The epiblast in the next stage is inflected for a small arc at the embryonic end of the blastoderm, where it becomes continuous with the lower layer cells; at the same time some of the lower layer cells of the embryonic end of the blastoderm assume a columnar form, and constitute the true hypoblast. The portion of the blastoderm, where epiblast and hypoblast are continuous, forms a projecting structure which will be called the embryonic rim ([fig. 16] B, er).
This rim is a very important structure, since it represents the dorsal portion of the lip of the blastopore of Amphioxus. The space between it and the yolk represents the commencing mesenteron, of which the hypoblast on the under side of the lip is the dorsal wall. The ventral wall of the mesenteron is at first formed solely of yolk held together by a protoplasmic network with numerous nuclei. The cavity under the lip becomes rapidly larger ([fig. 17], al), owing to the continuous conversion of lower layer cells into columnar hypoblast along an axial line passing from the middle of the embryonic rim towards the centre of the blastoderm. The continuous differentiation of the hypoblast towards the centre of the blastoderm corresponds with the invagination in Amphioxus. During the formation of the embryonic rim the blastoderm grows considerably larger, but, with the exception of the formation of the embryonic rim, retains its primitive constitution.
The segmentation cavity undergoes however important changes. There is formed below it a floor of lower layer cells, derived partly from an ingrowth from the two sides, but mainly from the formation of cells around the nuclei of the yolk ([fig. 16]). Shortly after the floor of cells has appeared, the whole segmentation cavity becomes obliterated ([fig. 17]).
The disappearance of the segmentation cavity corresponds in point of time with the formation of the hypoblast by the pseudo-invagination above described; and is probably due to this pseudo-invagination, in the same way that the disappearance of the segmentation cavity in Amphioxus is due to the true invagination of the hypoblast.
When the embryonic rim first appears there are no external indications of the embryo as distinguished from the blastoderm, but when it has attained to some importance the position of the embryo becomes marked out by the appearance of a shield-like area extending inwards from the edge of the embryonic rim, and formed of two folds with a groove between them ([fig. 28] B, mg), which is deepest at the edge of the blastoderm, and shallows out as it extends inwards. This groove is the medullary groove; and its termination at the edge of the blastoderm is placed at the hind end of the embryo.
At about the time of its appearance the mesoblast becomes first definitely established.
Fig. 18. Two transverse sections of an embryo of the same age as fig. 17.
A. Anterior section.
B. Posterior section.
mg. medullary groove; ep. epiblast; hy. hypoblast; n.al. cells formed round the nuclei of the yolk which have entered the hypoblast; m. mesoblast.
The sections shew the origin of the mesoblast.
At the edge of the embryonic rim the epiblast and lower layer cells are continuous. Immediately underneath the medullary groove, as is best seen in transverse section ([fig. 18]), the whole of the lower layer cells become converted into hypoblast, and along this line the columnar hypoblast is in contact with the epiblast above. At the sides however this is not the case; but at the junction of the epiblast and lower layer cells the latter remain undifferentiated. A short way from the edge the lower layer cells become divided into two distinct layers, a lower one continuous with the hypoblast in the middle line, and an upper one between this and the epiblast ([fig. 18] B). The upper layer is the commencement of the mesoblast (m). The mesoblast thus arises as two independent lateral plates, one on each side of the medullary groove, which are continuous behind with the undifferentiated lower layer cells at the edge of the embryonic rim. The mesoblast plates are at first very short, and do not extend to the front end of the embryo. They soon however grow forwards as two lateral ridges, attached to the hypoblast, one on each side of the medullary groove ([fig. 18] A, m). These ridges become separate from the hypoblast, and form two plates, thinner in front than behind; but still continuous at the edge of the blastoderm with the undifferentiated cells of the lip of the blastopore, and laterally with the lower layer cells of the non-embryonic part of the blastoderm. It results from the above mode of development of the mesoblast, that it may be described as arising in the form of a pair of solid outgrowths of the wall of the alimentary tract; which differ from the mesoblastic outgrowths of the wall of the archenteron in Amphioxus in not containing a prolongation of the alimentary cavity.