Special importance attaches to the fact that here again the four secondary germinal layers are already sharply distinct, and easily separated from each other. There is only one very restricted area in which they are connected, and actually pass into each other; this is the region of the primitive mouth, which is contracted in the amniotes into a dorsal longitudinal cleft, the primitive groove. Its two lateral lip-borders form the primitive streak, which has long been recognised as the most important embryonic source and starting-point of further processes. Sections through this primitive streak (Figs. 94 and 95) show that the two primary germinal layers grow at an early stage (in the discoid gastrula of the chick, a few hours after incubation) into the primitive streak (x), and that the two middle layers extend outward from this thickened axial plate (y) to the right and left between the former. The plates of the cœlom-layers, the parietal skin-fibre-layer (m) and the visceral gut-fibre-layer (f), are seen to be still pressed close together, and only diverge later to form the body-cavity. Between the inner borders of the two flat cœlom-pouches lies the chorda (Fig. 95 x), which here again develops from the middle line of the dorsal wall of the primitive gut.
Figs. 94 and 95—Transverse section of the primitive-streak (primitive mouth) of the chick. Fig. 94 a few hours after the commencement of incubation, Fig. 95 a little later. (From Waldeyer.) h horn-plate, n nerve-plate, m skin-fibre-layer, f gut-fibre-layer, d gut-gland-layer, y primitive streak or axial plate, in which all four germinal layers meet, x structure of the chorda, u region of the later primitive kidneys.
Cœlomation takes place in the vertebrates in just the same way as in the birds and reptiles. This was to be expected, as the characteristic gastrulation of the mammal has descended from that of the reptiles. In both cases a discoid gastrula with primitive streak arises from the segmented ovum, a two-layered germinal disk with long and small hinder primitive mouth. Here again the two primary germinal layers are only directly connected (Fig. 96 pr) along the primitive streak (at the folding-point of the blastula), and from this spot (the border of the primitive mouth) the middle germinal layers (mk) grow out to right and left between the preceding. In the fine illustration of the cœlomula of the rabbit which Van Beneden has given us (Fig. 96) one can clearly see that each of the four secondary germinal layers consists of a single stratum of cells.
Finally, we must point out, as a fact of the utmost importance for our anthropogeny and of great general interest, that the four-layered cœlomula of man has just the same construction as that of the rabbit (Fig. 96). A vertical section that Count Spee made through the primitive mouth or streak of a very young human germinal disk (Fig. 97) clearly shows that here again the four secondary germ-layers are inseparably connected only at the primitive streak, and that here also the two flattened cœlom-pouches (mk) extend outwards to right and left from the primitive mouth between the outer and inner germinal layers. In this case, too, the middle germinal layer consists from the first of two separate strata of cells, the parietal (mp) and visceral (mv) mesoblasts.
These concordant results of the best recent investigations (which have been confirmed by the observations of a number of scientists I have not enumerated) prove the unity of the vertebrate-stem in point of cœlomation, no less than of gastrulation. In both respects the invaluable amphioxus—the sole survivor of the acrania—is found to be the original model that has preserved for us in palingenetic form by a tenacious heredity these most important embryonic processes. From this primary model of construction we can cenogenetically deduce all the embryonic forms of the other vertebrates, the craniota, by secondary modifications. My thesis of the universal formation of the gastrula by folding of the blastula has now been clearly proved for all the vertebrates; so also has been Hertwig’s thesis of the origin of the middle germinal layers by the folding of a couple of cœlom-pouches which appear at the border of the primitive mouth. Just as the gastræa-theory explains the origin and identity of the two primary layers, so the cœlom-theory explains those of the four secondary layers. The point of origin is always the properistoma, the border of the original primitive mouth of the gastrula, at which the two primary layers pass directly into each other.
Fig. 96—Transverse section of the primitive groove (or primitive mouth) of a rabbit. (From Van Beneden.) pr primitive mouth, ul lips of same (primitive lips), ak and ik outer and inner germinal layers, mk middle germinal layer, mp parietal layer, mv visceral layer of the mesoderm.