The Evolution of Man, V.1., Ernst Haeckel [ebook pdf reader for pc .TXT] 📗

of these cells form a sum of mechanical processes which in the last analysis are due to movements of the smallest vital parts, or the molecules, of the living substance. If we agree to call this active substance plasson, and its molecules plastidules, we may say that the individual physiological character of each of these cells is due to its molecular plastidule-movement. HENCE, THE PLASTIDULE-MOVEMENT OF THE CYTULA IS THE RESULTANT OF THE COMBINED PLASTIDULE-MOVEMENTS OF THE FEMALE OVUM AND THE MALE SPERM-CELL.* (* The plasson of the stem-cell or cytula may, from the anatomical point of view, be regarded as homogeneous and structureless, like that of the monera. This is not inconsistent with our hypothetical ascription to the plastidules (or molecules of the plasson) of a complex molecular structure. The complexity of this is the greater in proportion to the complexity of the organism that is developed from it and the length of the chain of its ancestry, or to the multitude of antecedent processes of heredity and adaptation.) CHAPTER VIII(8. THE GASTRAEA THEORY.)

 

There is a substantial agreement throughout the animal world in the first changes which follow the impregnation of the ovum and the formation of the stem-cell; they begin in all cases with the segmentation of the ovum and the formation of the germinal layers. The only exception is found in the protozoa, the very lowest and simplest forms of animal life; these remain unicellular throughout life. To this group belong the amoebae, gregarinae, rhizopods, infusoria, etc. As their whole organism consists of a single cell, they can never form germinal layers, or definite strata of cells. But all the other animals--all the tissue-forming animals, or metazoa, as we call them, in contradistinction to the protozoa--construct real germinal layers by the repeated cleavage of the impregnated ovum. This we find in the lower cnidaria and worms, as well as in the more highly-developed molluscs, echinoderms, articulates, and vertebrates.

In all these metazoa, or multicellular animals, the chief embryonic processes are substantially alike, although they often seem to a superficial observer to differ considerably. The stem-cell that proceeds from the impregnated ovum always passes by repeated cleavage into a number of simple cells. These cells are all direct descendants of the stem-cell, and are, for reasons we shall see presently, called segmentation-cells. The repeated cleavage of the stem-cell, which gives rise to these segmentation-spheres, has long been known as "segmentation." Sooner or later the segmentation-cells join together to form a round (at first, globular) embryonic sphere (blastula); they then form into two very different groups, and arrange themselves in two separate strata--the two primary germinal layers. These enclose a digestive cavity, the primitive gut, with an opening, the primitive mouth. We give the name of the gastrula to the important embryonic form that has these primitive organs, and the name of gastrulation to the formation of it. This ontogenetic process has a very great significance, and is the real starting-point of the construction of the multicellular animal body.

The fundamental embryonic processes of the cleavage of the ovum and the formation of the germinal layers have been very thoroughly studied in the last thirty years, and their real significance has been appreciated. They present a striking variety in the different groups, and it was no light task to prove their essential identity in the whole animal world. But since I formulated the gastraea theory in 1872, and afterwards (1875) reduced all the various forms of segmentation and gastrulation to one fundamental type, their identity may be said to have been established. We have thus mastered the law of unity which governs the first embryonic processes in all the animals.

Man is like all the other higher animals, especially the apes, in regard to these earliest and most important processes. As the human embryo does not essentially differ, even at a much later stage of development--when we already perceive the cerebral vesicles, the eyes, ears, gill-arches, etc.--from the similar forms of the other higher mammals, we may confidently assume that they agree in the earliest embryonic processes, segmentation and the formation of germinal layers. This has not yet, it is true, been established by observation. We have never yet had occasion to dissect a woman immediately after impregnation and examine the stem-cell or the segmentation-cells in her oviduct. However, as the earliest human embryos we have examined, and the later and more developed forms, agree with those of the rabbit, dog, and other higher mammals, no reasonable man will doubt but that the segmentation and formation of layers are the same in both cases.

But the special form of segmentation and layer formation which we find in the mammal is by no means the original, simple, palingenetic form. It has been much modified and cenogenetically altered by a very complex adaptation to embryonic conditions. We cannot, therefore, understand it altogether in itself. In order to do this, we have to make a COMPARATIVE study of segmentation and layer-formation in the animal world; and we have especially to seek the original, PALINGENETIC form from which the modified CENOGENETIC (see

Chapter 1.

1) form has gradually been developed.

This original unaltered form of segmentation and layer-formation is found to-day in only one case in the vertebrate-stem to which man belongs--the lowest and oldest member of the stem, the wonderful lancelet or amphioxus (cf.

Chapters

2.16 and 2.17). But we find a precisely similar palingenetic form of embryonic development in the case of many of the invertebrate animals, as, for instance, the remarkable ascidia, the pond-snail (Limnaeus), and arrow-worm (Sagitta), and many of the echinoderms and cnidaria, such as the common star-fish and sea-urchin, many of the medusae and corals, and the simpler sponges (Olynthus). We may take as an illustration the palingenetic segmentation and germinal layer-formation in an eight-fold insular coral, which I discovered in the Red Sea, and described as Monoxenia Darwinii.

(FIGURE 1.29. Gastrulation of a coral (Monoxenia Darwinii). A, B, stem-cell (cytula) or impregnated ovum. In Figure A (immediately after impregnation) the nucleus is invisible. In Figure B (a little later) it is quite clear. C two segmentation-cells. D four segmentation-cells. E mulberry-formation (morula). F blastosphere (blastula). G blastula (transverse section). H depula, or hollowed blastula (transverse section). I gastrula (longitudinal section). K gastrula, or cup-sphere, external appearance.)

The impregnated ovum of this coral (Figure 1.29 A, B) first splits into two equal cells (C). First, the nucleus of the stem-cell and its central body divide into two halves. These recede from and repel each other, and act as centres of attraction on the surrounding protoplasm; in consequence of this, the protoplasm is constricted by a circular furrow, and, in turn, divides into two halves. Each of the two segmentation-cells thus produced splits in the same way into two equal cells. The four segmentation-cells (grand-daughters of the stem-cell) lie in one plane. Now, however, each of them subdivides into two equal halves, the cleavage of the nucleus again preceding that of the surrounding protoplasm. The eight cells which thus arise break into sixteen, these into thirty-two, and then (each being constantly halved) into sixty-four, 128, and so on.* (* The number of segmentation-cells thus produced increases geometrically in the original gastrulation, or the purest palingenetic form of cleavage. However, in different animals the number reaches a different height, so that the morula, and also the blastula, may consist sometimes of thirty-two, sometimes of sixty-four, and sometimes of 128, or more, cells.) The final result of this repeated cleavage is the formation of a globular cluster of similar segmentation-cells, which we call the mulberry-formation or morula. The cells are thickly pressed together like the parts of a mulberry or blackberry, and this gives a lumpy appearance to the surface of the sphere (Figure E).* (* The segmentation-cells which make up the morula after the close of the palingenetic cleavage seem usually to be quite similar, and to present no differences as to size, form, and composition. That, however, does not prevent them from differentiating into animal and vegetative cells, even during the cleavage.)

When the cleavage is thus ended, the mulberry-like mass changes into a hollow globular sphere. Watery fluid or jelly gathers inside the globule; the segmentation-cells are loosened, and all rise to the surface. There they are flattened by mutual pressure, and assume the shape of truncated pyramids, and arrange themselves side by side in one regular layer (Figures F, G). This layer of cells is called the germinal membrane (or blastoderm); the homogeneous cells which compose its simple structure are called blastodermic cells; and the whole hollow sphere, the walls of which are made of the preceding, is called the blastula or blastosphere.* (* The blastula of the lower animals must not be confused with the very different blastula of the mammal, which is properly called the gastrocystis or blastocystis. This cenogenetic gastrocystis and the palingenetic blastula are sometimes very wrongly comprised under the common name of blastula or vesicula blastodermica.)

In the case of our coral, and of many other lower forms of animal life, the young embryo begins at once to move independently and swim about in the water. A fine, long, thread-like process, a sort of whip or lash, grows out of each blastodermic cell, and this independently executes vibratory movements, slow at first, but quicker after a time (Figure F). In this way each blastodermic cell becomes a ciliated cell. The combined force of all these vibrating lashes causes the whole blastula to move about in a rotatory fashion. In many other animals, especially those in which the embryo develops within enclosed membranes, the ciliated cells are only formed at a later stage, or even not formed at all. The blastosphere may grow and expand by the blastodermic cells (at the surface of the sphere) dividing and increasing, and more fluid is secreted in the internal cavity. There are still to-day some organisms that remain throughout life at the structural stage of the blastula--hollow vesicles that swim about by a ciliary movement in the water, the wall of which is composed of a single layer of cells, such as the volvox, the magosphaera, synura, etc. We shall speak further of the great phylogenetic significance of this fact in

Chapter 2.

19.

A very important and remarkable process now follows--namely, the curving or invagination of the blastula (Figure H). The vesicle with a single layer of cells for wall is converted into a cup with a wall of two layers of cells (cf. Figures G, H, I). A certain spot at the surface of the sphere is flattened, and then bent inward. This depression sinks deeper and deeper, growing at the cost of the internal cavity. The latter decreases as the hollow deepens. At last the internal cavity disappears altogether, the inner side of the blastoderm (that which lines the depression) coming to lie close on the outer side. At the same time, the cells of the two sections assume different sizes and shapes; the inner cells are more round and the outer more oval (Figure I). In this way the embryo takes the form of a cup or jar-shaped body, with a wall made up of two layers of cells, the inner cavity of which opens to the outside at one end (the spot where the depression was originally formed). We call this very