The Evolution of Man, V.1. by Ernst Haeckel (most read book in the world .TXT) 📖

Then eight new longitudinal lines are formed at the north pole, and these proceed to divide, first the darker cells above and afterwards the lighter southern cells, and finally reach the south pole. In this way we get in succession forty, forty-eight, fifty-six, and at last sixty-four cells (I, K). In the meantime, the two hemispheres differ more and more from each other. Whereas the sluggish lower hemisphere long remains at thirty-two cells, the lively northern hemisphere briskly sub-divides twice, producing first sixty-four and then 128 cells (L, M). Thus we reach a stage in which we count on the surface of the ovum 128 small cells in the upper half and thirty-two large ones in the lower half, or 160 altogether. The dissimilarity of the two halves increases: while the northern breaks up into a great number of small cells, the southern consists of a much smaller number of larger cells. Finally, the dark cells of the upper half grow almost over the surface of the ovum, leaving only a small circular spot at the south pole, where the large and clear cells of the lower half are visible. This white region at the south pole corresponds, as we shall see afterwards, to the primitive mouth of the gastrula. The whole mass of the inner and larger and clearer cells (including the white polar region) belongs to the entoderm or ventral layer. The outer envelope of dark smaller cells forms the ectoderm or skin-layer.

In the meantime, a large cavity, full of fluid, has been formed within the globular body--the segmentation-cavity or embryonic cavity (blastocoel, Figures 1.41 to 1.44 F). It extends considerably as the cleavage proceeds, and afterwards assumes an almost semi-circular form (Figure 1.41 F). The frog-embryo now represents a modified embryonic vesicle or blastula, with hollow animal half and solid vegetal half.

Now a second, narrower but longer, cavity arises by a process of folding at the lower pole, and by the falling away from each other of the white entoderm-cells (Figures 1.41 to 1.44 N). This is the primitive gut-cavity or the gastric cavity of the gastrula, progaster or archenteron. It was first observed in the ovum of the amphibia by Rusconi, and so called the Rusconian cavity. The reason of its peculiar narrowness here is that it is, for the most part, full of yelk-cells of the entoderm. These also stop up the whole of the wide opening of the primitive mouth, and form what is known as the "yelk-stopper," which is seen freely at the white round spot at the south pole (P). Around it the ectoderm is much thicker, and forms the border of the primitive mouth, the most important part of the embryo (Figure 1.44 k, k apostrophe). Soon the primitive gut-cavity stretches further and further at the expense of the segmentation-cavity (F), until at last the latter disappears altogether. The two cavities are only separated by a thin partition (Figure 1.43 s). With the formation of the primitive gut our frog-embryo has reached the gastrula stage, though it is clear that this cenogenetic amphibian gastrula is very different from the real palingenetic gastrula we have considered (Figures 1.30 to 1.36).

In the growth of this hooded gastrula we cannot sharply mark off the various stages which we distinguish successively in the bell-gastrula as morula and gastrula. Nevertheless, it is not difficult to reduce the whole cenogenetic or disturbed development of this amphigastrula to the true palingenetic formation of the archigastrula of the amphioxus.

(FIGURE 1.45. Blastula of the water-salamander (Triton). fh segmentation-cavity, dz yelk-cells, rz border-zone. (From Hertwig.)

FIGURE 1.46. Embryonic vesicle of triton (blastula), outer view, with the transverse fold of the primitive mouth (u). (From Hertwig.)

FIGURE 1.47. Sagittal section of a hooded-embryo (depula) of triton (blastula at the commencement of gastrulation). ak outer germinal layer, ik inner germinal layer, fh segmentation-cavity, ud primitive gut, u primitive mouth, dl and vl dorsal and ventral lips of the mouth, dz yelk-cells. (From Hertwig.))

This reduction becomes easier if, after considering the gastrulation of the tailless amphibia (frogs and toads), we glance for a moment at that of the tailed amphibia, the salamanders. In some of the latter, that have only recently been carefully studied, and that are phylogenetically older, the process is much simpler and clearer than is the case with the former and longer known. Our common water-salamander (Triton taeniatus) is a particularly good subject for observation. Its nutritive yelk is much smaller and its formative yelk less obscured with black pigment-cells than in the case of the frog; and its gastrulation has better retained the original palingenetic character. It was first described by Scott and Osborn (1879), and Oscar Hertwig especially made a careful study of it (1881), and rightly pointed out its great importance in helping us to understand the vertebrate development. Its globular blastula (Figure 1.45) consists of loosely-aggregated, yelk-filled entodermic cells or yelk-cells (dz) in the lower vegetal half; the upper, animal half encloses the hemispherical segmentation-cavity (fh), the curved roof of which is formed of two or three strata of small ectodermic cells. At the point where the latter pass into the former (at the equator of the globular vesicle) we have the border zone (rz). The folding which leads to the formation of the gastrula takes place at a spot in this border zone, the primitive mouth (Figure 1.46 u).

Unequal segmentation takes place in some of the cyclostoma and in the oldest fishes in just the same way as in most of the amphibia. Among the cyclostoma ("round-mouthed") the familiar lampreys are particularly interesting. In respect of organisation and development they are half-way between the acrania (lancelet) and the lowest real fishes (Selachii); hence I divided the group of the cyclostoma in 1886 from the real fishes with which they were formerly associated, and formed of them a special class of vertebrates. The ovum-segmentation in our common river-lamprey (Petromyzon fluviatilis) was described by Max Schultze in 1856, and afterwards by Scott (1882) and Goette (1890).

Unequal total segmentation follows the same lines in the oldest fishes, the selachii and ganoids, which are directly descended from the cyclostoma. The primitive fishes (Selachii), which we must regard as the ancestral group of the true fishes, were generally considered, until a short time ago, to be discoblastic. It was not until the beginning of the twentieth century that Bashford Dean made the important discovery in Japan that one of the oldest living fishes of the shark type (Cestracion japonicus) has the same total unequal segmentation as the amphiblastic plated fishes (ganoides).* (* Bashford Dean, Holoblastic Cleavage in the Egg of a Shark, Cestracion japonicus Macleay. Annotationes zoologicae japonenses, volume 4 Tokio 1901.) This is particularly interesting in connection with our subject, because the few remaining survivors of this division, which was so numerous in paleozoic times, exhibit three different types of gastrulation. The oldest and most conservative forms of the modern ganoids are the scaly sturgeons (Sturiones), plated fishes of great evolutionary importance, the eggs of which are eaten as caviar; their cleavage is not essentially different from that of the lampreys and the amphibia. On the other hand, the most modern of the plated fishes, the beautifully scaled bony pike of the North American rivers (Lepidosteus), approaches the osseous fishes, and is discoblastic like them. A third genus (Amia) is midway between the sturgeons and the latter.

(FIGURE 1.48. Sagittal section of the gastrula of the water-salamander (Triton). (From Hertwig.) Letters as in Figure 1.47; except--p yelk-stopper, mk beginning of the middle germinal layer.)

The group of the lung-fishes (Dipneusta or Dipnoi) is closely connected with the older ganoids. In respect of their whole organisation they are midway between the gill-breathing fishes and the lung-breathing amphibia; they share with the former the shape of the body and limbs, and with the latter the form of the heart and lungs. Of the older dipnoi (Paladipneusta) we have now only one specimen, the remarkable Ceratodus of East Australia; its amphiblastic gastrulation has been recently explained by Richard Semon (cf.

Chapter 2.

21). That of the two modern dipneusta, of which Protopterus is found in Africa and Lepidosiren in America, is not materially different. (Cf. Figure 1.51.)

(FIGURE 1.49. Ovum-segmentation of the lamprey (Petromyzon fluviatalis), in four successive stages. The small cells of the upper (animal) hemisphere divide much more quickly than the cells of the lower (vegetal) hemisphere.

FIGURE 1.50. Gastrulation of the lamprey (Petromyzon fluviatilis). A blastula, with wide embryonic cavity (blastocoel, bl), g incipient invagination. B depula, with advanced invagination, from the primitive mouth (g). C gastrula, with complete primitive gut: the embryonic cavity has almost disappeared in consequence of invagination.)

All these amphiblastic vertebrates, Petromyzon and Cestracion, Accipenser and Ceratodus, and also the salamanders and batrachia, belong to the old, conservative groups of our stem. Their unequal ovum-segmentation and gastrulation have many peculiarities in detail, but can always be reduced with comparative ease to the original cleavage and gastrulation of the lowest vertebrate, the amphioxus; and this is little removed, as we have seen, from the very simple archigastrula of the Sagitta and Monoxenia (see Figures 1.29 to 1.36). All these and many other classes of animals generally agree in the circumstance that in segmentation their ovum divides into a large number of cells by repeated cleavage. All such ova have been called, after Remak, "whole-cleaving" (holoblasta), because their division into cells is complete or total.

(FIGURE 1.51. Gastrulation of ceratodus (from Semon). A and C stage with four cells, B and D with sixteen cells. A and B are seen from above, C and D sideways. E stage with thirty-two cells; F blastula; G gastrula in longitudinal section. fh segmentation-cavity. gh primitive gut or gastric cavity.)

In a great many other classes of animals this is not the case, as we find (in the vertebrate stem) among the birds, reptiles, and most of the fishes; among the insects and most of the spiders and crabs (of the articulates); and the cephalopods (of the molluscs). In all these animals the mature ovum, and the stem-cell that arises from it in fertilisation, consist of two different and separate parts, which we have called formative yelk and nutritive yelk. The formative yelk alone consists of living protoplasm, and is the active, evolutionary, and nucleated part of the ovum; this alone divides in segmentation, and produces the numerous cells which make up the embryo. On the other hand, the nutritive yelk is merely a passive part of the contents of the ovum, a subordinate element which contains nutritive material (albumin, fat, etc.), and so represents in a sense the provision-store of the developing embryo. The latter takes a quantity of food out of this store, and finally consumes it all. Hence the 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