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

important and interesting embryonic form the "cup-embryo" or "cup-larva" (gastrula, Figure 1.29, I longitudinal section, K external view). I have in my Natural History of Creation given the name of depula to the remarkable intermediate form which appears at the passage of the blastula into the gastrula. In this intermediate stage there are two cavities in the embryo--the original cavity (blastocoel) which is disappearing, and the primitive gut-cavity (progaster) which is forming.

I regard the gastrula as the most important and significant embryonic form in the animal world. In all real animals (that is, excluding the unicellular protists) the segmentation of the ovum produces either a pure, primitive, palingenetic gastrula (Figure 1.29 I, K) or an equally instructive cenogenetic form, which has been developed in time from the first, and can be directly reduced to it. It is certainly a fact of the greatest interest and instructiveness that animals of the most different stems--vertebrates and tunicates, molluscs and articulates, echinoderms and annelids, cnidaria and sponges--proceed from one and the same embryonic form. In illustration I give a few pure gastrula forms from various groups of animals (Figures 1.30 to 1.35, explanation given below each).

(FIGURES 1.30 TO 1.35. In each figure d is the primitive-gut cavity, o primitive mouth, s segmentation-cavity, i entoderm (gut-layer), e ectoderm (skin layer).

FIGURE 1.30. (A) Gastrula of a very simple primitive-gut animal or gastraead (gastrophysema). (Haeckel.)

FIGURE 1.31. (B) Gastrula of a worm (Sagitta). (From Kowalevsky.)

FIGURE 1.32. (C) Gastrula of an echinoderm (star-fish, Uraster), not completely folded in (depula). (From Alexander Agassiz.)

FIGURE 1.33. (D) Gastrula of an arthropod (primitive crab, Nauplius) (as 32).

FIGURE 1.34. (E) Gastrula of a mollusc (pond-snail, Linnaeus). (From Karl Rabl.)

FIGURE 1.35. (F) Gastrula of a vertebrate (lancelet, Amphioxus). (From Kowalevsky.) (Front view.))

In view of this extraordinary significance of the gastrula, we must make a very careful study of its original structure. As a rule, the typical gastrula is very small, being invisible to the naked eye, or at the most only visible as a fine point under very favourable conditions, and measuring generally 1/500 to 1/250 of an inch (less frequently 1/50 inch, or even more) in diameter. In shape it is usually like a roundish drinking-cup. Sometimes it is rather oval, at other times more ellipsoid or spindle-shaped; in some cases it is half round, or even almost round, and in others lengthened out, or almost cylindrical.

I give the name of primitive gut (progaster) and primitive mouth (prostoma) to the internal cavity of the gastrula-body and its opening; because this cavity is the first rudiment of the digestive cavity of the organism, and the opening originally served to take food into it. Naturally, the primitive gut and mouth change very considerably afterwards in the various classes of animals. In most of the cnidaria and many of the annelids (worm-like animals) they remain unchanged throughout life. But in most of the higher animals, and so in the vertebrates, only the larger central part of the later alimentary canal develops from the primitive gut; the later mouth is a fresh development, the primitive mouth disappearing or changing into the anus. We must therefore distinguish carefully between the primitive gut and mouth of the gastrula and the later alimentary canal and mouth of the fully developed vertebrate.* (* My distinction (1872) between the primitive gut and mouth and the later permanent stomach (metagaster) and mouth (metastoma) has been much criticised; but it is as much justified as the distinction between the primitive kidneys and the permanent kidneys. Professor E. Ray-Lankester suggested three years afterwards (1875) the name archenteron for the primitive gut, and blastoporus for the primitive mouth.)

(FIGURE 1.36. Gastrula of a lower sponge (olynthus). A external view, B longitudinal section through the axis, g primitive-gut cavity, a primitive mouth-aperture, i inner cell-layer (entoderm, endoblast, gut-layer), e external cell-layer (outer germinal layer, ectoderm, ectoblast, or skin-layer).

The two layers of cells which line the gut-cavity and compose its wall are of extreme importance. These two layers, which are the sole builders of the whole organism, are no other than the two primary germinal layers, or the primitive germ-layers. I have spoken in the introductory section (

Chapter 1.

3.) of their radical importance. The outer stratum is the skin-layer, or ectoderm (Figures 1.30 to 1.35 e); the inner stratum is the gut-layer, or entoderm (i). The former is often also called the ectoblast, or epiblast, and the latter the endoblast, or hypoblast. FROM THESE TWO PRIMARY GERMINAL LAYERS ALONE IS DEVELOPED THE ENTIRE ORGANISM OF ALL THE METAZOA OR MULTICELLULAR ANIMALS. The skin-layer forms the external skin, the gut-layer forms the internal skin or lining of the body. Between these two germinal layers are afterwards developed the middle germinal layer (mesoderma) and the body-cavity (coeloma) filled with blood or lymph.

The two primary germinal layers were first distinguished by Pander in 1817 in the incubated chick. Twenty years later (1849) Huxley pointed out that in many of the lower zoophytes, especially the medusae, the whole body consists throughout life of these two primary germinal layers. Soon afterwards (1853) Allman introduced the names which have come into general use; he called the outer layer the ectoderm ("outer-skin"), and the inner the entoderm ("inner-skin"). But in 1867 it was shown, particularly by Kowalevsky, from comparative observation, that even in invertebrates, also, of the most different classes--annelids, molluscs, echinoderms, and articulates--the body is developed out of the same two primary layers. Finally, I discovered them (1872) in the lowest tissue-forming animals, the sponges, and proved in my gastraea theory that these two layers must be regarded as identical throughout the animal world, from the sponges and corals to the insects and vertebrates, including man. This fundamental "homology [identity] of the primary germinal layers and the primitive gut" has been confirmed during the last thirty years by the careful research of many able observers, and is now pretty generally admitted for the whole of the metazoa.

As a rule, the cells which compose the two primary germinal layers show appreciable differences even in the gastrula stage. Generally (if not always) the cells of the skin-layer or ectoderm (Figures 1.36 c and 1.37 e) are the smaller, more numerous, and clearer; while the cells of the gut-layer, or entoderm (i), are larger, less numerous, and darker. The protoplasm of the ectodermic (outer) cells is clearer and firmer than the thicker and softer cell-matter of the entodermic (inner) cells; the latter are, as a rule, much richer in yelk-granules (albumen and fatty particles) than the former. Also the cells of the gut-layer have, as a rule, a stronger affinity for colouring matter, and take on a tinge in a solution of carmine, aniline, etc., more quickly and appreciably than the cells of the skin-layer. The nuclei of the entoderm-cells are usually roundish, while those of the ectoderm-cells are oval.

When the doubling-process is complete, very striking histological differences between the cells of the two layers are found (Figure 1.37). The tiny, light ectoderm-cells (e) are sharply distinguished from the larger and darker entoderm-cells (i). Frequently this differentiation of the cell-forms sets in at a very early stage, during the segmentation-process, and is already very appreciable in the blastula.

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, Figure 1.29), we find it in the lowest zoophyta (the gastrophysema, Figure 1.30), and the simplest sponges (olynthus, Figure 1.36); also in many of the medusae and hydrapolyps, lower types of worms of various classes (brachiopod, arrow-worm, Figure 1.31), tunicates (ascidia), many of the echinoderms (Figure 1.32), lower articulates (Figure 1.33), and molluscs (Figure 1.34), and, finally, in a slightly modified form, in the lowest vertebrate (the amphioxus, Figure 1.35).

(FIGURE 1.37. 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.

FIGURE 1.38. 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.))

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

2.16 and 2.17). 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 (Figure 1.29) 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 (Figure 1.38 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 (Figure 1.38 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 (Figure 1.38 D, E); it is most conspicuous when the invagination is complete and the segmentation-cavity has disappeared (Figure 1.38 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.

But the unequal gastrulation of the amphioxus diverges from the typical equal cleavage of the Sagitta, the Monoxenia (Figure 1.29), and the Olynthus (Figure 1.36), 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 (Figure 1.39 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