The whole embryonic area would seem to be employed in the formation of the body of the embryo. Its long axis has no very definite relation to that of the blastodermic vesicle. The first external trace of the embryo to appear is the medullary plate, bounded by the medullary folds, and occupying at first the anterior half of the embryonic area ([fig. 141]). The two medullary folds diverge behind and enclose the front end of the primitive streak. As the embryo elongates, the medullary folds nearly meet behind and so cut off the front portion of the primitive streak, which then appears as a projection in the hind end of the medullary groove. In an embryo rabbit, eight days after impregnation, the medullary groove is about 1.80 mm. in length. At this stage a division may be clearly seen in the lateral plates of mesoblast into a vertebral zone adjoining the embryo and a more peripheral lateral zone; and in the vertebral zone indications of two somites, about 0.37 mm. from the hinder end of the embryo, become apparent. The foremost of these somites marks the junction, or very nearly so, of the cephalic region and trunk. The small size of the latter as compared with the former is very striking, but is characteristic of Vertebrates generally. The trunk gradually elongates relatively to the head, by the addition behind of fresh somites. The embryo has not yet begun to be folded off from the yolk-sack. In a slightly older embryo of nine days there appears (Hensen, Kölliker) round the embryonic area a delicate clear ring which is narrower in front than behind ([fig. 144] A, ap). This ring is regarded by these authors as representing the peripheral part of the area pellucida of Birds, which does not become converted into the body of the embryo. Outside the area pellucida, an area vasculosa has become very well defined. In the embryo itself ([fig. 144] A) the disproportion between head and trunk is less marked than before; the medullary plate dilates anteriorly to form a spatula-shaped cephalic enlargement; and three or four somites are established. In the lateral parts of the mesoblast of the head there may be seen on each side a tube-like structure (hz). Each of these is part of the heart, which arises as two independent tubes. The remains of the primitive streak (pr) are still present behind the medullary groove.
In somewhat older embryos ([fig. 144] B) with about eight somites, in which the trunk considerably exceeds the head in length, the first distinct traces of the folding-off of the head end of the embryo become apparent, and somewhat later a fold also appears at the hind end. In the formation of the hind end of the embryo the primitive streak gives rise to a tail swelling and to part of the ventral wall of the postanal gut. In the region of the head the rudiments of the heart (h) are far more definite. The medullary groove is still open for its whole length, but in the head it exhibits a series of well-marked dilatations. The foremost of these (vh) is the rudiment of the fore-brain, from the sides of which there project the two optic vesicles (ab); the next is the mid-brain (mh), and the last is the hind-brain (hh), which is again divided into smaller lobes by successive constrictions. The medullary groove behind the region of the somites dilates into an embryonic sinus rhomboidalis like that of the Bird. Traces of the amnion (af) are now apparent both in front of and behind the embryo.
Fig. 144. Embryo Rabbits of about nine days from the dorsal side. (From Kölliker.)
A. magnified 22 times, and B. 21 times.
ap. area pellucida; rf. medullary groove; h´. medullary plate in the region of the future fore-brain; h´´. medullary plate in the region of the future mid-brain; vh. fore-brain; ab. optic vesicle; mh. mid-brain; hh. and h´´´. hind-brain; uw. mesoblastic somite; stz. vertebral zone; pz. lateral zone; hz. and h. heart; ph. pericardial section of body cavity; vo. vitelline vein; af. amnion fold.
The structure of the head and the formation of the heart at this age are illustrated in [fig. 145]. The widely-open medullary groove (rf) is shewn in the centre. Below it the hypoblast is thickened to form the notochord dd´; and at the sides are seen the two tubes, which, on the folding-in of the foregut, give rise to the unpaired heart. Each of these is formed of an outer muscular tube of splanchnic mesoblast (ahh), not quite closed towards the hypoblast, and an inner epithelioid layer (ihh); and is placed in a special section of the body cavity (ph), which afterwards forms the pericardial cavity.
Fig. 145. Transverse section through the head of a Rabbit of the same age as fig. 144 b. (From Kölliker.)
B. is a more highly magnified representation of part of A.
rf. medullary groove; mp. medullary plate; rw. medullary fold; h. epiblast; dd. hypoblast; dd´. notochordal thickening of hypoblast; sp. undivided mesoblast; hp. somatic mesoblast; dfp. splanchnic mesoblast; ph. pericardial section of body cavity; ahh. muscular wall of heart; ihh. epithelioid layer of heart; mes. lateral undivided mesoblast; sw. fold of hypoblast which will form the ventral wall of the pharynx; sr. commencing throat.
Before the ninth day is completed great external changes are usually effected. The medullary groove becomes closed for its whole length with the exception of a small posterior portion. The closure commences, as in Birds, in the region of the mid-brain. Anteriorly the folding-off of the embryo proceeds so far that the head becomes quite free, and a considerable portion of the throat, ending blindly in front, becomes established. In the course of this folding the, at first widely separated, halves of the heart are brought together, coalesce on the ventral side of the throat, and so give rise to a median undivided heart. The fold at the tail end of the embryo progresses considerably, and during its advance the allantois is formed in the same way as in Birds. The somites increase in number to about twelve. The amniotic folds nearly meet above the embryo.