 ; Lepidosteus, 113; Aves, 162; Lacerta, 203,
| |
| 206; general account of, 323; meaning
| |
| of, 3 2 3
| |
| | |
| Newt, ovum of, 120; development of,
| |
| I2 55 general growth of, 141
| |
| | |
| Notidanus, vertebral column of, 548;
| |
| branchial arches of, 572
| |
| | |
| Notochord of Amphioxus, 6; Ascidia,
| |
| II, 17; Elasmobranchii, 51; Teleostei,
| |
| 74; Petromyzon, 86, 94; Acipenser,
| |
| 104; Lepidosteus, 113; Amphibia, 128,
| |
| 129; Chick, 157; canal of, in Chick,
| |
| 163; Lacerta, 204, 205; Guinea-pig,
| |
| 226; comparative account of formation
| |
| of, 292, 325; sheath of, 545; later
| |
| histological changes in, 546; cartilaginous sheath of, 547; in head, 566;
| |
| absence of in region of trabeculas, 567
| |
| | |
| Notodelphys, brood-pouch of, 121 ; branchiae of, 140
| |
| | |
| Nototrema, brood-pouch of, 121
| |
| | |
| Nucleus pulposus, 559
| |
| | |
| Oceania, eye of, 471
| |
| | |
| Occipital bone, 595
| |
| | |
| CEsophagus, solid, of Elasmobranchii,
| |
| 61, 759; of Teleostei, 78
| |
| | |
| Olfactory capsules, 571
| |
| | |
| Olfactory lobes, development of, 444
| |
| | |
| Olfactory nerves, Ammoccetes, 99; general development of, 464
| |
| | |
| Olfactory organ, of aquatic forms, 531;
| |
| Insects and Crustacea, 531; of Tunicata, 532 ; of Amphioxus, 532 ; of
| |
| Vertebrata, 533; Petromyzon, 533;
| |
| of Myxine, 533
| |
| | |
| Olfactory sacks, of Elasmobranchii, 60;
| |
| Teleostei, 73; Petromyzon, 92, 97;
| |
| Acipenser, 106, 108; Lepidosteus, 116;
| |
| Chick, 176
| |
| | |
| Oligochreta, excretory organs of, 683
| |
| | |
| Olivary bodies, 426
| |
| | |
| Omentum, lesser and greater, 757
| |
| | |
| Onchidium, eye of, 473
| |
| | |
| Opercular bones, 593
| |
| | |
| Operculum, of Teleostei, 77; Acipenser,
| |
| 107; Lepidosteus, 117, 118; Amphibia,
| |
| | |
| r 3.5.
| |
| | |
| Ophidia, development of, 210; arterial
| |
| system of, 649 ; venous system of, 656
| |
| | |
| Optic chiasma, 430, 493
| |
| | |
| Optic cup, retinal part of, 488 ; ciliary
| |
| portion of, 489
| |
| | |
| Optic lobes, 428
| |
| | |
| Optic nerve, development of, 492 ; comparative development of, 500
| |
| | |
| Optic thalami, development of, 431
| |
| | |
| Optic vesicle, of Elasmobranchii, 57 59;
| |
| Teleostei, 74, 499 ; Petromyzon, 89, 92 ;
| |
| Acipenser, 106; Lepidosteus, 115;
| |
| Chick, 170; Rabbit, 229; general development of, 429 ; formation of secon
| |
| | |
| | |
| INDKX.
| |
| | |
| | |
| | |
| 7*9
| |
| | |
| | |
| | |
| dary, 487 ; obliteration of cavity of, 488 ;
| |
| comparative development of, 499; of
| |
| Lepidosteus and Teleostei, 499. See
| |
| also ' Eye '
| |
| | |
| Ora serrata, 488
| |
| | |
| Orbitosphenoid region of skull, 570
| |
| | |
| Organs, classification of, 391 ; derivation
| |
| of from germinal layers, 392
| |
| | |
| Orycteropus, placenta of, 249
| |
| | |
| Otic process of Axolotl, 583; of Frog,
| |
| 585 et seq.
| |
| | |
| Otoliths, 512
| |
| | |
| Oviposition, of Amphioxus, i ; Elasmobranchii, 40; Teleostei, 68; Petromyzon, 84; Amphibia, 121; Reptilia, 202
| |
| | |
| Ovum, of Amphioxus, i; Pyrosoma, 23;
| |
| Elasmobranchii, 40; Teleostei, 68;
| |
| Petromyzon, 83 ; Myxine, loo; Acipenser, 102; Lepidosteus, in; Amphibia,
| |
| 120; Chick, 146; Reptilia, 202 ; Mammalia, 214; of Porifera, 741; migration of in Ccelenterata, 742; Vertebrata, 746
| |
| | |
| Palatine bone, of Teleostei, 580; origin
| |
| of, 594
| |
| | |
| Pancreas, Acipenser, no; general development of, 770
| |
| | |
| Pancreatic caeca, of Teleostei, etc. 768
| |
| | |
| Papillae, oral, of Acipenser, 108; Lepidosteus, n6
| |
| | |
| Parachordals, 565, 566
| |
| | |
| Parasphenoid bone, 594
| |
| | |
| Parepididymis, 725
| |
| | |
| Parietal bones, 592
| |
| | |
| Paroophorori, 725
| |
| | |
| Parovarium, 725
| |
| | |
| Pectoral girdle, 599 ; of Elasmobranchs,
| |
| 600; of Teleostei, 600; of Amphibia
| |
| and Amniota, 60 1 ; comparison of with
| |
| pelvic, 608
| |
| | |
| Pecten, eye of, 479
| |
| | |
| Pecten, of Ammoccetes, 498; of Chick,
| |
| 501 ; Lizard, 501 ; Elasmobranchs, 501
| |
| | |
| Pedicle, of Axolotl, 484 ; of Frog, 485
| |
| | |
| Pelobates, branchial apertures of, 136;
| |
| vertebral column of, 556
| |
| | |
| Pelodytes, branchial chamber of, 135
| |
| | |
| Pelvic girdle, 606; of Fishes, 606; Amphibia and Amniota, 607 ; of Lacertilia, 607 ; of Mammalia, 608 ; comparison with pectoral, 608
| |
| | |
| Penis, development of, 727
| |
| | |
| Peribranchial cavity, of Amphioxus, 7;
| |
| of Ascidia, 18; Pyrosoma, 24
| |
| | |
| Pericardial cavity, of Pyrosoma, 26 ; Elasmobranchii, 49 ; Petromyzon, 94; general account of, 626; of Fishes, 627 ; of
| |
| Amphibia, Sauropsida and Mammalia,
| |
| 628
| |
| | |
| Perichordal formation of vertebral column,
| |
| 5^6
| |
| | |
| Perilymph of ear, 523
| |
| Periotic capsules, ossifications in, 595,
| |
| 596
| |
| | |
| | |
| | |
| Peripatus, nervous system of, 409 ; eye of
| |
| 480 ; excretory organs of, 688
| |
| | |
| Peritoneal membrane, 626
| |
| | |
| Petromyzon, development of, 83; affinities of, 83, 84; general development
| |
| of, 87; hatching of, 89; comparison of
| |
| gastrula of, 280; branchial skeleton of,
| |
| 312, 572; cerebellum of, 425; pineal
| |
| gland of, 434 ; pituitary body of, 436 ;
| |
| cerebrum of, 439; auditory organ of,
| |
| 517; olfactory organ of, 533; comparison of oral skeleton of with Tadpole,
| |
| 586; pericardial cavity of, 627; abdominal pores of, 626 ; venous system of,
| |
| 651 ; excretory organs of, 700; segmental duct of, 700; pronephros of, 700;
| |
| mesonephros of, 700 ; thyroid body of,
| |
| 760; postanalgut of, 774; stomodx-um
| |
| | |
| of, 775
| |
| | |
| Phosphorescence of larvae, 364
| |
| | |
| Phylogeny, of the Chordata, 327; of the
| |
| Metazoa, 384
| |
| | |
| Pig, placenta of, 251; mandibular and
| |
| hyoid arches of, 589
| |
| | |
| Pineal gland, of Petromyzon, 93 ; Chick,
| |
| 175; general development of, 432;
| |
| nature of, 432, 434
| |
| | |
| Pipa, brood-pouch of, 121 ; metamorphosis of, 139; yolk-sack of, 140; vertebral
| |
| column of, 556
| |
| | |
| Pituitary body, of Rabbit, 231 ; general
| |
| development of, 435 ; meaning of, 436 ;
| |
| Placenta, of Salpa, 29; Elasmobranchii, 66; of Mammalia, 232; villi of,
| |
| 235 ; deciduate and non-deciduate, 239;
| |
| comparative account of, 239 259 ; characters of primitive type of, 240; zonary, 248; non-deciduate, 250; histology of, 257; evolution of, 259
| |
| | |
| Placoid scales, 395
| |
| | |
| Planorbis, excretory organs of, 68 1
| |
| | |
| Planula, structure of, 367
| |
| | |
| Pleural cavities, 631
| |
| | |
| Pleuronectidae, development of, 80
| |
| | |
| Pneumatoccela, characters of, 327
| |
| | |
| Polygordius, excretory organs of, 684
| |
| | |
| Polyophthalmus, eye of, 479
| |
| | |
| Polypedates, brood-pouch of, 121
| |
| | |
| Polyzoa, excretory organs of, 682 ; generative cells of, 745 ; generative ducts
| |
| | |
| of, 751
| |
| | |
| Pons Varolii, 426, 427
| |
| | |
| Pori abdominales, Ammoccetes, 99
| |
| | |
| Porifera, ancestral form of, 345 ; development of generative cells of, 74!
| |
| | |
| Portal vein, 653
| |
| | |
| Postanal gut of Elasmobranchii, 58, 59,
| |
| 60; Teleostei, 75; Chick, 169; general account of, 323, 772
| |
| | |
| Prsemaxilla, 594
| |
| | |
| Praeopercular bone, 593
| |
| | |
| Prrcoral lobe, ganglion of, 377, 380
| |
| | |
| Prefrontals, 597
| |
| | |
| Presphenoid region of skull, 570
| |
| | |
| Primitive groove of Chick, 1 55
| |
| | |
| | |
| | |
| 790
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| Primitive streak, of Chick, 152, 161;
| |
| meaning of, 153; origin of mesoblast
| |
| form in Chick, 154; continuity of
| |
| hypoblast with epiblast at anterior end
| |
| of, in Chick, 156; comparison of with
| |
| blastopore, 165 ; fate of, in Chick, 165 ;
| |
| of Lacerta, 203; of Rabbit, 221; of
| |
| Guinea-pig, 223 ; fusion of layers at, in
| |
| Rabbit, 224; comparison of with blastopore of lower forms, 226, 287 ; of
| |
| Mammalia, 290
| |
| | |
| Processus falciformis of Ammoccetes, 498 ;
| |
| of Elasmobranch, 502 ; of Teleostei , 503
| |
| Proctodseum, 778
| |
| | |
| Pronephros, of Teleostei, 78, 701 ; Petromyzon, 95, 99, 700; Acipenser, 106,
| |
| no; Amphibia, 134, 707; general account of, 689 ; of Cyclostomata, 700 ;
| |
| of Myxine, 701 ; Ganoidei, 705 ; of
| |
| Amniota, 714; of Chick, 718; summary of and general conclusions as to,
| |
| 728; relation of, to mesonephros, 731 ;
| |
| cause of atrophy of, 729
| |
| Prootic, 596, 597
| |
| Propterygium, 616
| |
| Proteus, branchial arches of, 142
| |
| Protochordata, characters of, 327
| |
| Protoganoidei, characters of, 328
| |
| Protognathostomata, characters of, 328
| |
| Protopentadactyloidei, characters of, 329
| |
| Protovertebrata, characters of, 328
| |
| Pseudis, Tadpole of, 139; vertebral
| |
| | |
| column of, 556
| |
| | |
| Pseud ophryne, yolk-sack of, 140; Tadpole of, 140
| |
| Pterygoid bone, of Teleostei, 581; origin
| |
| | |
| of, 597
| |
| | |
| Pterygoquadrate bar, of Elasmobranchii,
| |
| 576; of Teleostei, 581; Axolotl, 584;
| |
| F r g, 584; ofSauropsida, 588; of Mammalia, 589
| |
| | |
| Pulmonary artery, origin of, 645 ; of
| |
| Amphibia, 645 ; of Amniota, 649
| |
| | |
| Pulmonary vein, 655
| |
| | |
| Pupil, 489
| |
| | |
| Pyrosoma, development of, 23
| |
| | |
| Quadrate bone of Teleostei, 581 ; of
| |
| Axolotl, 584; Frog, 585; Sauropsida,
| |
| 588
| |
| | |
| Quadratojugal bone, 594
| |
| | |
| Rabbit, development of, 214; general
| |
| growth of embryo of, 227 ; placenta of,
| |
| 248
| |
| | |
| Radiate symmetry, passage from to bilateral symmetry, 373 376
| |
| | |
| Raja, caudal vertebras of, 553
| |
| | |
| Rat, placenta of, 242
| |
| | |
| Recessus labyrinthi, 519
| |
| | |
| Reissner's membrane, 524
| |
| | |
| Reptilia, development of, 202; viviparous,
| |
| 202; cerebellum of, 426; infundibulum
| |
| of, 431; pituitary body of, 436; cerebrum of, 439; vertebral column of,
| |
| | |
| | |
| | |
| 556; arterial system of, 648; venous
| |
| system of, 656; mesonephros of, 713;
| |
| testicular network of, 723; spermatozoa
| |
| of, 747
| |
| | |
| Restiform tracts of Elasmobranchii and
| |
| Teleostei, 425
| |
| | |
| Retina, histogenesis of, 490
| |
| | |
| Retinulse, 482
| |
| | |
| Rhabdom, 482
| |
| | |
| Rhinoderma, brood-pouch of, 121; metamorphosis of, 1 39
| |
| | |
| Ribs, development of, 560
| |
| | |
| Roseniniiller's organ, 725
| |
| | |
| Rotifera, excretory organs of, 680
| |
| | |
| Round ligament of liver, 663
| |
| | |
| Ruminantia, placenta of, 253
| |
| | |
| Sacci vasculosi, 437
| |
| | |
| Sacculus hemisphericus, 519; of Mammals, 519, 520
| |
| | |
| Sagitta. See ' Chaetognatha'
| |
| | |
| Salpa, sexual development of, 29; asexual
| |
| development of, 33
| |
| | |
| Salamandra, larva of, 142; vertebral
| |
| column of, 553; limbs of, 619; mesonephros of, 708; Miillerian duct of,
| |
| 710
| |
| | |
| Salmonidse, hypoblast of, 71; generative
| |
| ducts of, 704
| |
| | |
| Sauropsida, gastrula of, 286; meaning of
| |
| primitive streak of, 288; blastopore of,
| |
| 289 ; mandibular and hyoid arches of,
| |
| 588 ; pectoral girdle of, 60 1
| |
| | |
| Scala, vestibuli, 522; tympani, 523;
| |
| media, 522
| |
| | |
| Scales, general development of, 396 ; development of placoid scales, 395
| |
| | |
| Scapula, 599
| |
| | |
| Sclerotic, 488
| |
| | |
| Scrotum, development of, 727
| |
| | |
| Scyllium, caudal vertebrse of, 553; mandibular and hyoid arches of, 578; pectoral girdle of, 600; limbs of, 610; pelvic fin of, 614; pectoral fin of, 615
| |
| | |
| Segmental duct, 690 ; development of in
| |
| Elasmobranchs, 690; of Cyclostomata,
| |
| 700; of Teleostei, 701; of Ganoidei,
| |
| 704, 705 ; of Amphibia, 707 ; of Amniota, 713
| |
| | |
| Segmental organs, 682
| |
| | |
| Segmental tubes, 690 ; development of in
| |
| Elasmobranchs, 691 ; rudimentary anterior in Elasmobranchs, 693 ; development of secondary, 731
| |
| | |
| Segmentation cavity, of Elasmobranchii,
| |
| 42 44; Teleostei, 69, 85, 86; Amphibia, 122, 125
| |
| | |
| Segmentation, meaning of, 331
| |
| | |
| Segmentation of ovum, in Amphioxus, 2 ;
| |
| Ascidia, 9 ; Molgula, 22 ; Pyrosoma,
| |
| 23; Salpa, 30; Elasmobranchii, 40;
| |
| Telostei, 69; Petromyzon, 84; Acipenser, IOT, Lcpidosteus, in; Amphibia, 122, 124; Newt, 125; Chick,
| |
| 146; Lizard, 202: Rabbit, 214
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| 791
| |
| | |
| | |
| | |
| Semicircular canals, 519
| |
| | |
| Sense organs, comparative account of
| |
| development of, 304
| |
| | |
| Septum lucidum, 443
| |
| | |
| Serous membrane, Lacerta, 209; of Rabbit, 237
| |
| | |
| Seventh nerve, development of, 459
| |
| | |
| Shell-gland of Crustacea, 689
| |
| | |
| Shield, embryonic, of Chick, 151 ; of
| |
| Lacerta, 202
| |
| | |
| SimiadiK, placenta of, 247
| |
| | |
| Sinus rhomboidalis, of Chick, 162
| |
| | |
| Sinus venosus, 637
| |
| | |
| Sirenia, placenta of, 255
| |
| | |
| Sixth nerve, 463
| |
| | |
| Skate, mandibular and hyoid arches of,
| |
| | |
| 577
| |
| | |
| Skeleton, elements of found in Vertebrata, 542
| |
| | |
| Skull, general development of, 564 ; historical account of, 564 ; development of
| |
| cartilaginous, 566; cartilaginous walls
| |
| of, 570; composition of primitive cartilaginous cranium, 565
| |
| | |
| Somatopleure, of Chick, 170
| |
| | |
| Spelerpes, branchial arches of, 142
| |
| | |
| Spermatozoa, of Porifera, 741; of Vertebrata, 746
| |
| | |
| Sphenoid bone, 595
| |
| | |
| Sphenodon, hyoid arch of, 588
| |
| | |
| Spinal cord, general account of, 415;
| |
| white matter of, 415; central canal of,
| |
| 417, 418; commissures of, 417; grey
| |
| matter of, 417; fissures of, 418
| |
| | |
| Spinal nerves, posterior roots of, 449;
| |
| anterior roots of, 453
| |
| | |
| Spiracle, of Elasmobranchii, 62 ; Acipenser, 105; Amphibia, 136
| |
| | |
| Spiral valve. See 'Valve'
| |
| | |
| Spleen, 664
| |
| | |
| Splenial bone, 595
| |
| | |
| Squamosal bone, 593
| |
| | |
| Stapes, 529; of Mammal, 590
| |
| | |
| Sternum, development of, 562
| |
| | |
| Stolon of Doliolum, 29 ; Salpa, 33
| |
| | |
| Stomodaeum, 774
| |
| | |
| Stria vascularis, 524
| |
| | |
| Styloid process, 591
| |
| | |
| Sub-intestinal vein, 65 1 ; meaning of,
| |
| | |
| 651
| |
| | |
| Syngnathus, brood-pouch of, 68
| |
| Subnotochordal rod, of Elasmobranchii,
| |
| | |
| 54; Petromyzon, 94; Acipenser, no;
| |
| | |
| Lepidosteus, 115; general account of,
| |
| | |
| 754; comparison of with siphon of
| |
| | |
| Chsetopods, 756
| |
| | |
| Subzonal membrane, 237; villi of, 236
| |
| Sulcus of Munro, 432
| |
| Supraclavicle, 600
| |
| Suprarenal bodies, 664
| |
| Supra-temporal bone, 593
| |
| Swimming bladder, see Air bladder
| |
| Sylvian aqueduct, 428
| |
| Sylvian fissure, 444
| |
| Sympathetic ganglia, development of, 467
| |
| | |
| | |
| | |
| Tadpole, 134, 139, 140; phylogenetic
| |
| meaning of, 137; metamorphosis of,
| |
| 137; m can ing of suctorial mouth of, 585
| |
| | |
| Tail of Teleostei, 80; Acipenser, 109;
| |
| Lepidosteus, 109; Amphibia, 132
| |
| | |
| Tarsus, development of, 620
| |
| | |
| Teeth, horny provisional, of Amphibia,
| |
| 136; general development of, 776;
| |
| origin of, 777
| |
| | |
| Teleostei, development of, 68; viviparous, 68; comparison of formation of
| |
| layers in, 286; restiform tracts of, 425 ;
| |
| mid-brain of, 425 ; infundibulum of,
| |
| 431 ; cerebrum of, 439; nares of, 534;
| |
| lateral line of, 538; notochord and
| |
| membrana elastica of, 549 ; vertebral
| |
| column of, 553; ribs of, 561; hyoid
| |
| and mandibular arches of, 579; pectoral girdle of, 601 : pelvic girdle of,
| |
| 606; limbs of, 618; heart of, 637;
| |
| arterial system of, 645; muscle-plates
| |
| of, 670; excretory organs of, 701 ; generative ducts of, 704, 735, 749; swimming bladder of, 763 ; postanal gut of,
| |
| | |
| Teredo, nervous system of, 414
| |
| | |
| Test of Ascidia, 14; Salpa, 31
| |
| | |
| Testicular network, of Elasmobranchs,
| |
| 697 ; of Amphibia, 712 ; Reptilia, 723 ;
| |
| of Mammals, 724
| |
| | |
| Testis of Vertebrata, 746
| |
| | |
| Testis, connection of with Wolffian body,
| |
| in Elasmobranchii, 697; in Amphibia,
| |
| 710; in Amniota, 723; origin of, 735
| |
| | |
| Thalamencephalon of Chick, 175; general development of, 430
| |
| | |
| Third nerve, development of, 461
| |
| | |
| Thymus gland, 762
| |
| | |
| Thyroid gland, Petromyzon, 92 ; general
| |
| account of, 759; nature of, 760; development of in Vertebrata, 761
| |
| | |
| Tooth. See 1 Teeth'
| |
| | |
| Tori semicirculares, 428
| |
| | |
| Tornaria, 372
| |
| | |
| Trabeculas, 565, 567; nature of, 568
| |
| | |
| Trachea, 766
| |
| | |
| Trematoda, excretory organs of, 68 1
| |
| | |
| Triton alpestris, sexual larva of, 143
| |
| | |
| Triton, development of limbs of, 619}
| |
| urinogenital organs of, 7 12
| |
| | |
| Truncus arteriosus, 638; of Amphibia,
| |
| 638; of Birds, 639
| |
| | |
| Turiicata, development of mesoblast of,
| |
| 293; test of, 394; eye of, 507; auditory organ of, 530; olfactory organ of,
| |
| 532; generative duct of, 749 ; intestine
| |
| of, 767; postanal gut of, 771; stomodseum of, 775
| |
| | |
| Turbellaria, excretory organs of, 68 1
| |
| | |
| Tympanic annulus of *'rog, 587
| |
| | |
| Tympanic cavity, of Amphibia, 135;
| |
| Chick, 1 80; Rabbit, 232; general development of, 528; of Mammals, 591
| |
| | |
| Tympanic membrane, of Chick, 180;
| |
| general development of, 528
| |
| | |
| | |
| | |
| 792
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| Tympanohyal, 591
| |
| | |
| Umbilical canal of Elasmobranchii, 54,
| |
| | |
| 57, 58, 59
| |
| | |
| Umbilical cord, 238; vessels of, 239
| |
| | |
| Ungulata, placenta of, 250
| |
| | |
| Urachus, 239, 726
| |
| | |
| Ureters, of Elasmobranchii, 696; development of, 723
| |
| | |
| Urethra, 727
| |
| | |
| Urinary bladder of Amphibia, "Jii; of
| |
| Amniota, 726
| |
| | |
| Urinogenital organs, see Excretory organs
| |
| | |
| Urinogenital sinus of Petromyzon, 700;
| |
| of Sauropsida, 726; of Mammalia,
| |
| 727
| |
| | |
| Urochorda, development of, 9
| |
| | |
| Uterus, development of, 726; of Marsupials, 726
| |
| | |
| Uterus masculinus, 726
| |
| | |
| Utriculus, 519
| |
| | |
| Uvea of iris, 489
| |
| | |
| Vagus nerve, development of, 456, 457;
| |
| intestinal branch of, 458; branch of to
| |
| lateral line, 459
| |
| | |
| Valve, spiral, of Petromyzon, 97; Acipenser, no; general account of, 767
| |
| | |
| Valves, semilunar, 641; auriculo-ventricular, 642
| |
| | |
| Vasa efferentia, of Elasmobranchs, 697 ;
| |
| of Amphibia, 711; general origin of,
| |
| 724
| |
| | |
| Vascular system, of Amphioxus, 8; Petromyzon, 97; Lepidosteus, 116; general
| |
| development of, 632
| |
| | |
| Vas deferens, of Elasmobranchii, 697 ;
| |
| of Amniota, 723
| |
| | |
| Vein, sub-intestinal of Petromyzon, 97 ;
| |
| Acipenser, no; Lepidosteus, 116
| |
| | |
| Velum of Petromyzon, 9 1
| |
| | |
| Vena cava inferior, development of, 655
| |
| | |
| Venous system of Petromyzon, 97; general development of, 651; of Fishes,
| |
| 651 ; of Amphibia and Amniota, 655 ;
| |
| of Reptilia, 656; of Ophidia, 656; of
| |
| Aves, 658; of Mammalia, 661
| |
| | |
| Ventricle, fourth, of Chick, 176; history
| |
| of, 424
| |
| | |
| Ventricle, lateral, 438, 440; fifth, 443
| |
| | |
| Ventricle, third, of Chick, 175
| |
| | |
| Vertebral bodies, of Chick, 183
| |
| | |
| Vertebral column, development of, 545,
| |
| 549; epichordal and perichordal development of in Amphibia, 556
| |
| | |
| Vespertilionidse, early development of,
| |
| 217
| |
| | |
| Vieussens, valve of, 426
| |
| | |
| Villi, placental, of zona radiata, 235 ;
| |
| subzonal membrane, 235; chorion, 237;
| |
| | |
| | |
| | |
| Man, 246; comparative account of,
| |
| 2 575 of young human ovum, 265, 269
| |
| | |
| Visceral arches, Amphioxus, 7 ; Elasmobranchii, 57 60; Teleostei, 77; Acipenser, 1 06; Lepidosteus, 116; Amphibia, 133; Chick, 177; Rabbit,
| |
| 231; prseoral, 570; relation of to head
| |
| cavities, 572; disappearance of posterior, 573; dental plates of in Teleostei, 574
| |
| | |
| Visual organs, evolution of, 470
| |
| | |
| Vitelline arteries of Chick, 195
| |
| | |
| Vitelline veins of Chick, 195
| |
| | |
| Vitreous humour, of Ammoccetes, 98 ;
| |
| general development of, 494; blood*
| |
| vessels of in Mammals, 503 ; mesoblastic ingrowth in Mammals, 503
| |
| | |
| Vomer, 594
| |
| | |
| White matter, of spinal cord, 415; of
| |
| brain, 423
| |
| | |
| Wolffian body, see ' Mesonephros '
| |
| | |
| Wolffian duct, first appearance of in
| |
| Chick, 183; general account of, 690;
| |
| of Elasmobranchs, 693 ; of Ganoids,
| |
| 704; of Amphibia, 710; of Amniota,
| |
| 713; atrophy of in Amniota, 724
| |
| | |
| Wolffian ridge, 185
| |
| | |
| Yolk blastopore, of Elasmobranchii, 64
| |
| | |
| Yolk, folding off of embryo from, in
| |
| Elasmobranchii, 55; in Teleostei, 76;
| |
| Acipenser, 106; Chick, 168, 170
| |
| | |
| Yolk nuclei, of Elasmobranchii, 41, 53;
| |
| Teleostei, 69, 75
| |
| | |
| Yolk, of Elasmobranchii, 40; Teleostei,
| |
| 68; Petromyzon, 96; Acipenser, 109;
| |
| Amphibia, 122, 129; Chick, 146; influence of on formation of layers, 278;
| |
| influence of on early development,
| |
| | |
| 341, 342
| |
| | |
| Yolk-sack, Amphibia, 131, 140, 141; enclosure of, 123
| |
| | |
| .Yolk-sack, development of in Rabbit,
| |
| 227; of Mammalia reduced, 227; circulation of in Rabbit, 233 ; enclosure
| |
| of in Sauropsida, 289
| |
| | |
| Yolk-sack, enclosure of, Petromyzon, 86
| |
| | |
| Yolk-sack, Lepidosteus, 118
| |
| | |
| Yolk-sack of Chick, enclosure of, 160;
| |
| stalk of, 174; general account of, 193;
| |
| circulation of, 195 ; later history of, 198
| |
| | |
| Yolk-sack of Elasmobranchii, enclosure
| |
| of, 62, 283; circulation of, 64
| |
| | |
| Yolk-sack of Lacerta, 209 ; circulation of,
| |
| 209
| |
| | |
| Yolk-sack, Teleostei, 75, 81; enclosure
| |
| of, 75 ; circulation of, 81
| |
| | |
| Zona radiata, villi of, 237
| |
| Zonula of Zinn, 495
| |
| | |
| | |
| | |
| BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| CEPHALOPODA.
| |
| | |
| (1) A. Kowalevsky. " Entwicklungsgeschichte des Amphioxus lanceolatus."
| |
| Mem. Acad. Imper. des Sciences de St Pttersbourg, Series vn. Tom. XI. 1867.
| |
| | |
| (2) A. Kowalevsky. "Weitere Studien iiber die Entwicklungsgeschichte des
| |
| Amphioxus lanceolatus." Archiv f. mikr. Anat., Vol. xui. 1877.
| |
| | |
| (3) Leuckart u. Pagenstecher. " Untersuchungen tiber niedere Seethiere."
| |
| Mutter's Archiv, 1858.
| |
| | |
| (4) Max Schultze. " Beobachtung junger Exemplare von Amphioxus." Zeit.
| |
| f. wiss. Zool., Bd. in. 1851.
| |
| | |
| (5) A. M. Marshall. "On the mode of Ovi position of Amphioxus." your,
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| of Anat. and Phys., Vol. x. 1876.
| |
| | |
| UROCHORDA.
| |
| | |
| (6) P. J. van Beneden. " Recherches s. 1'Embryogenie, 1'Anat. et la Physiol.
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| des Ascidies simples." Mem. Acad. Roy. de Belgique, Tom. xx.
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| | |
| (7) W. K. Brooks. "On the development of Salpa." Bull, of the Museum of
| |
| Comp. Anat. at Harvard College, Cambridge, Mass.
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| | |
| (8) H. Fol. Eludes surles Appendiculaires du detroit de Mcssine . Geneve et
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| Bale, 1872.
| |
| | |
| (9) Ganin. "Neue Thatsachen a. d. Entwicklungsgeschichte d. Ascidien."
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| Zeit.f. wiss. Zool., Vol. XX. 1870.
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| | |
| (10) C. Gegenbaur. " Ueber den Entwicklungscyclus von Doliolum nebst
| |
| Bemerkungen iiber die Larven dieser Thiere." Zeit.f. wiss. Zool., Bd. vu. 1856.
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| | |
| (11) A. Giard. "Etudes critiques des travaux d'embryogenie relatifs a la
| |
| parente des Vertebres et des Tuniciers." Archiv Zool. experiment., Vol. I. 1872.
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| (12) A. Giard. " Recherches sur les Synascidies. " Archiv Zool. exper., Vol. I.
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| 1872.
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| (13) O. Hertwig. "Untersuchungen lib. d. Bau u. d. Entwicklung des Cellulose-Mantels d. Tunicaten." Jenaische Zeitschrift, Bd. vn. 1873.
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| | |
| (14) Th. H. Huxley. " Remarks upon Appendicularia and Doliolum. " Phil.
| |
| Trans., 1851.
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| | |
| (15) Th. H.Huxley. " Observations on the anatomy and physiology of Salpa
| |
| and Pyrosoma." Phil. Trans., 1851.
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| | |
| (16) Th. H. Huxley. "Anatomy and development of Pyrosoma." Linnean
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| Trans., 1860, Vol. XXIII.
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| | |
| (17) Keferstein u. Ehlers. Zoologische Beitrage, 1861. Doliolum.
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| (18) A. Kowalevsky. "Entwicklungsgeschichte d. einfachen Ascidien." Mem.
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| Acad. Pctersbourg, VII. serie, T. x. 1866.
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| (19) A. Kowalevsky. "Beitrag z. Entwick. d. Tunicaten." Nachrichtcn d.
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| konigl. Gesell.zu Gottingen. 1868.
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| | |
| (20) A. Kowalevsky. "Weitere Studien iib. d. Entwicklung d. einfachen Ascidien." Archiv f. mikr. Anat., Vol. vn. 1871.
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| (21) A. Kowalevsky. "Ueber Knospung d. Ascidien." Archiv f. mikr. Anat.,
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| Vol. X. 1874.
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| (22) A. Kowalevsky. "Ueber die Entwicklungsgeschichte d. Pyrosoma."
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| Archiv f. mikr. Anat., Vol. xi. 1875.
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| (23) A. Krohn. "Ueber die Gattung Doliolum u. ihre Arten." Archiv f.
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| Natnrgeschichte, Bd. xvm. 1852.
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| B. Hi. a
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| BIBLIOGRAPHY.
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| (24) A. Krohn. "Ueber die Entwicklung d. Ascidien." Mailer's Archiv,
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| 1852.
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| | |
| (25) A. Krohn. "Ueber die Fortpfianzungsverhaltnisse d. Botrylliden. " Archiv
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| f. Naturgeschichte, Vol. xxxv. 1869.
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| | |
| (26) A. Krohn. "Ueber die fruheste Bildung d. Botryllenstocke." Archiv f.
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| Naturgeschichte, Vol. xxxv. 1869.
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| (27) C. Kupffer. " Die Stammverwandschaft zwischen Ascidien u. Wirbelthieren." Archiv f, mikr. Anat., Vol. vi. 1870.
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| | |
| (28) C. Kupffer. "Zur Entwicklung d. einfachen Ascidien." Archiv f. mikr.
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| Anat., Vol. vm. 187-2.
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| | |
| (29) H. Lacaze Duthiers. "Recherches sur 1'organisation et 1'Embryogenie
| |
| des Ascidies (Molgula tubulosa)." Comptes rendus, May 30, 1870, p. 1154.
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| | |
| (30) H. Lacaze Duthiers. "Les Ascidies simples des Cotes de France" (Development of Molgula). Archiv Zool. exper., Vol. ill. 1874.
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| | |
| (31) R. Leuckart. "Salpa u. Verwandte." Zoologischc Untcrsuchungen,
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| Heft u.
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| (32) E. Metschnikoff. " Observations sur le developpement de quelques animaux (Botryllus and Simple Ascidians)." Still, d. fAcad. Petersbottrg, Vol. xm.
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| 1869.
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| (33) H. Milne-Edwards. "Observations s. 1. Ascidies composees des cotes de
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| la Manche." Memoir es d. V Instittit, T. xvm. 1842.
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| | |
| (34) W. Salensky. "Ueber d.embryonaleEntwicklungsgeschichtederSalpen."
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| Zeit.f. wiss. Zool., B. xxvn. 1877.
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| (35) W. Salensky. "Ueber die Knospung d. Salpen." Morphol. Jahrbuch,
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| Bd. in. 1877.
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| (36) W. Salensky. "Ueber die Entwicklung d. Hoden u. iiber den Generationswechsel d. Salpen." Zeit.f. wiss. Zool., Bd. xxx. Suppl. 1878.
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| | |
| (37) C. Semper. " Ueber die Entstehung d. geschichteten Cellulose-Epidermis
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| d. Ascidien." Arbeit, a. d. zool.-zoot. Instit. Wiirzburg, Vol. ri. 1875.
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| (38) Fr. Todaro. Sopra lo sviluppo e F anatomia delle Salpc. Roma, 1875.
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| (39) Fr. Todaro. "Sui primi fenomeni dello sviluppo delle Salpe." Realc
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| Accadcmia dci Lincei, Vol. iv. 1880.
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| | |
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| ELASMOBRANCHII.
| |
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| (40) F. M. Balfour. " A preliminary account of the development of the Elasmobranch Fishes." Quart. J. of Micr. Science, Vol. xiv. 1876.
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| (41) F. M. Balfour. "A Monograph on the development of Elasmob ranch
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| Fishes." London, 1878. Reprinted from the Journal of Anat. and Fhysiol. for 1876,
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| 1877, and 1878.
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| | |
| (42) Z. Gerbe. " Recherches sur la segmentation de la cicatrule et la formation
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| des prodnits adventifs de Pceuf des Plagiostomes et particulierement des Rates." Vide
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| also Journal de FAnatomie et de la Physiologic, 1872.
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| Entwick., Vol. 11. 1877.
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| Irevis. " (Russian.) Transactions of the Kiew Society of Naturalists, Vol. I. 1870.
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| (45) R. Leuckart. "Ueber die allmahlige Bildung d. Korpergestalt bei d.
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| Rochen." Zeit. f. wiss. Zool., Bd. II., p. 258.
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| (46) Fr. Ley dig. Rochen u. Hate. Leipzig, 1852.
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| vetenskaps akademiens fo'rhandlingar. Stockholm, 1876.
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| | |
| (48) Joh. M tiller. Clatter Haie des Aristoteles und iiber die Verschiedenheitcn
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| unler den Haifachen und Rochen in der Entivicklung des Eies. Berlin, 1840.
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| Sitz. der k. Akad. Wien, Vol. LXXIII. 1873.
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| | |
| (50) Alex. Schultz. " Zur Entwicklungsgeschichte des Selachiereies. " Archiv
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| fiir micro. Anat., Vol. XI. 1875.
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| (51) Alex. Schultz. " Beitrag zur Entwicklungsgeschichte d. Knorpelfische. "
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| Archiv fiir micro. Anat., Vol. xni. 1877.
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| | |
| | |
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| BIBLIOGRAPHY.
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| | |
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| Ill
| |
| | |
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| (52) C. Semper. "Die Stammesverwandschaft d. Wirbelthiere u. Wirlwllosen. Arbeit, a. d. zool.-zoot. Instit. Wiirzburg, Vol. II. 1875.
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| (54) Wyman. " Observations on the Development of Raja batis." Memoirs of
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| the American Academy of Arts and Sciences, Vol. ix. 1864.
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| | |
| | |
| | |
| TELEOSTEI.
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| (55) Al. Agassiz. " On the young Stages of some Osseous Fishes. I. Development of the Tail." Proceedings of the American Academy of Arts and Sciences,
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| xiv BIBLIOGRAPHY.
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| Foetus des Menschen. Niirnberg, 1816.
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| Peripheral Nervous System of the Vertebrata.
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| (351) F. M. Balfour. "On the development of the spinal nerves in Elasmobranch Fishes." Philosophical Transactions, Vol. CLXVI. 1876; vide also, A monograph on the development of Elasmobranch Fishes. London, 1878, pp. 191216.
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| f. Anat. u. Physiol., 1879.
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| Quart, y. of Micr. Science, Vol. xvni. 1878.
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| Quart, y. of Micr. Science, Vol. xix. 1879.
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| (357) C. Schwalbe. "Das Ganglion oculomotorii. " Jenaische Zeitschrift,
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| Vol. xni. 1879.
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| Sympathetic Nervous System.
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| (360) F. M. Balfour. Monograph on the development of Elasmobranch Fishes.
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| London, 1878, p. 173.
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| III. 1879.
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| BIBLIOGRAPHY. XV
| |
| | |
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| THE EYE.
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| Eye of the Mollusca.
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| (362) N. Bobretzky. " Observations on the development of the Cephalopoda "
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| (Russian). Nachrichtcn d. kaiserlichen Gesell. d. Frennde der Natuna iss. Anthropolog.
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| Ethnogr. bei d. Universitdt Moskau.
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| f. wiss. Zool., Bd. xxiv. 1874.
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| Zool., Vol. xv. 1865.
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| 1877.
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| Eye of the Arthropoda.
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| Gottingen, 1879.
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| The Vertebrate Eye.
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| Accessory organs of the Vertebrate Eye.
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| Eye of the T2tnicata,
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| | |
| | |
| | |
| xvi BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| AUDITORY ORGANS.
| |
| Auditory organs of tlie Invertebrata.
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| (384) V. Hensen. "Studien lib. d. Gehororgan d. Decapoden." Zeil.f. wiss.
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| Zool., Vol. xui. 1863.
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| Leipzig, 1878.
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| Auditory organs of the Vertebrata.
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| (386) A. Boettcher. "Bau u. Entwicklung d. Schnecke." Denkschriften d.
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| kaiserl. Leap. Carol. Akad. d. Wissenschaft., Vol. xxxv.
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| (387) C. Hasse. Dievergleich. Morphologieu. Histologied. hciutigen Gehororgane
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| (388) V. Hensen. "Zur Morphologie d. Schnecke." Zeit. f, wiss. ZooI.,Vo\.
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| XIII. 1863.
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| bebrliteten Kiichlein." Isis von Oken, 1831, and Meckel's Archiv, Vol. VI.
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| (390) Reissner. De Auris internee formatione. Inaug. Diss. Dorpat, 1851.
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| Accessory parts of Vertebrate Ear.
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| (391) David Hunt. "A comparative sketch of the development of the ear and
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| (392) W. Moldenhauer. "Zur Entwick. d. mittleren u. ausseren Ohres."
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| Morphol. Jahrbiich, Vol. ill. 1877.
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| (393) V. Urbantschitsch. " Ueb. d. erste Anlage d. Mittelohres u. d. Trommelfelles." Mittheil. a. d. embryol. Instit. Wien, Heft I. 1877.
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| | |
| OLFACTORY ORGAN.
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| (394) G. Born. "Die Nasenhohlen u. d. Thranennasengang d. amnioten
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| Wirbelthiere." Parts I. and II. Morphologisches Jahrbuch, Bd. V., 1879.
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| (395) A. Kolliker. " Ueber die Jacobson'schen Organe des Menschen."
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| Quart. Journ. of Micr. Science, Vol. xix., 1879.
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| SENSE-ORGANS OF THE LATERAL LINE.
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| (397) F. M. Balfour. A Monograph on the development of Elasmobranch Fishes,
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| pp. 141 146. London, 1878.
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| ORIGIN OF THE SKELETON.
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| BIBLIOGRAPHY. xvii
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| RIBS AND STERNUM.
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| | |
| T. H. Huxley. "On Ceratodus Forsteri." Proc. Zool. Soc., 1876.
| |
| | |
| T. H. Huxley. " The nature of the craniofacial apparatus of Petromyzon."
| |
| | |
| | |
| | |
| Journ. of Anat. and Phys., Vol. X. 1876.
| |
| | |
| (448) T. H. Huxley. The Anatomy of Vertebrated Animals. London, 1871.
| |
| | |
| (449) W. K. Parker. "On the structure and development of the skull of the
| |
| Common Fowl (Gallus Domesticus). " Phil. Trans., 1869.
| |
| | |
| (450) W. K. Parker. "On the structure and development of the skull of the
| |
| Common Frog (Rana temporaria)." Phil. Trans., 1871.
| |
| | |
| (451) W. K. Parker. "On the structure and development of the skull in the
| |
| Salmon (Salmo salar)." Bakerian Lecture, Phil. Trans., 1873.
| |
| | |
| (452) W. K. Parker. "On the structure and development of the skull in the
| |
| Pig (Susscrofa)." Phil. Trans., 1874.
| |
| | |
| (453) W. K. Parker. "On the structure and development of the skull in the
| |
| Batrachia." Part II. Phil. Trans., 1876.
| |
| | |
| (454) W. K. Parker. "On the structure and development of the skull in the
| |
| Urodelous Amphibia." Part in. Phil. Trans., 1877.
| |
| | |
| (455) W. K. Parker. "On the structure and development of the skull in the
| |
| Common Snake (Tropidonotus natrix)." Phil. Trans. , 1878.
| |
| | |
| (456) W. K. Parker. "On the structure and development of the skull in Sharks
| |
| and Skates." Trans. Zoolog. Soc., 1878. Vol. x. pt. iv.
| |
| | |
| (1.17) W. K. Parker. "On the structure and development of the skull in the
| |
| Lacertilia." Pt. I. Lacerta agilis, L. viridis and Zootoca vivipara. Phil. Trans.,
| |
| 1879.
| |
| | |
| | |
| | |
| BIBLIOGRAPHY,
| |
| | |
| | |
| | |
| (458) W. K. Parker. "The development of the Green Turtle." The Zoolo-v
| |
| of the Voyage of H.M.S. Challenger. Vol. I. pt. v.
| |
| | |
| (459) W. K. Parker. "The structure and development of the skull in the
| |
| Batrachia." 1't. in. Phil. Trans., 1880.
| |
| | |
| (460) W. K. Parker and G. T. Bettany. The Morphology of the Skull.
| |
| London, 1877.
| |
| | |
| (460*) H. Rathke. Entwick. d. Natter. Konigsberg, 1830.
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| | |
| (461) C. B. Reichert. " Ueber die Visceralbogen d. Wirbelthiere." Mailer's
| |
| Archiv, 1837.
| |
| | |
| (462) W. Salensky. " Beitrage z. Entwick. d. knorpeligen Gehorknochelchen."
| |
| Morphol. Jahrbuch, Vol. VI. 1880.
| |
| | |
| Vide also Kolliker (No. 298), especially for the human and mammalian skull;
| |
| Gotte (No. 296).
| |
| | |
| THE PECTORAL GIRDLE.
| |
| | |
| (463) Bruch. "Ueber die Entwicklung der Clavicula und die Farbe des
| |
| Blutes." Zeit.f. wiss. Zool., IV. 1853.
| |
| | |
| (464) A. Duges. " Recherches sur 1'osteologie et la myologie des Batraciens a
| |
| leurs differents ages." Memoires des savants etrang. Academic royale des sciences de
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| Finstitut de France, Vol. VI. 1835.
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| | |
| (465) C. Gegenbaur. Unterstichungen zur vergleichenden Anatomic der Wirbelthiere, i Heft. Schultergilrtel der Wirbelthiere. Brustflosse der Fische. Leipzig,
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| 1865.
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| | |
| (466) A. Gotte. "Beitrage z. vergleich. Morphol. d. Skeletsystems d. Wirbelthiere : Brustbien u. Schultergiirtel. " Archiv f. mikr. Anat. Vol. XIV. 1877.
| |
| | |
| (467) C. K. Hoffmann. "Beitrage z. vergleichenden Anatomic d. Wirbelthiere." Niederldndisches Archiv f. Zool. , Vol. V. 1879.
| |
| | |
| (468) W. K. Parker. " A Monograph on the Structure and Development of the
| |
| Shoulder-girdle and Sternum in the Vertebrata." Ray Society, 1868.
| |
| | |
| (469) H. Rathke. Ueber die Entwicklung der Schildkroten. Braunschweig,
| |
| 1848.
| |
| | |
| (470) H. Rathke. Ueber den Bau und die Entwicklung des Brustbeins der
| |
| Satirier, 1853.
| |
| | |
| (471) A. Sab a tier. Comparaison des ceintures et des menibres anteneurs et posterieurs d. la Serie d. Vertebrcs. Montpellier, 1880.
| |
| | |
| (472) Georg 'Swirski. Untersuch. lib. d. Entwick. d. Schultergiirtels u. d.
| |
| Skelets d. Brustflosse d. Hechts. Inaug. Diss. Dorpat, 1880.
| |
| | |
| THE PELVIC GIRDLE.
| |
| | |
| (473) A. Bunge. Untersuch. z. Entwick. d. Beckengilrtels d. Amphibien,
| |
| Reptilien u. Vdgel. Inaug. Diss. Dorpat, 1880.
| |
| | |
| (474) C. Gegenbaur. " Ueber d. Ausschluss des Schambeins von d. Pfanne
| |
| d. Hiiftgelenkes." Morph. Jahrbuch, Vol. II. 1876.
| |
| | |
| (475) Th. H. Huxley. "The characters of the Pelvis in Mammalia, etc."
| |
| Proc. of Roy. Soc., Vol. xxvin. 1879.
| |
| | |
| (476) A. S aba tier. Comparaison des ceintures et des membres anterieurs ct
| |
| postb-ieurs dans la Serie d. Vertebres. Montpellier, 1880.
| |
| | |
| SKELETON OF THE LIMBS.
| |
| | |
| (477) M. v. Davidoff. "Beitrage z. vergleich. Anat. d. hinteren Gliedmaassen
| |
| d. Fische I." Morphol. Jahrbuch, Vol. v. 1879.
| |
| | |
| (478) C. Gegenbaur. Untersuchungen z. vergleich. Anat. d. Wirbelthiere.
| |
| Leipzig, 18645. Erstes Heft. Carpus u. Tarsus. Zweites Heft. Brustflosse d.
| |
| Fische.
| |
| | |
| (479) C. Gegenbaur. "Ueb. d. Skelet d. Gliedmaassen d. Wirbelthiere im
| |
| Allgemeinen u. d. Hintergliedmaassen d. Selachier insbesondere." Jenaische Zeilschrift, Vol. V. 1870.
| |
| | |
| | |
| | |
| XX BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| (480) C. Gegenbaur. " Ueb. d. Archipterygium." Jenaische Zeitschrift, Vol.
| |
| vn. 1873.
| |
| | |
| (481) C. Gegenbaur. "Zur Morphologic d. Gliedmaassen d. Wirbelthiere."
| |
| Morphologisches Jahrbuch, Vol. II. 1876.
| |
| | |
| (482) A. Gotte. Ueb. Entwick. u. Regeneration d. Gliedmaassenskelets d. Molche.
| |
| Leipzig, 1879.
| |
| | |
| (483) T. H. Huxley. "On Ceratodus Forsteri, with some observations on the
| |
| classification of Fishes." Proc. Zool. Soc. 1876.
| |
| | |
| (484) St George Mivart. "On the Fins of Elasmobranchii." Zoological
| |
| Trans., Vol. x.
| |
| | |
| (485) A. Rosenberg. "Ueb. d. Entwick. d. Extremitaten-Skelets bei einigen
| |
| d. Reduction ihrer Gliedmaassen charakterisirten Wirbelthiere." Zeit.f. wiss. Zool.,
| |
| Vol. xxin. 1873.
| |
| | |
| (486) E. Rosenberg. "Ueb. d. Entwick. d. Wirbelsaule u. d. centrale carpi
| |
| d. Menschen." Morphologisches Jahrbuch, Vol. I. 1875.
| |
| | |
| (487) H. Strasser. "Z. Entwick. d. Extremitatenknorpel bei Salamandern u.
| |
| Tritonen." Morphologisches Jahrbuch, Vol. V. 1879.
| |
| | |
| (488) G. 'S wirski. Unterstich. iib. d. Entwick. d. Schnltergiirtels u. d. Skelets d.
| |
| Brustflosse d. Hechts. Inaug. Diss. Dorpat, 1880.
| |
| | |
| (489) J. K. Thacker. "Median and paired fins. A contribution to the history
| |
| of the Vertebrate limbs." Trans, oftke Connecticut Acad., Vol. III. 1877.
| |
| | |
| (490) J. K. Thacker. "Ventral fins of Ganoids." Trans, of the Connecticut
| |
| Acad., Vol. IV. 1877.
| |
| | |
| PLEURAL AND PERICARDIAL CAVITIES.
| |
| | |
| (491) M. Cadiat. " Du developpement de la partie cephalothoracique de 1'embryon, de la formation du diaphragme, des pleures, du pericarde, du pharynx et de
| |
| 1'cesophage." Journal de FAnatomie et de la Physiologic, Vol. xiv. 1878.
| |
| | |
| VASCULAR SYSTEM.
| |
| The Heart.
| |
| | |
| (492) A. C. Bernays. " Entwicklungsgeschichte d. Atrioventricularklappen."
| |
| Morphol. Jahrbuch, Vol. 11. 1876.
| |
| | |
| (493) E. Gasser. " Ueber d. Entstehung d. Herzens beim Hiihn." Archiv f.
| |
| mikr. Anat., Vol. xiv.
| |
| | |
| (494) A. Thomson. "On the development of the vascular system of the foetus
| |
| of Vertebrated Animals." Edinb. New Phil. Journal, Vol. ix. 1830 and 1831.
| |
| | |
| (495) M. Tonge. "Observations on the development of the semilunar valves
| |
| of the aorta and pulmonary artery of the heart of the Chick." Phil. Trans. CLIX.
| |
| 1869.
| |
| | |
| Vide also Von Baer (291), Rathke (300), Hensen (182), Kolliker (298), Gotte (296),
| |
| and Balfour (292).
| |
| | |
| The Arterial System.
| |
| | |
| (496) H. Rathke. "Ueb. d. Entwick. d. Arterien w. bei d. Saugethiere von
| |
| d. Bogen d. Aorta ausgehen." Miiller's Archiv, 1843.
| |
| | |
| (41)7) PI. Rathke. " Untersuchungen iib. d. Aortenwurzeln d. Saurier."
| |
| Denkschriften d. k. Akad. Wien, Vol. xiil. 1857.
| |
| | |
| Vide also His (No. 232) and general works on Vertebrate Embryology.
| |
| | |
| The Venous System.
| |
| | |
| (498) J.Marshall. "On the development of the great anterior veins." Phil.
| |
| Trans., 1859.
| |
| | |
| | |
| | |
| BIHLIOGRAI'IIY. XXJ
| |
| | |
| | |
| | |
| (499) H. Rathke. " Ueb. d. Bildung d. Pfortader u. d. Lebervenen b. Sauge
| |
| thieren." Meckel 's Archiv, 1830.
| |
| | |
| (500) H. Rathke. "Ueb. d. Bau u. d. Entwick. d. Venensystems d. Wirbclthiere." Bericht. iib. d. natttrh. Seminar, d. Univ. Konigsberg, 1838.
| |
| | |
| Vide also Von Baer (No. 291), Gotte (No. 296), Kolliker (No. 298), and Rathke
| |
| (Nos. 299, 300, and 301).
| |
| | |
| THE SPLEEN.
| |
| | |
| (501) W. Miiller. "The Spleen." Strieker's Histology.
| |
| | |
| (502) Peremeschko. "Ueb. d. Entwick. d. Milz." Silz. d. Wien. Akad.
| |
| Wiss., Vol. LVI. 1867.
| |
| | |
| THE SUPRARENAL BODIES.
| |
| | |
| (503) M. Braun. "Bau u. Entwick. d. Nebennieren bei Reptilian." Arbeit,
| |
| a. d. zool.-zoot. Institut Wilrzburg, Vol. v. 1879.
| |
| | |
| (504) A. v. Brunn. "Ein Beitrag z. Kenntniss d. feinern Baues u. d. Entwick.
| |
| d. Nebennieren." Archiv f. mikr. Anat., Vol. vni. 1872.
| |
| | |
| (505) Fr. Leydig. Untersuch. ilb. Fische u. Reptilien. Berlin, 1853.
| |
| | |
| (506) Fr. Leydig. Rochen u. Haie. Leipzig, 1852.
| |
| | |
| Vide also F. M. Balfour (No. 292), Kolliker (No. 298), Remak (No. 302), etc.
| |
| | |
| THE MUSCULAR SYSTEM OF THE VERTEBRATA.
| |
| | |
| (507) G.M.Humphry. " Muscles in Vertebrate Animals." J our n. of Anat.
| |
| and Phys., Vol. vi. 1872.
| |
| | |
| (508) J. Miiller. "Vergleichende Anatomic d. Myxinoiden. Part I. Osteologie
| |
| u. Myologie." Akad. Wiss., Berlin, 1834.
| |
| | |
| (509) A. M. Marshall. "On the head cavities and associated nerves of
| |
| Elasmobranchs." Quart. J. of Micr. Science, Vol. XXI. 1881.
| |
| | |
| (510) A. Schneider. "Anat. u. Entwick. d. Muskelsystems d. Wirbelthiere."
| |
| Sitz. d. Oberhessischen Gesellschaft, 1873.
| |
| | |
| (511) A. Schneider. Beitrdge z. vergleich. Anat. u. Entwick. d. Wirbelthiere.
| |
| Berlin, 1879.
| |
| | |
| Vide also Gotte (No. 296), Kolliker (No. 298), Balfour (No. 292), Huxley, etc.
| |
| | |
| EXCRETORY ORGANS.
| |
| | |
| INVER TEBRA TA .
| |
| | |
| (512) H. Eisig. " Die Segmentalorgane d. Capitelliden." Mitth. a. d. zool.
| |
| Slat. z. Neapel, Vol. I. 1879.
| |
| | |
| (513) J. Fraipont. " Recherches s. 1'appareil excreteur des Irematc
| |
| Cestoides." Archives de Biologie, Vol. I. 1880.
| |
| | |
| (514) B. Hatschek. "Studien iib. Entwick. d. Annehden. Arbeit, a. d.
| |
| zool. Instil. Wien, Vol. I. 1878. .
| |
| | |
| (515) B. Hatschek. "Ueber Entwick. von Echmrus, etc. Arbeit, a.
| |
| | |
| zool. Instit. Wien, Vol. ill. 1880.
| |
| | |
| VERTEBRATA.
| |
| | |
| General.
| |
| | |
| (516) F. M. Balfour. "On the origin and history of the urinogenital organs of
| |
| Vertebrates." Journal of Anat. and Phys., Vol. X. 1876.
| |
| | |
| | |
| | |
| XXJi BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| (517) Max. Fiirbringer 1 . "Zur vergleichenden Anat. u. Entwick. d. Excretionsorgane d. Vertebraten." Morphol. Jahrbuch, Vol. IV. 1878.
| |
| | |
| (518) H. Meckel. Zur Morphol. d. Harn- u. Geschlechtswerkz.d. Wirbelthiere,
| |
| etc. Halle, 1848.
| |
| | |
| (519) Job. Mtiller. Bildungsgeschichte d. Genitalien, etc. Diisseldorf, 1830.
| |
| | |
| (520) H. Ratbke. "Beobachtungen u. Betrachtungen ii. d. Entwicklung d.
| |
| Geschlechtswerkzeuge bei den Wirbelthieren." N. Schriften d. naturf. Gesell. in
| |
| Dantzig, Bd. I. 1825.
| |
| | |
| (521) C. Semper 1 . "Das Urogenitalsystem d. Plagiostomen u. seine Bedeutung f. d. ubrigen Wirbelthiere." Arb. a. d. zool.-zoot. Insiit. Wiirzburg, Vol. u.
| |
| | |
| 1875
| |
| (522) W. Waldeyer 1 . Eierstock u. Ei. Leipzig, 1870.
| |
| | |
| ElasmobrancJdi.
| |
| | |
| (523) A. Schultz. "Zur Entwick. d. Selachiereies." Archiv f. mikr. Anal.,
| |
| Vol. xi. 1875.
| |
| | |
| Vide also Semper (No. 521) and Balfour (No. 292).
| |
| | |
| Cyclostomata.
| |
| | |
| (524) J. M uller. " Untersuchungen ii. d. Eingeweide d. Fische. " Abh. d. k.
| |
| Ak. Wiss. Berlin, 1845.
| |
| | |
| (525) W. Muller. "Ueber d. Persistenz d. Urniere b. Myxine glutinosa."
| |
| Jenaische Zeitschrift, Vol. VII. 1873.
| |
| | |
| (526) W. Muller. "Ueber d. Urogenitalsystem d. Amphioxus u. d. Cyclostomen." Jenaische Zeitschrift, Vol. ix. 1875.
| |
| | |
| (527) A. Schneider. Beitrdge z. vergleich. Anat. u. Entwick. d. Wirbelthiere.
| |
| Berlin, 1879.
| |
| | |
| (528) W. B. Scott. "Beitrage z. Entwick. d. Petromyzonten." Morphol.
| |
| Jahrbuch, Vol. vn. 1881.
| |
| | |
| Teleostei.
| |
| | |
| (529) J. Hyrtl. "Das uropoetische System d. Knochenfische." Denkschr. d.
| |
| k. k. Akad. Wiss. Wien, Vol. II. 1850.
| |
| | |
| (530) A. Rosenberg. Untersuchungen iib. die Enlwicklung d. Teleostierniere.
| |
| Dorpat, 1867.
| |
| | |
| Vide also Oellacher (No. 72).
| |
| | |
| Amphibia.
| |
| | |
| (531) F. H. Bidder. Vergleichend-anatomische u. histologisclie Untcrsiiclniii^cn
| |
| ii. die mdnnlichcn Geschlec/its- tmd Harmverkzeuge d. nackten Amphibien. Dorpat,
| |
| 1846.
| |
| | |
| (532) C. L. Duvernoy. "Fragments s. les Organes genito-urinaires des
| |
| Reptiles," etc. Mem. Acad. Sciences. Paris. Vol. xi. 1851, pp. 17 95.
| |
| | |
| (533) M. Fiirbringer. Zur Entwicklung d. Amphibienniere. Heidelberg, 1877.
| |
| | |
| (534) F. Ley dig. Analomie d. Amphibien u. Keptilien. Berlin, 1853.
| |
| | |
| (535) F. Leydig. Lehrbuch d. Histologie. Hamm, 1857.
| |
| | |
| (536) F. Meyer. "Anat. d. Urogenitalsystems d. Selachier u. Amphibien."
| |
| Sitz. d. naturfor. Gesellsch. Leipzig, 1875.
| |
| | |
| (537) J. W. Spengel. "Das Urogenitalsystem d. Amphibien." Arb. a. d.
| |
| zool.- zoot. Instil. Wiirzburg. Vol. in. 1876.
| |
| | |
| (538) Von Wittich. "Harn- u. Geschlechtswerkzeuge d. Amphibien." Zeit.
| |
| f. wiss. Zool., Vol. iv.
| |
| | |
| Vide also Gotte (No. 296).
| |
| | |
| 1 The papers of Fiirbringer, Semper and Waldeyer contain full references to the
| |
| literature of the Vertebrate excretory organs.
| |
| | |
| | |
| | |
| BIBLIOGRAPHY. xxiii
| |
| | |
| | |
| | |
| Amniota.
| |
| | |
| (539) F. M. Balfour and A. Sedgwick. "On the existence of ahead-kidney
| |
| in the embryo Chick," etc. Quart. J. of Micr. Science, Vol. XIX. 1878.
| |
| | |
| (540) Banks. On the Wolffian bodies of the foetus and their remains in the adult.
| |
| Edinburgh, 1864.
| |
| | |
| (541) Th. Bornhaupt. UntersucJnmgen iib. die Entwicklung d. Urogenitalsystems beim Hiihnchen. Inaug. Diss. Riga, 1867.
| |
| | |
| (542) Max Braun. "Das Urogenitalsystem d. einheimischen Reptilien."
| |
| Arbeiten a. d. zool.-zoot. Instit. Wiirzburg. Vol. IV. 1877.
| |
| | |
| (543) J. Dansky u. J. Kostenitsch. " Ueb. d. Entwick. d. Keimblatter u. d.
| |
| Wolffschen Ganges im Htihnerei." Me"m. Acad. Imp. Petersbourg, vn. Series, Vol.
| |
| xxvn. 1880.
| |
| | |
| (544) Th. Egli. Beitrdge zur Anat. tmd Entiuick. d. Geschlechtsorgane. Inaug.
| |
| Diss. Zurich, 1876.
| |
| | |
| (545) E. Gasser. Beitrdge zur Entwickhmgsgeschichte d. Allantois, der
| |
| MiUler' schen Giinge u. des Afters. Frankfurt, 1874.
| |
| | |
| (546) E. Gasser. " Beob. iib. d. Entstehung d. WolfFschen Ganges bei Embryonen von Hiihnern u. Gansen." Arch, fiir mikr. Anat., Vol. xiv. 1877.
| |
| | |
| (547) E. Gasser. "Beitrage z. Entwicklung d. Urogenitalsystems d. Htihnerembryonen." Sitz. d. Cesell. zur Beforderung d. gesam. Naturwiss. Marburg, 1879.
| |
| | |
| (548) C. Kupffer. " Untersuchung liber die Entwicklung des Harn- und Geschlechtssystems." Archiv fiir mikr. Anat., Vol. II. 1866.
| |
| | |
| (549) A. Sedgwick. "Development of the kidney in its relation to the
| |
| Wolffian body in the Chick." Quart. J. of Micros. Science, Vol. XX. 1880.
| |
| | |
| (550) A. Sedgwick. "On the development of the structure known as the
| |
| glomerulus of the head -kidney in the Chick." Quart. J. of Micros. Science, Vol. XX.
| |
| 1880.
| |
| | |
| (551) A. Sedgwick. "Early development of the Wolffian duct and anterior
| |
| Wolffian tubules in the Chick ; with some remarks on the vertebrate excretory
| |
| system." Quart. J. of Micros. Science, Vol. xxi. 1881.
| |
| | |
| (552) M. Watson. "The homology of the sexual organs, illustrated by comparative anatomy and pathology." Journal of Anat. and Phys., Vol. XIV. 1879.
| |
| | |
| (553) E. H. Weber. Zusdtze z, Lehre von Bane u. d. Verrichtungen d. Geschlechtsorgane. Leipzig, 1846.
| |
| | |
| Vide also Remak (No. 302), Foster and Balfour (No. 295), His (No. 297),
| |
| Kolliker (No. 298).
| |
| | |
| GENERATIVE ORGANS.
| |
| | |
| (554) G. Balbiani. Lemons s. la generation des Vertebres. Paris, 1879.
| |
| | |
| (555) F. M. Balfour. "On the structure and development of the Vertebrate
| |
| ovary." Quart. J. of Micr. Science, Vol. XVIII.
| |
| | |
| (556) E. van Beneden. "De la distinction originelledutecticuleet del'ovaire,
| |
| etc." Bull. Ac. roy. belgique, Vol. xxxvn. 1874.
| |
| | |
| (557) N. Kleinenberg. "Ueb. d. Entstehung d. Eier b. Eudendrhim." Zeit.
| |
| f. wiss. Zool., Vol. xxxv. 1 88 r.
| |
| | |
| (558) H. Ludwig. "Ueb. d. Eibildung im Theirreiche. " Arbeit, a. d. zool.zoot. Instit. Wiirzburg, Vol. I. 1874.
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| (559) C. Semper. "Das Urogenitalsystem d. Plagiostomen, etc." Arbeit, a.
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| d. zool.-zoot. Instit. Wiirzburg, Vol. II. 1875.
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| | |
| (560) A. Weismann. "Zur Frage nach clem Ursprung d. Geschlechtszellen bei
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| den Hydroiden." Zool. Anzeiger, No. 55, 1880.
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| Vide also O. and R. Hertwig (No. 271), Kolliker (No. 298), etc.
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| ALIMENTARY CANAL AND ITS APPENDAGES.
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| (561) B. Afanassiew. " Ueber Bau u. Entwicklung d. Thymus d. Saugeth."
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| Archiv f. mikr. Anat. Bd. XIV. 1877.
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| XXIV BIBLIOGRAPHY.
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| (562) Fr. Boll. Das Princip d. Wachsthums. Berlin, 1876.
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| (563) E. Gasser. "Die Entstehung d. Cloakenoffhung hei Hiihneremhryonen."
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| Archiv f. Anat. u. Physiol., Anat. Abth. 1880.
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| | |
| (564) A. Gotte. Beitrage zur Entwicklungsgeschichte 'd. Darmkanah im
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| Hithnchcn. 1867.
| |
| | |
| (565) W. Miiller. " Ueber die Entwickelung der Schilddriise." ycnaische
| |
| Zeitschrift, Vol. vi. 1871.
| |
| | |
| (566) W. Miiller. "Die Hypobranchialrinne d. Tunicaten." Jenaischc Zeitschrift, Vol. VII. 1872.
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| | |
| (567) S. L. Schenk. "Die Bauchspeicheldriise d. Embryo." Anatomischphysiologische UntersucJnmgcn. 1872.
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| | |
| (568) E. Selenka. " Beitrag zur Entwicklungsgeschichte d. Luftsacke d.
| |
| Huhns." Zeit.f. wiss. Zool. 1866.
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| | |
| (569) L. Stieda. Untersuch. lib. d. Entivick. d. Glandula Thymus, Glandula
| |
| thyroidea, u. Glandula carotica. Leipzig, 1881.
| |
| | |
| (570) C. Fr. Wolff. " De formatione intestinorum." Nov. Comment. Akad.
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| Petrop. 1766.
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| | |
| (571) A. Wblfler. Ueb. d. Entwick. it. d. Ban d. Schilddriise. Berlin, 1880.
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| Vide also Kolliker (298), Qotte (296), His (232 and 297), Foster and Balfour (2!)5),
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| Balfour (292), Remak (302), Schenk (303), etc.
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| Teeth.
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| (572) T. H. Huxley. "On the enamel and dentine of teeth." Quart. J. of
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| Micros. Science, Vol. III. 1855.
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| (573) R. Owen. Odontography. London, 1840 1845.
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| | |
| (574) Ch. S. Tomes. Manual of dental anatomy, human and comparative.
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| London, 1876.
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| (575) Ch. S. Tomes. " On the development of teeth." Quart. J. of Micros.
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| Science, Vol. xvi. 1876.
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| (576) W. Waldeyer. " Structure and development of teeth." Strieker 's Histology. 1870.
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| Vide also Kolliker (298), Gegenbaur (294), Hertwig (306), etc.
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| {{Footer}}
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| [[Category:Historic Embryology]][[Category:1800's]]
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