Part 3

Amongst these Nauplii—all belonging to the great group Crustacea, which includes crabs and shrimps—is one which gives rise to an animal decidedly degenerate, but not precisely parasitic in its habits. This Nauplius is the young of the ship’s Barnacle, a curious stalked body, inclosed in a shell of many pieces (Fig. 9). The egg of the Barnacle gives rise to an actively swimming Nauplius, the history of which is very astonishing. After swimming about for a time the Barnacle’s Nauplius fixes its head against a piece of wood, and takes to a perfectly fixed, immobile state of life (Fig. 10). The upper figures represent the Nauplius stage of animals closely resembling the Barnacle: the lower figures show the transformation of the Nauplius into the young Barnacle. Its organs of touch and of sight atrophy, its legs lose their locomotor function, and are simply used for bringing floating particles to the orifice of the stomach; so that an eminent naturalist has compared one of these animals to a man standing on his head and kicking his food into his mouth.

Were it not for the recapitulative phases in the development of the Barnacle, we may doubt whether naturalists would ever have guessed that it was a degenerate Crustacean. It was in fact for a long time regarded as quite remote from them, and placed among the snails and oysters; its true nature was only admitted when the young form was discovered.

Other parasitic organisms, which exhibit extreme degeneration as compared with their free-living relatives, might be cited and figured in profusion, did our limits permit. Very noteworthy are the degenerate Spiders—the mites, leading to still more degenerate forms, the Linguatulæ.

We have two of these represented in Figs. 11 and 12. The one (Fig. 11), as compared with a spider is seen still to possess the eight walking legs, small, it is true, whilst the palps and daggers of the spider have dwindled to a beak projecting from the front of the globular unjointed body. In the other the eight legs have become mere stumps, and the body is elongated like that of a worm.

The instances of degeneration which we have so far examined are due to parasitism, except in the example of the Barnacle, where we have an instance of degeneration due to sessile and immobile habit of life. We may now proceed to look at some sessile or immobile animals which are not usually regarded as degenerate, but which, I think, there is every reason to believe are the degenerate descendants of very much higher and more elaborate ancestors. These are certain marine animals, the Ascidians, or sea-squirts. These animals are found encrusting rocks, stones, and weeds on the sea bottom. Sometimes they are solitary (Fig. 13), but many of them produce buds, like plants, and so form compound masses or sheets of individuals all connected and continuous with one another, like the buds on a creeping plant (Fig. 14).

We will examine one of the simple forms—a tough mass like a leather bottle with two openings; water is continually passing in at the one and out at the other of these apertures. If we remove the leathery outer-case (Fig. 15), we find that there is a soft creature within which has the following parts:—Leading from the mouth a great throat, followed by an intestine. The throat is perforated by innumerable slits, through which the water passes into a chamber—the cloaca: in passing, the water aërates the blood which circulates in the framework of the slits. The intestine takes a sharp bend, which causes it to open also into the cloaca. Between the orifice of the mouth and of the cloaca there is a nerve-ganglion.

My object in the next place is to show that the structure and life-history of these Ascidians may be best explained on the hypothesis that they are instances of degeneration; that they are the modified descendants of animals of higher, that is more elaborate structure, and in fact are degenerate Vertebrata, standing in the same relation to fishes, frogs, and men, as do the barnacles to shrimps, crabs, and lobsters.

The young of some, but by no means of all these Ascidians, have a form totally different from that of their parents. The egg of Phallusia gives rise to a tadpole, a drawing of which placed side by side with the somewhat larger tadpole of the common frog is seen in the adjoining figure (Fig. 16). The young Ascidian has the same general shape as the young frog, but not only this; the resemblance extends into details, the internal organs agreeing closely in the two cases. Further still as shown by the beautiful researches of the Russian naturalist, Kowalewsky, the resemblance reaches absolute identity when we examine the way in which the various organs arise from the primitive egg-cell. Tail, body, spiracle, eye, and mouth, agree in the two tadpoles, the only important difference being in the position of the two mouths and in the fact that the Ascidian has one eye while the frog has two.

Now let us look at the internal organs (Fig. 17). There are four structures, which are all four possessed at some time of their lives by all those animals which we call the Vertebrata, the great branch of the pedigree to which fishes, reptiles, birds, beasts, and men belong. And the combination of these marks or structural peculiarities is an overwhelming piece of evidence in favour of the supposition that the creatures which possess this combination are derived from one common ancestor. Just as one would conclude that a man whom one might meet, say on Salisbury Plain, must belong to the New Zealand race, if it were found not only that he had the colour, and the hair, and the shape of head of a New Zealander, but also that he was tattooed like a New Zealander, carried the weapons of a New Zealander, and, over and above in addition to these proofs, that he talked the Maori language and none other; so here, in the case of the vertebrate race, there are certain qualities and possessions, the accumulation of which cannot be conceived of as occurring in any animal but one belonging to that race. These four great structural features are—first, the primitive backbone or notochord; second, the throat perforated by gill-slits; third, the tubular nerve-centre or spinal cord and brain placed along the back; and lastly, and perhaps most distinctive and clinching as an evidence of affinity, the myelonic or cerebral eye.

Now let us convince ourselves that these four features exist not only in the frog’s tadpole, as they do in all fishes, reptiles, birds, and beasts, but that they also exist in the Ascidian tadpole, and, it may be added, co-exist in no other animals at all.

The corresponding parts are named in Figs. 16 and 17, in such a way as to render their agreement tolerably clear, whilst in Fig. 18 a more detailed representation of the head of an Ascidian tadpole is given.

It is clear then that the Ascidians must be admitted to be Vertebrates, and must be classified in that great sub-kingdom or branch of the animal pedigree. The Ascidian tadpole is very unlike its parent the Ascidian, and has to go through a process of degeneration in order to arrive at the adult structure. The diagrams which are reproduced in Figs. 19 and 20, show how this degeneration proceeds. It will be observed, that in somewhat the same manner as the young barnacle, the young Ascidian fixes itself to a stone by its head: then the tail with its notochord and nerve-chord atrophies. The body grows and gradually changes its shape, whilst the cloacal chamber forms. The brain remains quite small and undeveloped, and the remarkable myelonic eye (the eye in the brain) disappears. The number of gill-slits increases as the animal grows in size and its outer skin becomes tough and leather-like.

Before saying anything further on the subject of degeneration, it seems desirable once more to direct attention to the myelonic or cerebral eye which the Ascidian tadpole possesses in common with all Vertebrates. All other animals which have eyes develop the retina or sensitive part of the eye from their outer skin (See Figs. 21 and 22, and explanation.) It is easy to understand that an organ which is to be affected by the light should form on the surface of the body where the light falls. It has long been known as a very puzzling and unaccountable peculiarity of Vertebrates, that the retina or sensitive part of the eye grows out in the embryo as a bud or vesicle of the brain, and thus forms deeply below the surface and away from the light (see Fig. 23, and explanation). The Ascidian tadpole helps us to understand this, for it is perfectly transparent and has its eye actually inside its brain. The light passes through the transparent tissues and acts on the pigmented eye, lying deep in the brain. We are thus led to the conclusion—and I believe this inference to be now for the first time put into so many words—that the original Vertebrate must have been a transparent animal, and had an eye or pair of eyes inside its brain, like that of the Ascidian tadpole. As the tissues of this ancestral Vertebrate grew denser and more opaque, the eye-bearing part of the brain was forced by natural selection to grow outwards towards the surface, in order that it might still be in a position to receive the influence of the sun’s rays. Thus the very peculiar mode of development of the Vertebrate eye from two parts, a brain-vesicle (Fig. 23, A a, and B p r), and a skin-vesicle (Fig. 23, B e, l), is accounted for.

The cases of degeneration which I have up to this point brought forward, are cases which admit of very little dispute or doubt. They are attested by either the history of the individual development of the organisms in question, as in Sacculina, in the Barnacle, and in the Ascidian, or they are cases where the comparison of the degenerate animal, with others like it in structure, but not degenerate, renders the hypothesis of degeneration an unassailable one. Such cases are the Acarus or mite, and the skin-worm (Demodex).

We have seen that degeneration, or the simplification of the general structure of an animal, may be due to the ancestors of that animal having taken to one of two new habits of life, either the parasitic or the immobile. Other new habits of life appear also to be such as to lead to degeneration. Let us suppose a race of animals fitted and accustomed to catch their food, and having a variety of organs to help them in this chase—suppose such animals suddenly to acquire the power of feeding on the carbonic acid dissolved in the water around them just as green plants do. This would lead to a degeneration; they would cease to hunt their food, and would bask in the sunlight, taking food in by the whole surface, as plants do by their leaves. Certain small flat worms, by name Convoluta, of a bright green colour, appear to be in this condition. Their green colour is known to be the same substance as leaf-green; and Mr. Patrick Geddes has recently shown that by the aid of this green substance they feed on carbonic acid, making starch from it as plants do. As a consequence we find that their stomachs and intestines as well as their locomotive organs become simplified, since they are but little wanted. These vegetating animals, as Mr. Geddes calls them, are the exact complement of the carnivorous plants, and show how a degeneration of animal forms may be caused by vegetative nutrition.

Another possible cause of degeneration appears to be the indirect one of minute size. It cannot be doubted that natural selection has frequently acted on a race of animals so as to reduce the size of the individuals. The smallness of size has been favourable to their survival in the struggle for existence, and in some cases they have been reduced to even microscopic proportions. But this reduction of size has, when carried to an extreme, resulted in the loss or suppression of some of the most important organs of the body. The needs of a very minute creature are limited as compared with those of a large one, and thus we may find heart and blood-vessels, gills and kidneys, besides legs and muscles, lost by the diminutive degenerate descendants of a larger race. That this is a possible course of change all will, I think, admit. It is actually exemplified in Appendicularia—the only adult representative of the Ascidian tadpole—still tadpole-like in form and structure, but curiously degenerate and simplified in its internal organs. This kind of degeneration is also exemplified in the Rotifers, or wheel animalcules, in the minute Crustacean water-fleas (Ostracoda), and in the Moss-polyps, or Polyzoa. Roughly then we may sum up the immediate antecedents of degenerative evolution as, 1, Parasitism; 2, Fixity or immobility; 3, Vegetative nutrition; 4, Excessive reduction of size. This is not a logical enumeration, for each of these causes involves, or may be inseparably connected with, one or more of the others. It will serve for the present as well as a more exhaustive analysis. (See Note C.)

And now we have to note an important fact with regard to the evidence which we can obtain of the occurrence of this process of degeneration. We have seen that the most conclusive evidence is that of the recapitulative development of the individual. The Ascidian Phallusia shows itself to be a degenerate Vertebrate by beginning life as a tadpole. But such recapitulative development is by no means the rule. Quite arbitrarily, we find, it is exhibited in one animal and not in a nearly allied kind. Thus very many animals belonging to the Ascidian group have no tadpole young—just as some tree-frogs have no tadpoles. It is quite possible, and often, more often than not, occurs, that the most important part of the recapitulative phases are absent from the developmental history of an animal. The egg proceeds very rapidly to produce the adult form, and all the wonderful series of changes showing the animal’s ancestry are absolutely and completely omitted; that is to say, all those stages which are of importance for our present purpose. Just as certain bodies pass from the solid to the liquid state at a bound, omitting all intermediate phases of consistence, but giving evidence of “internal work” by the suggestive phenomenon of latent heat—so do these embryos skip long tracts in the historically continuous phases of form, and present to us only the intangible correlative “internal work” in place of the tangible series of embryonic changes of shape.

Now I want to put this case—a supposition—before the reader who has so far followed me in these pages. Suppose, as might well have happened, that the Barnacles, one and all, instead of recapitulating in their early life, were to develop directly from the egg to the adult form, as so many animals do; should we have ever made out that they were degenerate Crustaceans? Possibly we should: their adult structure still bears important marks of affinities with crabs and shrimps; but as a matter of fact before their recapitulative development had been discovered they were classed by the great Cuvier and other naturalists with the Molluscs, the mussels and snails.

Suppose again that all the existing Ascidians, as many of them actually have, had long ago lost their recapitulative history in growth from the egg: suppose that no such a thing as an Ascidian tadpole existed, but that the Ascidian’s egg grew as directly as possible into an Ascidian, in every living species of the group. This might easily be the case. Then most assuredly we should not have the least notion that the Ascidians were degenerate Vertebrates. We should still class them where they used to be classed before the Russian naturalist Kowalewsky discovered the true history and structure of the Ascidian tadpole. I believe that I shall have the assent of every naturalist when I say that the vertebrate character of the Ascidians and the history of their degeneration would never have been suspected, or even dreamed of, had the Ascidian tadpoles ceased to appear in the course of the Ascidian development at a geological period anterior to the present epoch.

This being the case, it must be admitted that it is quite possible—I do not say more than possible—that other groups of animals besides parasites, Barnacles, and Ascidians, are degenerate. It is quite possible that animals with considerable complexity of structure, at least as complex as the Ascidians, may have been produced by degeneration from still more highly-organized ancestors. Any group of animals to which we can turn may possibly be the result of degeneration, and yet offer no evidence of that degeneration in its growth from the egg.

Accordingly, wherever we can note that a group of organisms is characterized by habits likely to lead to degeneration, such as I have enumerated, viz., parasitism or immobility, or certain special modes of nutrition, or again, by minute size of its representatives—there we are justified in applying the hypothesis of degeneration, even in the absence of any confirmatory evidence from embryology. When we so apply this hypothesis we find in not a few cases, in working over the details of the organization of many different animals by the light which it affords—that much becomes clear and assignable to cause which, on the hypothesis either of “balance” or of “elaboration,” is quite hopelessly obscure. As examples of groups of animals which can thus be satisfactorily explained I may cite first of all the Sponges: as only somewhat less degenerate, we have all the Polyps and Coral-animals, also the Starfishes. Amongst the Mollusca—the group of headless bivalves, the oysters, mussels and clams, known as the Lamellibranchs, are, when one once looks at their structure in this light, clearly enough explained as degenerated from a higher type of head-bearing active creatures like the Cuttle-fish; whilst the Polyzoa or Moss-polyps stand in precisely the same kind of relation to the higher Mollusca as do the Ascidians to the higher Vertebrates: they have greatly degenerated, and become minute encrusting organisms which, like some of the Ascidians, build up colonies by plant-like budding growth. The Rotifers, or wheel animalcules, I have already mentioned as best explained by the supposition that they are the descendants of far larger and more fully-organized animals provided with locomotive appendages or limbs: they have dwindled and degenerated to their present minute size and curiously suggestive structure.

Besides these there are other very numerous cases of animal structure which can best be explained by the hypothesis of degeneration. A discussion of these, and a due exposition of the application of the hypothesis of degeneration to the various groups just cited, would involve a complete treatise on comparative anatomy and embryology, and lead far beyond the limitations of this little volume.

All that has been, thus far, here said on the subject of Degeneration is so much zoological specialism, and may appear but a narrow restriction of the discussion to those who are not zoologists. Though we may establish the hypothesis most satisfactorily by the study of animal organization and development, it is abundantly clear that degenerative evolution is by no means limited in its application to the field of zoology. It clearly offers an explanation of many vegetable phenomena, and is already admitted by botanists as the explanation of the curious facts connected with the reproductive process in the higher plants. As a further example of its application in this field, the yeast-plant may be adduced, which is in all probability a degenerate floating form derived from a species of mould (Mucor). In other fields, wherever in fact the great principle of evolution has been recognised, degeneration plays an important part. In tracing the development of languages, philologists have long made use of the hypothesis of degeneration. Under certain conditions, in the mouths and minds of this or that branch of a race, a highly elaborate language has sometimes degenerated and become no longer fit to express complex or subtle conceptions, but only such as are simpler and more obvious. (See Note D.)

The traditional history of mankind furnishes us with notable examples of degeneration. High states of civilisation have decayed and given place to low and degenerate states. At one time it was a favourite doctrine that the savage races of mankind were degenerate descendants of the higher and civilised races. This general and sweeping application of the doctrine of degeneration has been proved to be erroneous by careful study of the habits, arts, and beliefs of savages; at the same time there is no doubt that many savage races as we at present see them are actually degenerate and are descended from ancestors possessed of a relatively elaborate civilisation. As such we may cite some of the Indians of Central America, the modern Egyptians, and even the heirs of the great oriental monarchies of præ-Christian times. Whilst the hypothesis of universal degeneration as an explanation of savage races has been justly discarded, it yet appears that degeneration has a very large share in the explanation of the condition of the most barbarous races, such as the Fuegians, the Bushmen, and even the Australians. They exhibit evidence of being descended from ancestors more cultivated than themselves.

With regard to ourselves, the white races of Europe, the possibility of degeneration seems to be worth some consideration. In accordance with a tacit assumption of universal progress—an unreasoning optimism—we are accustomed to regard ourselves as necessarily progressing, as necessarily having arrived at a higher and more elaborated condition than that which our ancestors reached, and as destined to progress still further. On the other hand, it is well to remember that we are subject to the general laws of evolution, and are as likely to degenerate as to progress. As compared with the immediate forefathers of our civilisation—the ancient Greeks—we do not appear to have improved so far as our bodily structure is concerned, nor assuredly so far as some of our mental capacities are concerned. Our powers of perceiving and expressing beauty of form have certainly not increased since the days of the Parthenon and Aphrodite of Melos. In matters of the reason, in the development of intellect, we may seriously inquire how the case stands. Does the reason of the average man of civilised Europe stand out clearly as an evidence of progress when compared with that of the men of bygone ages? Are all the inventions and figments of human superstition and folly, the self-inflicted torturing of mind, the reiterated substitution of wrong for right, and of falsehood for truth, which disfigure our modern civilisation—are these evidences of progress? In such respects we have at least reason to fear that we may be degenerate. Possibly we are all drifting, tending to the condition of intellectual Barnacles or Ascidians. It is possible for us—just as the Ascidian throws away its tail and its eye and sinks into a quiescent state of inferiority—to reject the good gift of reason with which every child is born, and to degenerate into a contented life of material enjoyment accompanied by ignorance and superstition. The unprejudiced, all-questioning spirit of childhood may not inaptly be compared to the tadpole tail and eye of the young Ascidian: we have to fear lest the prejudices, pre-occupations, and dogmatism of modern civilisation should in any way lead to the atrophy and loss of the valuable mental qualities inherited by our young forms from primæval man.

There is only one means of estimating our position, only one means of so shaping our conduct that we may with certainty avoid degeneration and keep an onward course. We are as a race more fortunate than our ruined cousins—the degenerate Ascidians. For us it is possible to ascertain what will conduce to our higher development, what will favour our degeneration. To us has been given the power to know the causes of things, and by the use of this power it is possible for us to control our destinies. It is for us by ceaseless and ever hopeful labour to try to gain a knowledge of man’s place in the order of nature. When we have gained this fully and minutely, we shall be able by the light of the past to guide ourselves in the future. In proportion as the whole of the past evolution of civilised man, of which we at present perceive the outlines, is assigned to its causes, we and our successors on the globe may expect to be able duly to estimate that which makes for, and that which makes against, the progress of the race. The full and earnest cultivation of Science—the Knowledge of Causes—is that to which we have to look for the protection of our race—even of this English branch of it—from relapse and degeneration.

FOOTNOTES

These pages formed a discourse delivered before the British Association at Sheffield on the evening of August 22nd, 1879, under the presidency of Professor Allman, LL.D., F.R.S.