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

two stages rather agree in this.

When the blastula is fully formed, we have again in this case the important folding or invagination that determines gastrulation. The space between the skin-layer and the gut-layer (the remainder of the segmentation-cavity) remains full of food-yelk, which is gradually used up. This is the only material difference between our vesicular gastrula (perigastrula) and the original form of the bell-gastrula (archigastrula). Clearly the one has been developed from the other in the course of time, owing to the accumulation of food-yelk in the centre of the ovum.* (* On the reduction of all forms of gastrulation to the original palingenetic form see especially the lucid treatment of the subject in Arnold Lang's Manual of Comparative Anatomy (1888), Part 1.)

We must count it an important advance that we are thus in a position to reduce all the various embryonic phenomena in the different groups of animals to these four principal forms of segmentation and gastrulation. Of these four forms we must regard one only as the original palingenetic, and the other three as cenogenetic and derivative. The unequal, the discoid, and the superficial segmentation have all clearly arisen by secondary adaptation from the primary segmentation; and the chief cause of their development has been the gradual formation of the food-yelk, and the increasing antithesis between animal and vegetal halves of the ovum, or between ectoderm (skin-layer) and entoderm (gut-layer).

(FIGURE 1.72. Gastrula of the placental mammal (epigastrula from the rabbit), longitudinal section through the axis. e ectodermic cells (sixty-four, lighter and smaller), i entodermic cells (thirty-two, darker and larger), d central entodermic cell, filling the primitive gut-cavity, o peripheral entodermic cell, stopping up the opening of the primitive mouth (yelk-stopper in the Rusconian anus).)

(FIGURE 1.73. Gastrula of the rabbit. A as a solid, spherical cluster of cells, B changing into the embryonic vesicle, bp primitive mouth, ep ectoderm, hy entoderm.)

The numbers of careful studies of animal gastrulation that have been made in the last few decades have completely established the views I have expounded, and which I first advanced in the years 1872 to 1876. For a time they were greatly disputed by many embryologists. Some said that the original embryonic form of the metazoa was not the gastrula, but the "planula"--a double-walled vesicle with closed cavity and without mouth-aperture; the latter was supposed to pierce through gradually. It was afterwards shown that this planula (found in several sponges, etc.) was a later evolution from the gastrula. It was also shown that what is called delamination--the rise of the two primary germinal layers by the folding of the surface of the blastoderm (for instance, in the Geryonidae and other medusae)--was a secondary formation, due to cenogenetic variations from the original invagination of the blastula. The same may be said of what is called "immigration," in which certain cells or groups of cells are detached from the simple layer of the blastoderm, and travel into the interior of the blastula; they attach themselves to the inner wall of the blastula, and form a second internal epithelial layer--that is to say, the entoderm. In these and many other controversies of modern embryology the first requisite for clear and natural explanation is a careful and discriminative distinction between palingenetic (hereditary) and cenogenetic (adaptive) processes. If this is properly attended to, we find evidence everywhere of the biogenetic law.

CHAPTER X(10. THE COELOM THEORY.)

 

The two "primary germinal layers" which the gastraea theory has shown to be the first foundation in the construction of the body are found in this simplest form throughout life only in animals of the lowest grade--in the gastraeads, olynthus (the stem-form of the sponges), hydra, and similar very simple animals. In all the other animals new strata of cells are formed subsequently between these two primary body-layers, and these are generally comprehended under the title of the middle layer, or mesoderm. As a rule, the various products of this middle layer afterwards constitute the great bulk of the animal frame, while the original entoderm, or internal germinal layer, is restricted to the clothing of the alimentary canal and its glandular appendages; and, on the other hand, the ectoderm, or external germinal layer, furnishes the outer clothing of the body, the skin and nervous system.

In some large groups of the lower animals, such as the sponges, corals, and flat-worms, the middle germinal layer remains a single connected mass, and most of the body is developed from it; these have been called the three-layered metazoa, in opposition to the two-layered animals described. Like the two-layered animals, they have no body-cavity--that is to say, no cavity distinct from the alimentary system. On the other hand, all the higher animals have this real body-cavity (coeloma), and so are called coelomaria. In all these we can distinguish four secondary germinal layers, which develop from the two primary layers. To the same class belong all true vermalia (excepting the platodes), and also the higher typical animal stems that have been evolved from them--molluscs, echinoderms, articulates, tunicates, and vertebrates.

(FIGURES 1.74 AND 1.75. Diagram of the four secondary germinal layers, transverse section through the metazoic embryo: Figure 1.74 of an annelid, Figure 1.75 of a vermalian. a primitive gut, dd ventral glandular layer, df ventral fibre-layer, hm skin-fibre-layer, hs skin-sense-layer, u beginning of the rudimentary kidneys, n beginning of the nerve-plates.)

The body-cavity (coeloma) is therefore a new acquisition of the animal body, much younger than the alimentary system, and of great importance. I first pointed out this fundamental significance of the coelom in my Monograph on the Sponges (1872), in the section which draws a distinction between the body-cavity and the gut-cavity, and which follows immediately on the germ-layer theory and the ancestral tree of the animal kingdom (the first sketch of the gastraea theory). Up to that time these two principal cavities of the animal body had been confused, or very imperfectly distinguished; chiefly because Leuckart, the founder of the coelenterata group (1848), has attributed a body-cavity, but not a gut-cavity, to these lowest metazoa. In reality, the truth is just the other way about.

The ventral cavity, the original organ of nutrition in the multicellular animal-body, is the oldest and most important organ of all the metazoa, and, together with the primitive mouth, is formed in every case in the gastrula as the primitive gut; it is only at a much later stage that the body-cavity, which is entirely wanting in the coelenterata, is developed in some of the metazoa between the ventral and the body wall. The two cavities are entirely different in content and purport. The alimentary cavity (enteron) serves the purpose of digestion; it contains water and food taken from without, as well as the pulp (chymus) formed from this by digestion. On the other hand, the body-cavity, quite distinct from the gut and closed externally, has nothing to do with digestion; it encloses the gut itself and its glandular appendages, and also contains the sexual products and a certain amount of blood or lymph, a fluid that is transuded through the ventral wall.

As soon as the body-cavity appears, the ventral wall is found to be separated from the enclosing body-wall, but the two continue to be directly connected at various points. We can also then always distinguish a number of different layers of tissue in both walls--at least two in each. These tissue-layers are formed originally from four different simple cell-layers, which are the much-discussed four secondary germinal layers. The outermost of these, the skin-sense-layer (Figures 1.74 and 1.75 hs), and the innermost, the gut-gland-layer (dd), remain at first simple epithelia or covering-layers. The one covers the outer surface of the body, the other the inner surface of the ventral wall; hence they are called confining or limiting layers. Between them are the two middle-layers, or mesoblasts, which enclose the body-cavity.

(FIGURE 1.76. Coelomula of sagitta (gastrula with a couple of coelom-pouches. (From Kowalevsky.) bl.p primitive mouth, al primitive gut, pv coelom-folds, m permanent mouth.)

The four secondary germinal layers are so distributed in the structure of the body in all the coelomaria (or all metazoa that have a body-cavity) that the outer two, joined fast together, constitute the body-wall, and the inner two the ventral wall; the two walls are separated by the cavity of the coelom. Each of the walls is made up of a limiting layer and a middle layer. The two limiting layers chiefly give rise to epithelia, or covering-tissues, and glands and nerves, while the middle layers form the great bulk of the fibrous tissue, muscles, and connective matter. Hence the latter have also been called fibrous or muscular layers. The outer middle layer, which lies on the inner side of the skin-sense-layer, is the skin fibre-layer; the inner middle layer, which attaches from without to the ventral glandular layer, is the ventral fibre layer. The former is usually called briefly the parietal, and the latter the visceral layer or mesoderm. Of the many different names that have been given to the four secondary germinal layers, the following are those most in use to-day:--

Skin-sense-layer (outer limiting layer) and 2. Skin-fibre-layer (outer middle layer). Neural layer (neuroblast) and II. Parietal layer (myoblast). The two secondary germinal layers of the body-wall: 1. Epithelial. 2. Fibrous. Gut-fibre-layer (inner middle layer) and 4. Gut-gland-layer (inner limiting layer).

III. Visceral layer (gonoblast) and IV. Enteral layer (enteroblast). The two secondary germinal layers of the gut-wall: 3. Fibrous. 4. Epithelial.

The first scientist to recognise and clearly distinguish the four secondary germinal layers was Baer. It is true that he was not quite clear as to their origin and further significance, and made several mistakes in detail in explaining them. But, on the whole, their great importance did not escape him. However, in later years his view had to be given up in consequence of more accurate observations. Remak then propounded a three-layer theory, which was generally accepted. These theories of cleavage, however, began to give way thirty years ago, when Kowalevsky (1871) showed that in the case of Sagitta (a very clear and typical subject of gastrulation) the two middle germinal layers and the two limiting layers arise not by cleavage, but by folding--by a secondary invagination of the primary inner germ-layer. This invagination or folding proceeds from the primitive mouth, at the two sides of which (right and left) a couple of pouches are formed. As these coelom-pouches or coelom-sacs detach themselves from the primitive gut, a double body-cavity is formed (Figures 1.74 to 1.76).

(FIGURE 1.77. Coelomula of sagitta, in section. (From Hertwig.) D dorsal side, V ventral side, ik inner germinal layer, mv visceral mesoblast, lh body-cavity, mp parietal mesoblast, ak outer germinal layer.)

The same kind of coelom-formation as in sagitta was afterwards found by Kowalevsky in brachiopods and other invertebrates, and in the lowest vertebrate--the amphioxus. Further instances were discovered by two English embryologists, to whom we owe very considerable advance in ontogeny--E. Ray-Lankester and F. Balfour. On the strength of these and other studies, as well as most extensive research of their own, the brothers Oscar and Richard Hertwig constructed in 1881 the Coelom Theory. In order to appreciate fully the great merit of this illuminating and helpful theory, one must remember what a chaos of contradictory views was then represented by the "problem of the mesoderm," or