The Works of Francis Balfour 3-1
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Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.
- Cephalochorda | Urochorda | Elasmobranchii | Teleostei | Cyclostomata | Ganoidei | Amphibia | Aves | Reptilia | Mammalia | Comparison of the Formation of Germinal Layers and Early Stages in Vertebrate Development | Ancestral form of the Chordata | General Conclusions | Epidermis and Derivatives | The Nervous System | Organs of Vision | Auditory, Olfactory, and Lateral Line Sense Organs | Notochord, Vertebral Column, Ribs, and Sternum | The Skull | Pectoral and Pelvic Girdles and Limb Skeleton | Body Cavity, Vascular System and Glands | The Muscular System | Excretory Organs | Generative Organs and Genital Ducts | The Alimentary Canal and Appendages in Chordata
Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.
Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.
Foster M. and Sedgwick A. The Works of Francis Balfour Vol. IV. Plates (1885) MacMillan and Co., London.
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Vol. III. A Treatise on Comparative Embryology 2 (1885)
Chapter I. Cephalochorda
THE developmental history of the Chordata has been studied far more completely than that of any of the groups so far considered ; and the results which have been arrived at are of striking interest and importance. Three main subdivisions of this group can be recognized : (i) the Cephalochorda containing the single genus Amphioxus ; (2) the Urochorda or Tunicata ; and (3) the Vertebrata 1 . The members of the second and probably of the first of these groups have undergone degeneration, but at the same time the members of the first group especially undergo a less modified development than that of other Chordata.
Our knowledge of the development of Amphioxus is mainly due to Kowalevsky (Nos. 1 and 2). The ripe eggs appear to be dehisced into the branchial or atrial cavity, and to be transported thence through the branchial clefts into the pharynx, and so through the mouth to the exterior. (Kowalevsky, No. 1, and Marshall, No. 5.)
1 The term Vertebrata is often used to include the Cephalochorda. It is in many ways convenient to restrict its use to the forms which have at any rate some indications of vertebrae ; a restriction which has the further convenience of restoring to the term its original limitations. In the first volume of this work the term Craniata was used for the forms which I now propose to call Vertebrata.
Formation of the Layers
When laid the egg is about O'iO5 mm. in diameter. It is invested by a delicate membrane, and is somewhat opaque owing to the presence of yolk granules, which are however uniformly distributed through it, and proportionately less numerous than in the ova of most Chordata. Impregnation is external and the segmentation is nearly regular (fig. i). A small segmentation
FIG. i. THE SEGMENTATION OF AMPHIOXUS. (Copied from Kowalevsky. )
A. Stage with two equal segments.
B. Stage with four equal segments.
C. Stage after the four segments have become divided by an equatorial furrow into eight equal segments.
D. Stage in which a single layer of cells encloses a central segmentation cavity.
E. Somewhat older stage in optical section. sg. segmentation cavity.
cavity is visible at the stage with four segments, and increases during the remainder of the segmentation ; till at the close (fig. I E) the embryo consists of a blastosphere formed of a single layer of cells enclosing a large segmentation cavity. One side of the blastosphere next becomes invaginated, and during the process the embryo becomes ciliated, and commences to rotate. The cells forming the invaginated layer become gradually more columnar than the remaining cells, and constitute the hypoblast; and a structural distinction between the epiblast and hypoblast is thus established. In the course of the invagination the segmentation cavity becomes gradually obliterated, and the embryo first assumes a cup-shaped form with a wide blastopore, but soon becomes elongated, while the communication of the archenteron, or cavity of invagination, with the exterior is reduced to a small blastopore (fig. 2 A), placed at the pole of the long axis which the subsequent development shews to be the hinder end oj the
FIG. i. EMBRYOS OF AMPHIOXUS. (After Kowalevsky.)
The parts in black with white lines are epiblastic; the shaded parts are hypoblastic.
A. Gastrula stage in optical section.
B. Slightly later stage after the neural plate np has become differentiated, seen as a transparent object from the dorsal side.
C. Lateral view of a slightly older larva in optical section.
D. Dorsal view of an older larva with the neural canal completely closed except for a small pore (no) in front.
E. Older larva seen as a transparent object from the side.
bl. blastopore (which becomes in D the neurenteric canal) ; ne. neurenteric canal ; np. neural or medullary plate ; no. anterior opening of neural canal ; ch. notochord ; so 1 , so 11 , first and second mesoblastic somites.
embryo. The blastopore is often known in other Chordata as the anus of Rusconi. Before the invagination is completed the larva throws off the egg-membrane, and commences to lead a free existence.
Up to this stage the larva, although it has acquired a cylindrical elongated form, has only the structure of a simple two-layered gastrula; but the changes which next take place give rise on the one hand to the formation of the central nervous system, and on the other to the formation of the notochord and mesoblastic somites 1 . The former structure is developed from the epiblast and the two latter from the hypoblast.
The formation of the central nervous system commences with the flattening of the dorsal surface of the embryo. The flattened area forms a plate (fig. 2 B and fig. 3 A, /), extending backwards to the blastopore, which has in the meantime passed round to the dorsal surface. The sides of the plate become raised as two folds, which are most prominent posteriorly, and meet behind the blastopore, but shade off in front. The two folds next unite dorsally, so as to convert the previous groove into a canal 2 the neural or medullary canal. They unite first of all over the blastopore, and their line of junction extends from this point forwards (fig. 2 C, D, E). There is in this way formed a tube on the floor of which the blastopore opens behind, and which is itself open in front. Finally the medullary canal is formed for the whole length of the embryo. The anterior opening persists however for some time. The communication between the neural and alimentary tracts becomes interrupted when the caudal fin appears and the anus is formed. The neural canal then extends round the end of the notochord to the ventral side, but subsequently retreats to the dorsal side and terminates in a slight dilatation.
In the formation of the medullary canal there are two points deserving notice viz. (i) the connection with the blastopore; (2) the relation of the walls of the canal to the adjoining epiblast. With reference to the first of these points it is clear that the fact of the blastopore opening on the floor of the neural canal causes a free communication to exist between the archenteron or gastrula cavity and the neural canal ; and that, so long as the anterior pore of the neural canal remains open, the archenteron communicates indirectly with the exterior (vide fig. 2 E). It must not however be supposed (as has been done by some embryologists) that the pore at the front end of the neural canal represents the blastopore carried forwards. It is even probable that what Kowalevsky describes as the carrying of the blastopore to the dorsal side is really the commencement of the formation of the neural canal, the walls of which are continuous with the lips of the blastopore. This interpretation receives support from the fact that at a later stage, when the neural and alimentary canals become separated, the neural canal extends round the posterior end of the notochord to the ventral side. The embryonic communication between the neural and alimentary canals is common to most Chordata ; and the tube connecting them will be called the neurenteric canal. It is always formed in fundamentally the same manner as in Amphioxus. With reference to the second point it is to be noted that Amphioxus is exceptional amongst the Chordata in the fact that, before the closure of the neural groove, the layer of cells which will form the neural tube becomes completely separated from the adjoining epiblast (fig. 3 A), and forms a structure which may be spoken of as the medullary plate ; and that in the closure of the neural canal the lateral epiblast forms a complete layer above this plate before the plate itself is folded over into a closed canal. This peculiarity will be easily understood from an examination of fig. 3 A, B and C.
1 The protovertebrae of most embryologists will be spoken of as mesoblastic somites.
2 The details of this process are spoken of below.
FIG. 3. SECTIONS OF AN AMPHIOXUS EMBRYO AT THREE STAGES. (After Kowalevsky.)
A. Section at gastrula stage.
B. Section of an embryo slightly younger than that represented in fig. 2 D.
C. Section through the anterior part of an embryo at the stage represented in fig. 2 E.
np. neural plate ; nc. neural canal ; mes. archenteron in A and B, and mesenteron in C ; ch. notochord ; so. mesoblastic somite.
The formation of the mesoblastic somites commences, at about the same time as that of the neural canal, as a pair of hollow outgrowths of the walls of the archenteron. These outgrowths, which are shewn in surface view in fig. 2 B and D, so, and in section in fig. 3 B and C, so, arise near the front end of the body and gradually extend backwards as wing-like diverticula of the archenteric cavity. As they grow backwards their dorsal part becomes divided by transverse constrictions into cubical bodies (fig. 2 D and E), which, with the exception of the foremost, soon cease to open into what may now be called the mesenteron, and form the mesoblastic somites. Each mesoblastic somite, after its separation from the mesenteron, is constituted of two layers, an inner one the splanchnic and an outer the somatic, and a cavity between the two which was originally continuous with the cavity of the mesenteron. Eventually the dorsal parts of the outgrowths become separated from the ventral, and form the muscle-plates, while their cavities atrophy. The cavity of the ventral part, which is not divided into separate sections by the above described constrictions, remains as the true body cavity. The ventral part of the inner layer of the mesoblastic outgrowths gives rise to the muscular and connective tissue layers of the alimentary tract, and the dorsal part to a section of the voluntary muscular system. The ventral part of the outer layer gives rise to the somatic mesoblast, and the dorsal to a section of the voluntary muscular system. The anterior mesoblastic somite long retains its communication with the mesenteron, and was described by Max Schultze, and also at first by Kowalevsky, as a glandular organ. While the mesoblastic somites are becoming formed the dorsal wall of the mesenteron develops a median longitudinal fold (fig. 3 B, c/i), which is gradually separated off from before backwards as a rod (fig. 3 C, c/i), underlying the central nervous system. This rod is the notochord. After the separation of those parts the remainder of the hypoblast forms the wall of the mesenteron.
With the formation of the central nervous system, the mesoblastic somites, the notochord, and the alimentary tract the main systems of organs are established, and it merely remains briefly to describe the general changes of form which accompany the growth of the larva into the adult. By the time the larva is but twenty-four hours old there are formed about seventeen mesoblastic somites. The body, during the period in which these are being formed, remains cylindrical, but shortly afterwards it becomes pointed at both ends, and the caudal fin appears. The fine cilia covering the larva also become replaced by long cilia, one to each cell. The mesenteron is still completely closed, but on the right side of the body, at the level of the front end of the mesenteron, the hypoblast and epiblast now grow together, and a perforation becomes formed through their point of contact, which becomes the mouth. The anus is probably formed about the same time if not somewhat earlier 1 .
FIG. 4. SECTIONS THROUGH TWO ADVANCED EMBRYOS OF AMPHIOXUS TO SHEW THE FORMATION OF THE PERIBRANCHIAL CAVITY. (After Kowalevsky.)
In A are seen two folds of the body wall with a prolongation of the body cavity. In B the two folds have coalesced ventrally, forming a cavity into which a branchial cleft is seen to open.
tttes. mesenteron ; br.c. branchial cavity; //. body cavity.
Of the subsequent changes the two most important are (i) the formation of the gill slits or clefts ; (2) the formation of the peribranchial or atrial cavity.
The formation of the gill slits is, according to Kowalevsky's description, so peculiar that one is almost tempted to suppose that his observations were made on pathological specimens. The following is his account of the process. Shortly after the formation of the mouth there appears on the ventral line a coalescence between the epiblast and hypoblast. Here an opening is formed, and a visceral cleft is thus established, which passes to the left side, viz. the side opposite the mouth. A second and apparently a third slit are formed in the same way. The stages immediately following were not observed, but in the next stage twelve slits were present, no longer however on the left side, but in the median ventral line. There now appears on the side opposite the mouth, and the same therefore as that originally occupied by the first three clefts, a series of fresh clefts, which in their growth push the original clefts over to the same side as the mouth. Each of the fresh clefts becomes divided into two, which form the permanent clefts of their side.
1 The lateral position of the mouth in the embryo Amphioxus has been regarded as proving that the mouth represents a branchial cleft, but the general asymmetry of the organs is such that no great stress can, I think, be laid on the position of the mouth.
The gill slits at first open freely to the exterior, but during their formation two lateral folds of the body wall, containing a prolongation of the body cavity, make their appearance (fig. 4 A), and grow downwards over the gill clefts, and finally meet and coalesce along the ventral line, leaving a widish cavity between themselves and the body wall. Into this cavity, which is lined by epiblast, the gill clefts open (fig. 4 B, br.c). This cavity which forms a true peribranchial cavity is completely closed in front, but owing to the folds not uniting completely behind it remains in communication with the exterior by an opening known as the atrial or abdominal pore.
The vascular system of Amphioxus appears at about the same time as the first visceral clefts.
(1) A. Kowalevsky. " Entwicklungsgeschichte des Amphioxus lanceolatus." AIJHI. Acad. Imper. des Sciences de St Petersbourg, Series vil. Tom. xi. 1867.
(2) A. Kowalevsky. " Weitere Studien iiber die Entwicklungsgeschichte des Amphioxus lanceolatus." Archivf. mikr. Anat., Vol. XIII. 1877.
(3) Leuckart u. Pagenstecher. " Untersuchungen iiber niedere Seethiere." Mailer's Arckiv, 1858.
(4) Max Schultze. " Beobachtung junger Exemplare von Amphioxus." Zeit. f. wiss. Zool., Bd. in. 1851.
(5) A. M. Marshall. "On the mode of Oviposition of Amphioxus." your, of Anat. and Phys., Vol. X. 1876.