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

(Baer's "muscular layer") the "skin layer" (it would be better to say, skin-fibre layer); it forms the outer wall of the body (the true skin, the muscles, etc.). To the inner secondary leaf (Baer's "vascular layer") he gave the name of the "alimentary-fibre layer"; this forms the outer envelope of the alimentary canal, with the mesentery, the heart, the blood-vessels, etc.

On this firm foundation provided by Remak for histogeny, or the science of the formation of the tissues, our knowledge has been gradually built up and enlarged in detail. There have been several attempts to restrict and even destroy Remak's principles. The two anatomists, Reichert (of Berlin) and Wilhelm His (of Leipzic), especially, have endeavoured in their works to introduce a new conception of the embryonic development of the vertebrate, according to which the two primary germinal layers would not be the sole sources of formation. But these efforts were so seriously marred by ignorance of comparative anatomy, an imperfect acquaintance with ontogenesis, and a complete neglect of phylogenesis, that they could not have more than a passing success. We can only explain how these curious attacks of Reichert and His came to be regarded for a time as advances by the general lack of discrimination and of grasp of the true object of embryology.

Wilhelm His published, in 1868, his extensive Researches into the Earliest Form of the Vertebrate Body,* (* None of His's works have been translated into English.) one of the curiosities of embryological literature. The author imagines that he can build a "mechanical theory of embryonic development" by merely giving an exact description of the embryology of the chick, without any regard to comparative anatomy and phylogeny, and thus falls into an error that is almost without parallel in the history of biological literature. As the final result of his laborious investigations, His tells us "that a comparatively simple law of growth is the one essential thing in the first development. Every formation, whether it consist in cleavage of layers, or folding, or complete division, is a consequence of this fundamental law." Unfortunately, he does not explain what this "law of growth" is; just as other opponents of the theory of selection, who would put in its place a great "law of evolution," omit to tell us anything about the nature of this. Nevertheless, it is quite clear from His's works that he imagines constructive Nature to be a sort of skilful tailor. The ingenious operator succeeds in bringing into existence, by "evolution," all the various forms of living things by cutting up in different ways the germinal layers, bending and folding, tugging and splitting, and so on.

His's embryological theories excited a good deal of interest at the time of publication, and have evoked a fair amount of literature in the last few decades. He professed to explain the most complicated parts of organic construction (such as the development of the brain) in the simplest way on mechanical principles, and to derive them immediately from simple physical processes (such as unequal distribution of strain in an elastic plate). It is quite true that a mechanical or monistic explanation (or a reduction of natural processes) is the ideal of modern science, and this ideal would be realised if we could succeed in expressing these formative processes in mathematical formulae. His has, therefore, inserted plenty of numbers and measurements in his embryological works, and given them an air of "exact" scholarship by putting in a quantity of mathematical tables. Unfortunately, they are of no value, and do not help us in the least in forming an "exact" acquaintance with the embryonic phenomena. Indeed, they wander from the true path altogether by neglecting the phylogenetic method; this, he thinks, is "a mere by-path," and is "not necessary at all for the explanation of the facts of embryology," which are the direct consequence of physiological principles. What His takes to be a simple physical process--for instance, the folding of the germinal layers (in the formation of the medullary tube, alimentary tube, etc.)--is, as a matter of fact, the direct result of the growth of the various cells which form those organic structures; but these growth-motions have themselves been transmitted by heredity from parents and ancestors, and are only the hereditary repetition of countless phylogenetic changes which have taken place for thousands of years in the race-history of the said ancestors. Each of these historical changes was, of course, originally due to adaptation; it was, in other words, physiological, and reducible to mechanical causes. But we have, naturally, no means of observing them now. It is only by the hypotheses of the science of evolution that we can form an approximate idea of the organic links in this historic chain.

All the best recent research in animal embryology has led to the confirmation and development of Baer and Remak's theory of the germinal layers. One of the most important advances in this direction of late was the discovery that the two primary layers out of which is built the body of all vertebrates (including man) are also present in all the invertebrates, with the sole exception of the lowest group, the unicellular protozoa. Huxley had detected them in the medusa in 1849. He showed that the two layers of cells from which the body of this zoophyte is developed correspond, both morphologically and physiologically, to the two original germinal layers of the vertebrate. The outer layer, from which come the external skin and the muscles, was then called by Allman (1853) the "ectoderm" (outer layer, or skin); the inner layer, which forms the alimentary and reproductory organs, was called the "entoderm" (= inner layer). In 1867 and the following years the discovery of the germinal layers was extended to other groups of the invertebrates. In particular, the indefatigable Russian zoologist, Kowalevsky, found them in all the most diverse sections of the invertebrates--the worms, tunicates, echinoderms, molluscs, articulates, etc.

In my monograph on the sponges (1872) I proved that these two primary germinal layers are also found in that group, and that they may be traced from it right up to man, through all the various classes, in identical form. This "homology of the two primary germinal layers" extends through the whole of the metazoa, or tissue-forming animals; that is to say, through the whole animal kingdom, with the one exception of its lowest section, the unicellular beings, or protozoa. These lowly organised animals do not form germinal layers, and therefore do not succeed in forming true tissue. Their whole body consists of a single cell (as is the case with the amoebae and infusoria), or of a loose aggregation of only slightly differentiated cells, though it may not even reach the full structure of a single cell (as with the monera). But in all other animals the ovum first grows into two primary layers, the outer or animal layer (the ectoderm, epiblast, or ectoblast), and the inner or vegetal layer (the entoderm, hypoblast, or endoblast); and from these the tissues and organs are formed. The first and oldest organ of all these metazoa is the primitive gut (or progaster) and its opening, the primitive mouth (prostoma). The typical embryonic form of the metazoa, as it is presented for a time by this simple structure of the two-layered body, is called the gastrula; it is to be conceived as the hereditary reproduction of some primitive common ancestor of the metazoa, which we call the gastraea. This applies to the sponges and other zoophyta, and to the worms, the mollusca, echinoderma, articulata, and vertebrata. All these animals may be comprised under the general heading of "gut animals," or metazoa, in contradistinction to the gutless protozoa.

I have pointed out in my Study of the Gastraea Theory [not translated] (1873) the important consequences of this conception in the morphology and classification of the animal world. I also divided the realm of metazoa into two great groups, the lower and higher metazoa. In the first are comprised the coelenterata (also called zoophytes, or plant-animals). In the lower forms of this group the body consists throughout life merely of the primary germinal layers, with the cells sometimes more and sometimes less differentiated. But with the higher forms of the coelentarata (the corals, higher medusae, ctenophorae, and platodes) a middle layer, or mesoderm, often of considerable size, is developed between the other two layers; but blood and an internal cavity are still lacking.

To the second great group of the metazoa I gave the name of the coelomaria, or bilaterata (or the bilateral higher forms). They all have a cavity within the body (coeloma), and most of them have blood and blood-vessels. In this are comprised the six higher stems of the animal kingdom, the annulata and their descendants, the mollusca, echinoderma, articulata, tunicata, and vertebrata. In all these bilateral organisms the two-sided body is formed out of four secondary germinal layers, of which the inner two construct the wall of the alimentary canal, and the outer two the wall of the body. Between the two pairs of layers lies the cavity (coeloma).

Although I laid special stress on the great morphological importance of this cavity in my Study of the Gastraea Theory, and endeavoured to prove the significance of the four secondary germinal layers in the organisation of the coelomaria, I was unable to deal satisfactorily with the difficult question of the mode of their origin. This was done eight years afterwards by the brothers Oscar and Richard Hertwig in their careful and extensive comparative studies. In their masterly Coelum Theory: An Attempt to Explain the Middle Germinal Layer [not translated] (1881) they showed that in most of the metazoa, especially in all the vertebrates, the body-cavity arises in the same way, by the outgrowth of two sacs from the inner layer. These two coelom-pouches proceed from the rudimentary mouth of the gastrula, between the two primary layers. The inner plate of the two-layered coelom-pouch (the visceral layer) joins itself to the entoderm; the outer plate (parietal layer) unites with the ectoderm. Thus are formed the double-layered gut-wall within and the double-layered body-wall without; and between the two is formed the cavity of the coelom, by the blending of the right and left coelom-sacs. We shall see this more fully in

Chapter 1.

10.

The many new points of view and fresh ideas suggested by my gastraea theory and Hertwig's coelom theory led to the publication of a number of writings on the theory of germinal layers. Most of them set out to oppose it at first, but in the end the majority supported it. Of late years both theories are accepted in their essential features by nearly every competent man of science, and light and order have been introduced into this once dark and contradictory field of research. A further cause of congratulation for this solution of the great embryological controversy is that it brought with it a recognition of the need for phylogenetic study and explanation.

Interest and practice in embryological research have been remarkably stimulated during the past thirty years by this appreciation of phylogenetic methods. Hundreds of assiduous and able observers are now engaged in the development of comparative embryology and its establishment on a basis of evolution, whereas they numbered only a few dozen not many decades ago. It would take too long to enumerate even the most important of the countless valuable works which have enriched embryological literature since that time. References to them will be found in the latest manuals of embryology of Kolliker, Balfour, Hertwig, Kollman, Korschelt, and Heider.

Kolliker's Entwickelungsgeschichte des Menschen und der hoherer Thiere, the first edition of which appeared forty-two years ago, had the rare merit at that time of gathering into presentable form the scattered attainments of the science, and expounding them in some sort of unity on the basis of the cellular theory and the theory of germinal layers. Unfortunately, the distinguished Wurtzburg anatomist, to whom comparative anatomy, histology, and ontogeny owe so much, is opposed to the theory of descent generally and