The Evolution of Man

by Ernst Haeckel

We have, up to the present, only considered that form of segmentation and gastrulation which, for many and weighty reasons, we may regard as the original, primordial, or palingenetic form. We might call it “equal” or homogeneous segmentation, because the divided cells retain a resemblance to each other at first (and often until the formation of the blastoderm). We give the name of the “bell-gastrula,” or archigastrula, to the gastrula that succeeds it. In just the same form as in the coral we considered (Monoxenia, Fig. 29), we find it in the lowest zoophyta (the gastrophysema, Fig. 30), and the simplest sponges (olynthus, Fig. 36); also in many of the medusæ and hydrapolyps, lower types of worms of various classes (brachiopod, arrow-worm, Fig. 31), tunicates (ascidia), many of the echinoderms (Fig. 32), lower articulates (Fig. 33), and molluscs (Fig. 34), and, finally, in a slightly modified form, in the lowest vertebrate (the amphioxus, Fig. 35).

Fig. 37—Cells from the two primary germinal layers of the mammal (from both layers of the blastoderm). i larger and darker cells of the inner stratum, the vegetal layer or entoderm. e smaller and clearer cells from the outer stratum, the animal layer or ectoderm.

The gastrulation of the amphioxus is especially interesting because this lowest and oldest of all the vertebrates is of the highest significance in connection with the evolution of the vertebrate stem, and therefore with that of man (compare Chapters XVI and XVII). Just as the comparative anatomist traces the most elaborate features in the structures of the various classes of vertebrates to divergent development from this simple primitive vertebrate, so comparative embryology traces the various secondary forms of vertebrate gastrulation to the simple, primary formation of the germinal layers in the amphioxus. Although this formation, as distinguished from the cenogenetic modifications of the vertebrate, may on the whole be regarded as palingenetic, it is nevertheless different in some features from the quite primitive gastrulation such as we have, for instance, in the Monoxenia (Fig. 29) and the Sagitta. Hatschek rightly observes that the segmentation of the ovum in the amphioxus is not strictly equal, but almost equal, and approaches the unequal. The difference in size between the two groups of cells continues to be very noticeable in the further course of the segmentation; the smaller animal cells of the upper hemisphere divide more quickly than the larger vegetal cells of the lower (Fig. 38 A, B). Hence the blastoderm, which forms the single-layer wall of the globular blastula at the end of the cleavage-process, does not consist of homogeneous cells of equal size, as in the Sagitta and the Monoxenia; the cells of the upper half of the blastoderm (the mother-cells of the ectoderm) are more numerous and smaller, and the cells of the lower half (the mother-cells of the entoderm) less numerous and larger. Moreover, the segmentation-cavity of the blastula (Fig. 38 C, h) is not quite globular, but forms a flattened spheroid with unequal poles of its vertical axis. While the blastula is being folded into a cup at the vegetal pole of its axis, the difference in the size of the blastodermic cells increases (Fig. 38 D, E); it is most conspicuous when the invagination is complete and the segmentation-cavity has disappeared (Fig. 38 F). The larger vegetal cells of the entoderm are richer in granules, and so darker than the smaller and lighter animal cells of the ectoderm.

Fig. 38—Gastrulation of the amphioxus, from Hatschek (vertical section through the axis of the ovum). A, B, C three stages in the formation of the blastula; D, E curving of the blastula; F complete gastrula. h segmentation-cavity. g primitive gut-cavity.

But the unequal gastrulation of the amphioxus diverges from the typical equal cleavage of the Sagitta, the Monoxenia (Fig. 29), and the Olynthus (Fig. 36), in another important particular. The pure archigastrula of the latter forms is uni-axial, and it is round in its whole length in transverse section. The vegetal pole of the vertical axis is just in the centre of the primitive mouth. This is not the case in the gastrula of the amphioxus. During the folding of the blastula the ideal axis is already bent on one side, the growth of the blastoderm (or the increase of its cells) being brisker on one side than on the other; the side that grows more quickly, and so is more curved (Fig. 39 v), will be the anterior or belly-side, the opposite, flatter side will form the back (d). The primitive mouth, which at first, in the typical archigastrula, lay at the vegetal pole of the main axis, is forced away to the dorsal side; and whereas its two lips lay at first in a plane at right angles to the chief axis, they are now so far thrust aside that their plane cuts the axis at a sharp angle. The dorsal lip is therefore the upper and more forward, the ventral lip the lower and hinder. In the latter, at the ventral passage of the entoderm into the ectoderm, there lie side by side a pair of very large cells, one to the right and one to the left (Fig. 39 p): these are the important polar cells of the primitive mouth, or “the primitive cells of the mesoderm.” In consequence of these considerable variations arising in the course of the gastrulation, the primitive uni-axial form of the archigastrula in the amphioxus has already become tri-axial, and thus the two-sidedness, or bilateral symmetry, of the vertebrate body has already been determined. This has been transmitted from the amphioxus to all the other modified gastrula-forms of the vertebrate stem.

Apart from this bilateral structure, the gastrula of the amphioxus resembles the typical archigastrula of the lower animals (Figs. 30–36) in developing the two primary germinal layers from a single layer of cells. This is clearly the oldest and original form of the metazoic embryo. Although the animals I have mentioned belong to the most diverse classes, they nevertheless agree with each other, and many more animal forms, in having retained to the present day, by a conservative heredity, this palingenetic form of gastrulation which they have from their earliest common ancestors. But this is not the case with the great majority of the animals. With these the original embryonic process has been gradually more or less altered in the course of millions of years by adaptation to new conditions of development. Both the segmentation of the ovum and the subsequent gastrulation have in this way been considerably changed. In fact, these variations have become so great in the course of time that the segmentation was not rightly understood in most animals, and the gastrula was unrecognised. It was not until I had made an extensive comparative study, lasting a considerable time (in the years 1866–75), in animals of the most diverse classes, that I succeeded in showing the same common typical process in these apparently very different forms of gastrulation, and tracing them all to one original form. I regard all those that diverge from the primary palingenetic gastrulation as secondary, modified, and cenogenetic. The more or less divergent form of gastrula that is produced may be called a secondary, modified gastrula, or a metagastrula. The reader will find a scheme of these different kinds of segmentation and gastrulation at the close of this chapter.