The space underneath the entoderm corresponds to the primitive gut-cavity, and is filled with the decreasing food-yelk (n). Thus the formation of the gastrula of our fish is complete. In contrast to the two chief forms of gastrula we considered previously, we give the name of discoid gastrula (discogastrula, Fig. 54) to this third principal type.
Very similar to the discoid gastrulation of the bony fishes is that of the hags or myxinoida, the remarkable cyclostomes that live parasitically in the body-cavity of fishes, and are distinguished by several notable peculiarities from their nearest relatives, the lampreys. While the amphiblastic ova of the latter are small and develop like those of the amphibia, the cucumber-shaped ova of the hag are about an inch long, and form a discoid gastrula. Up to the present it has only been observed in one species (Bdellostoma Stouti), by Dean and Doflein (1898).
It is clear that the important features which distinguish the discoid gastrula from the other chief forms we have considered are determined by the large food-yelk. This takes no direct part in the building of the germinal layers, and completely fills the primitive gut-cavity of the gastrula, even protruding at the mouth-opening. If we imagine the original bell-gastrula (Figs. 30–36) trying to swallow a ball of food which is much bigger than itself, it would spread out round it in discoid shape in the attempt, just as we find to be the case here (Fig. 54). Hence we may derive the discoid gastrula from the original bell-gastrula, through the intermediate stage of the hooded gastrula. It has arisen through the accumulation of a store of food-stuff at the vegetal pole, a “nutritive yelk” being thus formed in contrast to the “formative yelk.” Nevertheless, the gastrula is formed here, as in the previous cases, by the folding or invagination of the blastula. We can, therefore, reduce this cenogenetic form of the discoid segmentation to the palingenetic form of the primitive cleavage.
Fig. 53—Ovum-segmentation of a bony fish. A first cleavage of the stem-cell (cytula), B division of same into four segmentation-cells (only two visible), C the germinal disk divides into the blastoderm (b) and the periblast (p). d nutritive yelk, f fat-globule, c ovolemma, z space between the ovolemma and the ovum, filled with a clear fluid.)
This reduction is tolerably easy and confident in the case of the small ovum of our deep-sea bony fish, but it becomes difficult and uncertain in the case of the large ova that we find in the majority of the other fishes and in all the reptiles and birds. In these cases the food-yelk is, in the first place, comparatively colossal, the formative yelk being almost invisible beside it; and, in the second place, the food-yelk contains a quantity of different elements, which are known as “yelk-granules, yelk-globules, yelk-plates, yelk-flakes, yelk-vesicles,” and so on. Frequently these definite elements in the yelk have been described as real cells, and it has been wrongly stated that a portion of the embryonic body is built up from these cells. This is by no means the case. In every case, however large it is—and even when cell-nuclei travel into it during the cleavage of the border—the nutritive yelk remains a dead accumulation of food, which is taken into the gut during embryonic development and consumed by the embryo. The latter develops solely from the living formative yelk of the stem-cell. This is equally true of the ova of our small bony fishes and of the colossal ova of the primitive fishes, reptiles, and birds.
The gastrulation of the primitive fishes or selachii (sharks and rays) has been carefully studied of late years by Ruckert, Rabl, and H.E. Ziegler in particular, and is very important in the sense that this group is the oldest among living fishes, and their gastrulation can be derived directly from that of the cyclostoma by the accumulation of a large quantity of food-yelk. The oldest sharks (Cestracion) still have the unequal segmentation inherited from the cyclostoma. But while in this case, as in the case of the amphibia, the small ovum completely divides into cells in segmentation, this is no longer so in the great majority of the selachii (or Elasmobranchii). In these the contractility of the active protoplasm no longer suffices to break up the huge mass of the passive deutoplasm completely into cells; this is only possible in the upper or dorsal part, but not in the lower or ventral section. Hence we find in the primitive fishes a blastula with a small eccentric segmentation-cavity (Fig. 55 b), the wall of which varies greatly in composition. The circular border of the germinal disk which connects the roof and floor of the segmentation-cavity corresponds to the border-zone at the equator of the amphibian ovum. In the middle of its hinder border we have the beginning of the invagination of the primitive gut (Fig. 56 ud); it extends gradually from this spot (which corresponds to the Rusconian anus of the amphibia) forward and around, so that the primitive mouth becomes first crescent-shaped and then circular, and, as it opens wider, surrounds the ball of the larger food-yelk.
