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

outer, structureless envelope that encloses it is the original ovolemma or zona pellucida, modified, and clothed with a layer of albumin that has been deposited on the outside. From this stage the envelope is called the external membrane, the primary chorion or prochorion (a). The real wall of the vesicle enclosed by it consists of a simple layer of ectodermic cells (b), which are flattened by mutual pressure, and generally hexagonal; a light nucleus shines through their fine-grained protoplasm (Figure 1.108). At one part (c) inside this hollow ball we find a circular disc, formed of darker, softer, and rounder cells, the dark-grained entodermic cells (Figure 1.109).

(FIGURE 1.108. Four entodermic cells from the embryonic vesicle of the rabbit.

FIGURE 1.109. Two entodermic cells from the embryonic vesicle of the rabbit.)

The characteristic embryonic form that the developing mammal now exhibits has up to the present usually been called the “blastula”

(Bischoff), “sac-shaped embryo” (Baer), “vesicular embryo” (vesicula blastodermica, or, briefly, blastosphaera). The wall of the hollow vesicle, which consists of a single layer of cells, was called the “blastoderm,” and was supposed to be equivalent to the cell-layer of the same name that forms the wall of the real blastula of the amphioxus and many of the invertebrates (such as Monoxenia, Figure 1.29 F, G). Formerly this real blastula was generally believed to be equivalent to the embryonic vesicle of the mammal. However, this is by no means the case. What is called the “blastula” of the mammal and the real blastula of the amphioxus and many of the invertebrates are totally different embryonic structures. The latter (blastula) is palingenetic, and precedes the formation of the gastrula. The former (blastodermic vesicle) is cenogenetic, and follows gastrulation. The globular wall of the blastula is a real blastoderm, and consists of homogeneous (blastodermic) cells; it is not yet differentiated into the two primary germinal layers. But the globular wall of the mammal vesicle is the differentiated ectoderm, and at one point in it we find a circular disk of quite different cells—the entoderm. The round cavity, filled with fluid, inside the real blastula is the segmentation-cavity. But the similar cavity within the mammal vesicle is the yelk-sac cavity, which is connected with the incipient gut-cavity. This primitive gut-cavity passes directly into the segmentation-cavity in the mammals, in consequence of the peculiar cenogenetic changes in their gastrulation, which we have considered previously (Chapter 1.9). For these reasons it is very necessary to recognise the secondary embryonic vesicle in the mammal (gastrocystis or blastocystis) as a characteristic structure peculiar to this class, and distinguish it carefully from the primary blastula of the amphioxus and the invertebrates.

(FIGURE 1.110. Ovum of a rabbit from the uterus, one sixth of an inch in diameter. The embryonic vesicle (b) has withdrawn a little from the smooth ovolemma (a). In the middle of the ovolemma we see the round germinal disk (blastodiscus, c), at the edge of which (at d) the inner layer of the embryonic vesicle is already beginning to expand.

(Figures 1.110 to 1.114 from Bischoff.) FIGURE 1.111. The same ovum, seen in profile. Letters as in Figure 1.110.

FIGURE 1.112. Ovum of a rabbit from the uterus, one-fourth of an inch in diameter. The blastoderm is already for the most part two-layered (b). The ovolemma, or outer envelope, is tufted (a).

FIGURE 1.113. The same ovum, seen in profile. Letters as in Figure 1.112.

FIGURE 1.114. Ovum of a rabbit from the uterus, one-third of an inch in diameter. The embryonic vesicle is now nearly everywhere two-layered (k) only remaining one-layered below (at d).

FIGURE 1.115. Round germinative area of the rabbit, divided into the central light area (area pellucida) and the peripheral dark area (area opaca). The light area seems darker on account of the dark ground appearing through it.)

The small, circular, whitish, and opaque spot which the gastric disk (Figure 1.106) forms at a certain part of the surface of the clear and transparent embryonic vesicle has long been known to science, and compared to the germinal disk of the birds and reptiles. Sometimes it has been called the germinal disk, sometimes the germinal spot, and usually the germinative area. From the area the further development of the embryo proceeds. However, the larger part of the embryonic vesicle of the mammal is not directly used for building up the later body, but for the construction of the temporary umbilical vesicle. The embryo separates from this in proportion as it grows at its expense; the two are only connected by the yelk-duct (the stalk of the yelk-sac), and this maintains the direct communication between the cavity of the umbilical vesicle and the forming visceral cavity (Figure 1.105).

The germinative area or gastric disk of the animal consists at first (like the germinal disk of birds and reptiles) merely of the two primary germinal layers, the ectoderm and entoderm. But soon there appears in the middle of the circular disk between the two a third stratum of cells, the rudiment of the middle layer or fibrous layer (mesoderm). This middle germinal layer consists from the first, as we have seen in Chapter 1.10, of two separate epithelial plates, the two layers of the coelom-pouches (parietal and visceral). However, in all the amniotes (on account of the large formation of yelk) these thin middle plates are so firmly pressed together that they seem to represent a single layer. It is thus peculiar to the amniotes that the middle of the germinative area is composed of four germinal layers, the two limiting (or primary) layers and the middle layers between them (Figures 1.96 and 1.97). These four secondary germinal layers can be clearly distinguished as soon as what is called the sickle-groove (or “embryonic sickle”) is seen at the hind border of the germinative area. At the borders, however, the germinative area of the mammal only consists of two layers. The rest of the wall of the embryonic vesicle consists at first (but only for a short time in most of the mammals) of a single layer, the outer germinal layer.

(FIGURE 1.116. Oval area, with the opaque whitish border of the dark area without.)

From this stage, however, the whole wall of the embryonic vesicle becomes two-layered. The middle of the germinative area is much thickened by the growth of the cells of the middle layers, and the inner layer expands at the same time, and increases at the border of the disk all round. Lying close on the outer layer throughout, it grows over its inner surface at all points, covers first the upper and then the lower hemisphere, and at last closes in the middle of the inner layer (Figures 1.110 to 1.114). The wall of the embryonic vesicle now consists throughout of two layers of cells, the ectoderm without and the entoderm within. It is only in the centre of the circular area, which becomes thicker and thicker through the growth of the middle layers, that it is made up of all four layers. At the same time, small structureless tufts or warts are deposited on the surface of the outer ovolemma or prochorion, which has been raised above the embryonic vesicle (Figures 1.112 to 1.114 a).

(FIGURE 1.117. Oval germinal disk of the rabbit, magnified about ten times. As the delicate, half-transparent disk lies on a black ground, the pellucid area looks like a dark ring, and the opaque area (lying outside it) like a white ring. The oval shield in the centre also looks whitish, and in its axis we see the dark medullary groove. (From Bischoff.))

We may now disregard both the outer ovolemma and the greater part of the vesicle, and concentrate our attention on the germinative area and the four-layered embryonic disk. It is here alone that we find the important changes which lead to the differentiation of the first organs. It is immaterial whether we examine the germinative area of the mammal (the rabbit, for instance) or the germinal disk of a bird or a reptile (such as a lizard or tortoise). The embryonic processes we are now going to consider are essentially the same in all members of the three higher classes of vertebrates which we call the amniotes.

Man is found to agree in this respect with the rabbit, dog, ox, etc.; and in all these animals the germinative area undergoes essentially the same changes as in the birds and reptiles. They are most frequently and accurately studied in the chick, because we can have incubated hens’ eggs in any quantity at any stage of development.

Moreover, the round germinal disk of the chick passes immediately after the beginning of incubation (within a few hours) from the two-layered to the four-layered stage, the two-layered mesoderm developing from the median primitive groove between the ectoderm and entoderm (Figures 1.82 to 1.95).

The first change in the round germinal disk of the chick is that the cells at its edges multiply more briskly, and form darker nuclei in their protoplasm. This gives rise to a dark ring, more or less sharply set off from the lighter centre of the germinal disk (Figure 1.115).

From this point the latter takes the name of the “light area” (area pellucida), and the darker ring is called the “dark area” (area opaca). (In a strong light, as in Figures 1.115 to 1.117, the light area seems dark, because the dark ground is seen through it; and the dark area seems whiter). The circular shape of the area now changes into elliptic, and then immediately into oval (Figures 1.116 and 1.117). One end seems to be broader and blunter, the other narrower and more pointed; the former corresponds to the anterior and the latter to the posterior section of the subsequent body. At the same time, we can already trace the characteristic bilateral form of the body, the antithesis of right and left, before and behind. This will be made clearer by the “primitive streak,” which appears at the posterior end.

(FIGURE 1.118. Pear-shaped germinal shield of the rabbit (eight days old), magnified twenty times. rf medullary groove. pr primitive groove (primitive mouth). (From Kolliker.)

FIGURE 1.119. Median longitudinal section of the gastrula of four vertebrates. (From Rabl.) A discogastrula of a shark (Pristiurus). B

amphigastrula of a sturgeon (Accipenser). C amphigastrula of an amphibium (Triton). D epigastrula of an amniote (diagram). a ventral, b dorsal lip of the primitive mouth.)

At an early stage an opaque spot is seen in the middle of the clear germinative area, and this also passes from a circular to an oval shape. At first this shield-shaped marking is very delicate and barely perceptible; but it soon becomes clearer, and now stands out as an oval shield, surrounded by two rings or areas (Figure 1.117). The inner and brighter ring is the remainder of the pellucid area, and the dark outer ring the remainder of the opaque area; the opaque shield-like spot itself is the first rudiment of the dorsal part of the embryo. We give it briefly the name of embryonic shield or dorsal shield. In most works this embryonic shield is described as “the first rudiment or trace of the embryo,” or “primitive embryo.” But this is wrong, though it rests on the authority of Baer and Bischoff. As a matter of fact, we already have the embryo in the stem-cell, the gastrula, and all the subsequent stages. The embryonic shield is simply the first rudiment of the dorsal part, which is the earliest to develop. As the older names of “embryonic rudiment” and “germinative area” are used in many different senses—and this has led to a fatal confusion in embryonic literature—we must explain very clearly the real significance of these important embryonic parts of the amniote.

It will be useful to do so in a series of formal principles:—

1. The so-called “first trace of the embryo” in the amniotes, or the embryonic shield, in the centre of the pellucid area,