Walter Holbrook Gaskell

“ At first sight it seems too great an absurdity even to imagine the possibility of any genetic connection between a fish and an arthropod, for to the mind's eye there arises immediately the picture of a salmon or a shark and a lobster or a spider. ”

“ Not having been educated in a morphological laboratory and taught that the one organ which is homologous throughout the animal kingdom is the gut, and that therefore the gut of the invertebrate ancestor must continue on as the gut of the vertebrate, the conception that the central nervous system has grown round and enclosed the original ancestral gut, and that the vertebrate has formed a new gut did not seem to me so impossible as to prevent my taking it as a working hypothesis, and seeing to what it would lead. ”

“ To those who believe in Evolution and the Darwinian theory of the survival of the fittest, it is simply inconceivable that a soft-bodied animal living in the mud, blind, with a rudimentary brain and rudimentary olfactory organs, such as is postulated when we think of Balanoglossus and Amphioxus, should hold its own and come victorious out of the struggle for existence at a time when the sea was peopled with powerful predaceous scorpion- and crab-like armour-plated animals possessing a well-developed brain, good eyes and olfactory organs, and powerful means of locomotion. ”

“ In the whole scheme of evolution we can recognize, not only an upward progress in the organization of the animal as a whole, but also a distinct advance in the structure of the tissues composing an individual, which accompanies that upward progress. Thus it is possible to speak of an evolution of the supporting tissues from the simplest form of connective tissue up to cartilage and thence to bone ”

“ In order, therefore, to compare two adult animals for the purpose of finding kinship between them, it is necessary to find whether parts such as the gut, which in both cases have the same function, arise from the same germinal layer in the embryo. We can, in fact, have no certainty of kinship, even although the two animals are built up as far as the adult state is concerned on a remarkably similar plan, unless we can study their respective embryos and find out what parts arise from the hypoblast and what from the epiblast. ”

“ The great value of the study of embryology for questions of the sequence of the evolution of animals is to be found in what is known as the Law of Recapitulation, which asserts that every animal gives some indication in the stages of its individual development of its ancestral history. Naturally enough it cannot pass through all the stages of its past history with equal clearness, for what has taken millions of years to be evolved has to be compressed into an evolution lasting only a few months or weeks, or even less. ”

“ If we take Limulus as the only living type of the Palæostraca, we are struck with the fact that the animal consists to all intents and purposes of prosomatic and mesosomatic regions only; the metasoma consisting of the segments posterior to the mesosoma is very insignificant, so that the large mass of the animal consists of what has become the head-region in the vertebrate; the spinal region, which has become in the higher vertebrates by far the largest region of the body, can hardly be said to exist in such an animal as Limulus. ”

“ Thus the suggestion arises that in the embryo a cellular lining is formed to a cœlomic cavity (mesothelium) composed of cells which have no communication with the nervous system, and are capable of a separate existence as free individuals, either in the form of germinal cells or of lymphocytes, hæmocytes, and plasma-cells, so that these latter free cells may be considered as living an independent existence in the body, and ministering to it in the same sense as the germ-cells live an independent existence in the body. ”

“ I have been told that it is impossible for one man to consider so vast a subject with that thoroughness which is necessary, before any theory can be accepted as the true solution of the problem. I acknowledge the vastness of the task, and feel keenly enough my own shortcomings. ”

“ For all questions of evolution the presence of vestigial organs in the embryo is the important consideration, for they give an indication of near ancestry; the early formation of the embryo concerns a much more remote ancestral period, all vestigial organs of which may well have been lost and obscured by cœnogenetic changes. Let us, then, consider the two things—the vestigial organs and the early formation of the embryo—separately, and see how far my opponents are justified in their statement that my theory contravenes the principles of embryology. ”

“ We must endeavour to dismiss from our imagination such forms as the salmon and shark as representatives of the fish-tribe, and the lobster and spider of the arthropods, and try to picture the kind of animals living in the seas in the early Devonian and Upper Silurian times, and then we find, to our surprise, that instead of the contrast between fishes and arthropods being so striking as to make any comparison between the two seem an absurdity, the difficulty in the last century, and even now, is to decide in many cases whether a fossil is an arthropod or a fish. ”

“ One of the principles of evolution is that any change which is supposed to have taken place in the process of formation of one animal or group of animals from a lower group must be in harmony with changes which are known to have occurred in that lower group. On the assumption, therefore, that the vertebrate branchiæ represent the branchial portion of the arthropod mesosomatic appendages which have sunk in and so become internal, we ought to find that in members of this very group such inclusion of branchial appendages has taken place. ”

“ I have not been able to obtain any evidence of a fusiform nerve-cell on the course of the terminal nerve-fibres as depicted by him; fusiform cells there are in plenty, as depicted in my drawing, but none with a large nucleus resembling those of the main ganglion. In no case, either in the flabellum or in the branchial organs of Limulus, or in the pecten-organs, have I ever seen a ganglion-cell within the chitin-layer; all the nuclei seen there resemble those of the cells of the hypodermis or else the elongated nuclei characteristic of the presence of nerve-fibres. ”

“ The growth of the individual, whether arthropod or vertebrate, spreads from the position of the central nervous system, regardless of whether that position is a ventral or dorsal one with respect to the yolk-mass. Where the pabulum is, there is the definite gut, the lining walls of which are called in the embryo, hypoblast; but when the pabulum is no longer there, although a tube is formed in the same manner as the alimentary canal of the arthropod, it is now called an epiblastic tube, and is known as the neural tube of the vertebrate. ”

“ What are the lines of investigation most likely to meet with success? Is it possible to lay down any laws of evolution? It is instructive to consider the nature of the investigations which have led to the two theories just mentioned, for the fundamental starting-point is remarkably different in the two cases. The one theory is based upon the study of the vertebrate itself, and especially of its central nervous system, and its supporters and upholders have been and are essentially anatomists, whose chief study is that of vertebrate and human anatomy. ”

“ It does not seem to me possible to conceive of the gonads arising from cells of the epiblast or of the hypoblast, in the sense that such cells are differentiated cells belonging to a layer with a definite meaning. When we consider that the gonad gives origin to the whole of a new individual, that in the protozoan ancestors of the Metazoa their ultimate aim and object was the formation of gonads, it seems a wrong conception to speak of the gonads as formed from cells belonging either to the gut-wall or to the external epithelium. The gonads must stand in a category by themselves ”

“ In any organ the nerves which are specially of value in determining its innervation, are of necessity the efferent or motor nerves, for the limits of their distribution in the organ are much more easily determined than those of the afferent or sensory nerves. The question therefore of primary importance in endeavouring to determine the nature of the origin of the alimentary canal from its innervation is the determination of the efferent supply to the musculature of its walls. ”

“ If we regard only the retinal and nervous elements, and for the moment pay no attention to the existence of the tube, Götte's observation that the true retina has been formed from the optic plate (Sinnes-platte) to which the retinal portion of the brain (retinal ganglion) has become firmly fixed, and that then the optic nerve has been formed by the withdrawal of the rest of the brain (optic lobes) , is word for word applicable to the description of the development of the compound retina of the arthropod eye, as has been already stated. ”

“ If it is a right conception that the excretory organs of the protostracan group, which gave origin to the vertebrates as well as to the crustaceans and arachnids, were of the nature of coxal glands, then it follows that such coxal glands must have existed originally on every segment, because they themselves were derived from the segmental organs of the annelids; it is therefore worth while making an attempt to trace the fate of such segmental organs in the vertebrate as well as in the crustacean and arachnid. ”

“ By the vertebrate skeleton I mean the bony and cartilaginous structures which form the backbone, the cranio-facial skeleton, the pectoral and pelvic girdles, and the bones of the limbs. I do not include the notochord in these skeletal tissues, because there is not the slightest evidence that the notochord played any part in the formation of these structures; the notochordal tissue is something sui generis, and never gives rise to cartilage or bone. The notochord happens to lie in the middle line of the body and is very conspicuous in the lowest vertebrate ”

“ But there is one most important fact which ought to make us hesitate before accepting the homology of the dorsal heart of the arthropod with the ventral heart of the vertebrate—The heart in all invertebrates is a systemic heart, i.e. drives the arterial blood to the different organs of the body, and then the veins carry it back to the respiratory organ, from whence it passes to the heart. ”

“ If it is permissible to speak of a dorsal and ventral surface to an egg, and we may imagine the egg held with such dorsal surface uppermost, then the yolk would be situated ventrally to the embryo, as in the vertebrate, if the protoplasmic cells of the embryo rose from their central position to the surface through the yolk, while if they sank through the yolk, the yolk would be situated dorsally to the embryo, as in the arthropod. ”

“ The neurones of the third order, the cells of which form the ganglionic layer and their neuraxons constitute the fibres of the optic nerve which end in the optic lobes of the brain.
Strictly speaking, of course, the visual cells with their elongated processes have no right to be called neurones: I only use Parker's phraseology in order to show how closely the two retinas agree even to the formation of synapses between the fine drawn-out processes of the visual cells and the neurones of the ganglion of the retina. ”

“ In fact, we find in the nearest living representative of the trilobites a retina and retinal ganglion and optic nerve, closely resembling that of the vertebrate, in close connection with an epithelial tube which has nothing to do with the organ of sight, but is one of a pair of anterior gut-diverticula. It is impossible to obtain more decisive evidence that the trilobites possessed a pair of gut-diverticula surrounded to a greater or less extent by the retina and optic nerve of each lateral eye. ”

“ When in the highest vertebrate a vestigial organ, such as the pineal gland, can be traced back without leaving the vertebrate kingdom to a distinct median eye, such as is found in the lamprey, that rudimentary organ is evidence of an organ which was functional in the earliest vertebrates or their immediate ancestors. So it is generally with well defined vestigial organs found in the adult animal; they always indicate an organ which was functional in the near ancestor. ”

“ Does the history of evolution pick out any particular organ or group of organs as more necessary than another for upward progress? If so, it is upon that organ or group of organs that special stress must be laid.
Since Darwin wrote the "Origin of Species," and laid down that the law of the 'survival of the fittest' is the factor upon which evolution depends, it has gradually dawned upon the scientific mind that 'the fittest' may be produced in two diametrically opposite ways: {17}either by progress upwards to a superior form, or by degeneration to a lower type of animal. ”

“ We see, in fact, that in the compound crustacean eye an extra layer of hypodermal cells has become inserted between the cornea and the retina to form a lens. So also in the vertebrate eye the lens is formed by an extra layer of the epidermal cells between the cornea and the retina. The fact that the vertebrate eye possesses a single lens, though its retina is composed of a number of ommatidia, while the crustacean eye possesses a lens to each ommatidium, may well be a consequence of the inversion of the vertebrate retina. ”