The Evolution of Man, vol 1 by Ernst Haeckel (paper ebook reader .txt) 📕

nutritive yelk is of great indirect importance in embryonic development, though it has no direct share in it. It either does not divide at all, or only later on, and does not generally consist of cells. It is sometimes large and sometimes small, but generally many times larger than the formative yelk; and hence it is that it was formerly thought the more important of the two. As the respective significance of these two parts of the ovum is often wrongly described, it must be borne in mind that the nutritive yelk is only a secondary addition to the primary cell, it is an inner enclosure, not an external appendage. All ova that have this independent nutritive yelk are called, after Remak, “partially-cleaving” (meroblasta). Their segmentation is incomplete or partial.

(FIGURE 1.52. Ovum of a deep-sea bony fish. b protoplasm of the stem-cell, k nucleus of same, d clear globule of albumin, the nutritive yelk, f fat-globule of same, c outer membrane of the ovum, or ovolemma.)

There are many difficulties in the way of understanding this partial segmentation and the gastrula that arises from it. We have only recently succeeded, by means of comparative research, in overcoming these difficulties, and reducing this cenogenetic form of gastrulation to the original palingenetic type. This is comparatively easy in the small meroblastic ova which contain little nutritive yelk—for instance, in the marine ova of a bony fish, the development of which I observed in 1875 at Ajaccio in Corsica. I found them joined together in lumps of jelly, floating on the surface of the sea; and, as the little ovula were completely transparent, I could easily follow the development of the germ step by step. These ovula are glossy and colourless globules of little more than the 50th of an inch. Inside a structureless, thin, but firm membrane (ovolemma, Figure 1.52 c) we find a large, quite clear, and transparent globule of albumin (d). At both poles of its axis this globule has a pit-like depression. In the pit at the upper, animal pole (which is turned downwards in the floating ovum) there is a bi-convex lens composed of protoplasm, and this encloses the nucleus (k); this is the formative yelk of the stem-cell, or the germinal disk (b). The small fat-globule (f) and the large albumin-globule (d) together form the nutritive yelk. Only the formative yelk undergoes cleavage, the nutritive yelk not dividing at all at first.

The segmentation of the lens-shaped formative yelk (b) proceeds quite independently of the nutritive yelk, and in perfect geometrical order.

When the mulberry-like cluster of cells has been formed, the border-cells of the lens separate from the rest and travel into the yelk and the border-layer. From this the blastula is developed; the regular bi-convex lens being converted into a disk, like a watch-glass, with thick borders. This lies on the upper and less curved polar surface of the nutritive yelk like the watch glass on the yelk. Fluid gathers between the outer layer and the border, and the segmentation-cavity is formed. The gastrula is then formed by invagination, or a kind of turning-up of the edge of the blastoderm.

In this process the segmentation-cavity disappears.

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, Figure 1.54) 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 (Figures 1.30 to 1.36) 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 (Figure 1.54). 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.

(FIGURE 1.53. 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.

(FIGURE 1.54. Discoid gastrula (discogastrula) of a bony fish. e ectoderm, i entoderm, w border-swelling or primitive mouth, n albuminous globule of the nutritive yelk, f fat-globule of same, c external membrane (ovolemma), d partition between entoderm and ectoderm (earlier the segmentation-cavity).) 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 (Figure 1.55 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 (Figure 1.56 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.

Essentially different from the wide-mouthed discoid gastrula of most of the selachii is the narrow-mouthed discoid gastrula (or epigastrula) of the amniotes, the reptiles, birds, and monotremes; between the two—as an intermediate stage—we have the amphigastrula of the amphibia. The latter has developed from the amphigastrula of the ganoids and dipneusts, whereas the discoid amniote gastrula has been evolved from the amphibian gastrula by the addition of food-yelk.

This change of gastrulation is still found in the remarkable ophidia (Gymnophiona, Coecilia, or Peromela), serpent-like amphibia that live in moist soil in the tropics, and in many respects represent the transition from the gill-breathing amphibia to the lung-breathing reptiles. Their embryonic development has been explained by the fine studies of the brothers Sarasin of Ichthyophis glutinosa at Ceylon (1887), and those of August Brauer of the Hypogeophis rostrata in the Seychelles (1897). It is only by the historical and comparative study of these that we can understand the difficult and obscure gastrulation of the amniotes.

The bird’s egg is particularly important for our purpose, because most of the chief studies of the development of the vertebrates are based on observations of the hen’s egg during hatching. The mammal ovum is much more difficult to obtain and study, and for this practical and obvious reason very rarely thoroughly investigated. But we can get hens’ eggs in any quantity at any time, and, by means of artificial incubation, follow the development of the embryo step by step. The bird’s egg differs considerably from the tiny mammal ovum in size, a large quantity of food-yelk accumulating within the original yelk or the protoplasm of the ovum. This is the yellow ball which we commonly call the yolk of the egg. In order to understand the bird’s egg aright—for it is very often quite wrongly explained—we must examine it in its original condition, and follow it from the very beginning of its development in the bird’s ovary. We then see that the original ovum is a quite small, naked, and simple cell with a nucleus, not differing in either size or shape from the original ovum of the mammals and other animals (cf. Figure 1.13 E). As in the case of all the craniota (animals with a skull), the original or primitive ovum (protovum) is covered with a continuous layer of small cells. This membrane is the follicle, from which the ovum afterwards issues.

Immediately underneath it the structureless yelk-membrane is secreted from the yelk.

(FIGURE 1.55. Longitudinal section through the blastula of a shark (Pristiuris). (From Ruckert.) (Looked at from the left; to the right is the hinder end, H, to the left the fore end, V.) B

segmentation-cavity, kz cells of the germinal membrane, dk yelk-nuclei.

FIGURE 1.56. Longitudinal section of the blastula of a shark (Pristiurus) at the beginning of gastrulation. (From Ruckert.) (Seen from the left.) V fore end, H hind end, B segmentation-cavity, ud first trace of the primitive gut, dk yelk-nuclei, fd fine-grained yelk, gd coarse-grained yelk.)

The small primitive ovum of the bird begins very early to take up into itself a quantity of food-stuff through the yelk-membrane, and work it up into the “yellow yelk.” In this way the ovum enters on its second stage (the metovum), which is many times larger than the first, but still only a single enlarged cell. Through the accumulation of the store of