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Darwin, and After Darwin (Vol. 1 and 3, of 3)
by George John Romanes
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Once more, we have seen that aggregates of characters presenting resemblances to one another have always been found to be of special importance as guides to classification. This, of course, is what we should have expected, if the real meaning of classification be that of tracing lines of pedigree; but on the theory of special creation no reason can be assigned why single characters are not such sure tokens of a natural arrangement as are aggregates of characters, however trivial the latter may be. For it is obvious that unity of ideal might have been even better displayed by everywhere maintaining the pattern of some one important structure, than by doing so in the case of several unimportant structures. Take an analogous instance from human contrivances. Unity of ideal in the case of gun-making would be shown by the same principles of mechanism running through all the different sizes and shapes of gun-locks, rather than by the ornamental patterns engraved upon the outside. Yet it must be supposed that in the mechanisms assumed to have been constructed by special creation, it was the trivial details rather than the fundamental principles of these mechanisms which were chosen by the Divinity to display his ideals.

And this leads us to the next consideration—namely, that when in two different lines of descent animals happen to adopt similar habits of life, the modifications which they undergo in order to fit them for these habits often induces striking resemblances of structure between the two animals, as in the case of whales and fish. But in all such instances it is invariably found that the resemblance is only superficial and apparent: not anatomical or real. In other words, the resemblance does not extend further than it is necessary that it should, if both sets of organs are to be adapted to perform the same functions. Now this, again, is just what one would expect to find as the universal rule on the theory of descent, with modification of ancestral characters. But, on the opposite theory of special creation, I know not how it is to be explained that among so many instances of close superficial resemblance between creatures belonging to different branches of the tree of life, there are no instances of any real or anatomical resemblance. So far as their structures are adapted to perform a common function, there is in all such cases what may be termed a deceptive appearance of some unity of ideal; but, when carefully examined, it is always found that two apparently identical structures occurring on different branches of the classificatory tree are in fact fundamentally different in respect of their structural plan.

Lastly, we have seen that one of the guiding principles of classification has been empirically found to consist in setting a high value on "chains of affinities." That is to say, naturalists not unfrequently meet with a long series of progressive modifications of type, which, although it cannot be said that the continuity is anywhere broken, at last leads to so much divergence of character that, but for the intermediate links, the members at each end of the chain could not be suspected of being in any way related. Well, such cases of chains of affinity obviously tell most strongly in favour of descent with continuous modification; while it is impossible to suggest why, if all the links were separately forged by as many acts of special creation, there should have been this gradual transmutation of characters carried to the point where the original creative ideal has been so completely transformed that, but for the accident of the chain being still complete, no one of nature's interpreters could possibly have discovered the connexion. For, as we have seen, this is not a case in which any appeal can be logically made to the argument from ignorance of divine method, unless some independent evidence could be adduced in favour of special creation. And that no such independent evidence exists, it will be the object of future chapters to show.



CHAPTER III.

MORPHOLOGY.

The theory of evolution supposes that hereditary characters admit of being slowly modified wherever their modification will render an organism better suited to a change in its conditions of life. Let us, then, observe the evidence which we have of such adaptive modifications of structure, in cases where the need of such modification is apparent. We may begin by again taking the case of the whales and porpoises. The theory of evolution infers, from the whole structure of these animals, that their progenitors must have been terrestrial quadrupeds of some kind, which gradually became more and more aquatic in their habits. Now the change in the conditions of their life thus brought about would have rendered desirable great modifications of structure. These changes would have begun by affecting the least typical—that is, the least strongly inherited—structures, such as the skin, claws, and teeth. But, as time went on, the adaptation would have extended to more typical structures, until the shape of the body would have become affected by the bones and muscles required for terrestrial locomotion becoming better adapted for aquatic locomotion, and the whole outline of the animal more fish-like in shape. This is the stage which we actually observe in the seals, where the hind legs, although retaining all their typical bones, have become shortened up almost to rudiments, and directed backwards, so as to be of no use for walking, while serving to complete the fish-like taper of the body. (Fig. 2.) But in the whales the modification has gone further than this so that the hind legs have ceased to be apparent externally, and are only represented internally—and even this only in some species—by remnants so rudimentary that it is difficult to make out with certainty the homologies of the bones; moreover, the head and the whole body have become completely fish-like in shape. (Fig. 3.) But profound as are these alterations, they affect only those parts of the organism which it was for the benefit of the organism to have altered, so that it might be adapted to an aquatic mode of existence. Thus the arm, which is used as a fin, still retains the bones of the shoulder, fore-arm, wrist, and fingers, although they are all enclosed in a fin-shaped sack, so as to render them useless for any purpose other than swimming (Fig. 4.) Similarly, the head, although it so closely resembles the head of a fish in shape, still retains the bones of the mammalian skull in their proper anatomical relations to one another; but modified in form so as to offer the least possible resistance to the water. In short, it may be said that all the modifications have been effected with the least possible divergence from the typical mammalian type, which is compatible with securing so perfect an adaptation to a purely aquatic mode of life.







Now I have chosen the case of the whale and porpoise group, because they offer so extreme an example of profound modification of structure in adaptation to changed conditions of life. But the same thing may be seen in hundreds and hundreds of other cases. For instance, to confine our attention to the arm, not only is the limb modified in the whale for swimming, but in another mammal—the bat—it is modified for flying, by having the fingers enormously elongated and overspread with a membranous web.

In birds, again, the arm is modified for flight in a wholly different way—the fingers here being very short and all run together, while the chief expanse of the wing is composed of the shoulder and fore-arm. In frogs and lizards, again, we find hands more like our own; but in an extinct species of flying reptile the modification was extreme, the wing having been formed by a prodigious elongation of the fifth finger, and a membrane spread over it and the rest of the hand. (Fig. 5.) Lastly, in serpents the hand and arm have disappeared altogether.



Thus, even if we confine our attention to a single organ, how wonderful are the modifications which it is seen to undergo, although never losing its typical character. Everywhere we find the distinction between homology and analogy which was explained in the last chapter—the distinction, that is, between correspondence of structure and correspondence of function. On the one hand, we meet with structures which are perfectly homologous and yet in no way analogous: the structural elements remain, but are profoundly modified so as to perform wholly different functions. On the other hand, we meet with structures which are perfectly analogous, and yet in no way homologous: totally different structures are modified to perform the same functions. How, then, are we to explain these things? By design manifested in special creation, or by descent with adaptive modification? If it is said by design manifested in special creation, we must suppose that the Deity formed an archetypal plan of certain structures, and that he determined to adhere to this plan through all the modifications which those structures exhibit. But, if so, why is it that some structures are selected as typical and not others? Why should the vertebral skeleton, for instance, be tortured into every conceivable variety of modification in order to subserve as great a variety of functions; while another structure, such as the eye, is made in different sub-kingdoms on fundamentally different plans, notwithstanding that it has throughout to perform the same function? Will any one have the hardihood to assert that in the case of the skeleton the Deity has endeavoured to show his ingenuity, by the manifold functions to which he has made the same structure subservient; while in the case of the eye he has endeavoured to show his resources, by the manifold structures which he has adapted to serve the same function? If so, it becomes a most unfortunate circumstance that, throughout both the vegetable and animal kingdoms, all cases which can be pointed to as showing ingenious adaptation of the same typical structure to the performance of widely different functions—or cases of homology without analogy,—are cases which come within the limits of the same natural group of plants and animals, and therefore admit of being equally well explained by descent from a common ancestry; while all cases of widely different structures performing the same function—or cases of analogy without homology,—are to be found in different groups of plants or animals, and are therefore suggestive of independent variations arising in the different lines of hereditary descent.

To take a specific illustration. The octopus, or devil-fish, belongs to a widely different class of animals from a true fish; and yet its eye, in general appearance, looks wonderfully like the eye of a true fish. Now, Mr. Mivart pointed to this fact as a great difficulty in the way of the theory of evolution by natural selection, because it must clearly be a most improbable thing that so complicated a structure as the eye of a fish should happen to be arrived at through each of two totally different lines of descent. And this difficulty would, indeed, be a formidable one to the theory of evolution, if the similarity were not only analogical but homological. Unfortunately for the objection, however, Darwin clearly showed in his reply that in no one anatomical or homologous feature do the two structures resemble one another; so that, in point of fact, the two organs do not resemble one another in any particular further than it is necessary that they should, if both are to be analogous, or to serve the same function as organs of sight. But now, suppose that this had not been the case, and that the two structures, besides presenting the necessary superficial or analogical resemblance, had also presented an anatomical or homologous resemblance, with what force might it have then been urged,—Your hypothesis of hereditary descent with progressive modification being here excluded by the fact that the animals compared belong to two widely different branches of the tree of life, how are we to explain the identity of type manifested by these two complicated organs of vision? The only hypothesis open to us is intelligent adherence to an ideal plan or mechanism. But as this cannot now be urged in any comparable case throughout the whole organic world, we may on the other hand present it as a most significant fact, that while within the limits of the same large branch of the tree of life we constantly find the same typical structures modified so as to perform very different functions, we never find any of these particular types of structure in other large branches of the tree. That is to say, we never find typical structures appearing except in cases where their presence may be explained by the hypothesis of hereditary descent; while in thousands of such cases we find these structures undergoing every conceivable variety of adaptive modification.

Consequently, special creationists must fall back upon another position and say,—Well, but it may have pleased the Deity to form a certain number of ideal types, and never to have allowed the structures occurring in one type to appear in any of the others. We answer,—Undoubtedly such may have been the case; but, if so, it is a most unfortunate thing for your theory, because the fact implies that the Deity has planned his types in such a way as to suggest the counter-theory of descent. For instance, it would seem most capricious on the part of the Deity to have made the eyes of an innumerable number of fish on exactly the same ideal type, and then to have made the eye of the octopus so exactly like these other eyes in superficial appearance as to deceive so accomplished a naturalist as Mr. Mivart, and yet to have taken scrupulous care that in no one ideal particular should the one type resemble the other. However, adopting for the sake of argument this great assumption, let us suppose that God did lay down these arbitrary rules for his own guidance in creation, and then let us see to what the assumption leads. If the Deity formed a certain number of ideal types, and determined that on no account should he allow any part of one type to appear in any part of another, surely we should expect that within the limits of the same type the same typical structures should always be present. Thus, remember what efforts, so to speak, have been made to maintain the uniformity of type in the case of the fore-limb as previously explained, and should we not expect that in other and similar cases a similar method should have been followed? Yet we repeatedly find that this is not the case. Even in the whale, as we have seen, the hind-limbs are either altogether absent or dwindled almost to nothing; and it is impossible to see in what respect the hind-limbs are of any less ideal value than the fore-limbs—which are carefully preserved in all vertebrated animals except the snakes, and the extinct Dinornis, where again we meet in this particular with a sudden and sublime indifference to the maintenance of a typical structure. (Fig. 6.)[4] Now I say that if the theory of ideal types is true, we have in these facts evidence of a most unreasonable inconsistency. But the theory of descent with continued adaptive modification fully explains all the known cases; for in every case the degree of divergence from the typical structure which an organism presents corresponds, in a general way, with the length of time during which the divergence has been going on. Thus we scarcely ever meet with any great departure from the typical form with respect to one of the organs, without some of the other organs being so far modified as of themselves to indicate, on the supposition of descent with modification, that the animal or plant must have been subject to the modifying influences for an enormously long series of generations. And this combined testimony of a number of organs in the same organism is what the theory of descent would lead us to expect, while the rival theory of design can offer no explanation of the fact, that when one organ shows a conspicuous departure from the supposed ideal type, some of the other organs in the same organism should tend to keep it company by doing likewise.

[4] It is, however, probable that all species of the genus retained a tiny rudiment of wings in greatly dwindled scapulo-coracoid bones. And Mr. H. O. Forbes has detected, in a recently exhumed specimen of the latter, an indication of the glenoid cavity, for the articulation of an extremely aborted humerus. (See Nature, Jan. 14th, 1892.)



As an illustration both of this and of other points which have been mentioned, I may draw attention to what seems to me a particularly suggestive case. So-called soldier-or hermit-crabs, are crabs which have adopted the habit of appropriating the empty shells of mollusks. In association with this peculiar habit, the structure of these animals differs very greatly from that of all other crabs. In particular, the hinder part of the body, which occupies the mollusk-shell, and which therefore has ceased to require any hard covering of its own, has been suffered to lose its calcareous integument, and presents a soft fleshy character, quite unlike that of the more exposed parts of the animal. Moreover, this soft fleshy part of the creature is specially adapted to the particular requirements of the creature by having its lateral appendages—i. e. appendages which in other crustacea perform the function of legs—modified so as to act as claspers to the inside of the mollusk-shell; while the tail-end of the part in question is twisted into the form of a spiral, which fits into the spiral of the mollusk-shell. Now, in Keeling Island there is a large kind of crab called Birgus latro, which lives upon land and there feeds upon cocoa-nuts. The whole structure of this crab, it seems to me, unmistakeably resembles the structure of a hermit-crab (see drawings on the next page, Fig. 7). Yet this crab neither lives in the shell of a mollusk, nor is the hinder part of its body in the soft and fleshy condition just described: on the contrary, it is covered with a hard integument like all the other parts of the animal. Consequently, I think we may infer that the ancestors of Birgus were hermit-crabs living in mollusk-shells; but that their descendants gradually relinquished this habit as they gradually became more and more terrestrial, while, concurrently with these changes in habit, the originally soft posterior parts acquired a hard protective covering to take the place of that which was formerly supplied by the mollusk-shell. So that, if so, we now have, within the limits of a single organism, evidence of a whole series of morphological changes in the past history of its species. First, there must have been the great change from an ordinary crab to a hermit-crab in all the respects previously pointed out. Next, there must have been the change back again from a hermit-crab to an ordinary crab, so far as living without the necessity of a mollusk-shell is concerned. From an evolutionary point of view, therefore, we appear to have in the existing structure of Birgus a morphological record of all these changes, and one which gives us a reasonable explanation of why the animal presents the extraordinary appearance which it does. But, on the theory of special creation, it is inexplicable why this land-crab should have been formed on the pattern of a hermit-crab, when it never has need to enter the shell of a mollusk. In other words, its peculiar structure is not specially in keeping with its present habits, although so curiously allied to the similar structure of certain other crabs of totally different habits, in relation to which the peculiarities are of plain and obvious significance.



* * * * *

I will devote the remainder of this chapter to considering another branch of the argument from morphology, to which the case of Birgus serves as a suitable introduction: I mean the argument from rudimentary structures.

Throughout both the animal and vegetable kingdoms we constantly meet with dwarfed and useless representatives of organs, which in other and allied kinds of animals and plants are of large size and functional utility. Thus, for instance, the unborn whale has rudimentary teeth, which are never destined to cut the gums; and throughout its life this animal retains, in a similarly rudimentary condition, a number of organs which never could have been of use to any kind of creature save a terrestrial quadruped. The whole anatomy of its internal ear, for example, has reference to hearing in air—or, as Hunter long ago remarked, "is constructed upon the same principle as in the quadruped"; yet, as Owen says, "the outer opening and passage leading therefrom to the tympanum can rarely be affected by sonorous vibrations of the atmosphere, and indeed they are reduced, or have degenerated, to a degree which makes it difficult to conceive how such vibrations can be propagated to the ear-drum during the brief moments in which the opening may be raised above the water."



Now, rudimentary organs of this kind are of such frequent occurrence, that almost every species presents one or more of them—usually, indeed, a considerable number. How, then, are they to be accounted for? of course the theory of descent with adaptive modification has a simple answer to supply—namely, that when, from changed conditions of life, an organ which was previously useful becomes useless, it will be suffered to dwindle away in successive generations, under the influence of certain natural causes which we shall have to consider in future chapters. On the other hand, the theory of special creation can only maintain that these rudiments are formed for the sake of adhering to an ideal type. Now, here again the former theory appears to be triumphant over the latter; for, without waiting to dispute the wisdom of making dwarfed and useless structures merely for the whimsical motive assigned, surely if such a method were adopted in so many cases, we should expect that in consistency it would be adopted in all cases. This reasonable expectation, however, is far from being realized. We have already seen that in numberless cases, such as that of the fore-limbs of serpents, no vestige of a rudiment is present. But the vacillating policy in the matter of rudiments does not end here; for it is shown in a still more aggravated form where within the limits of the same natural group of organisms a rudiment is sometimes present and sometimes absent. For instance, although in nearly all the numerous species of snakes there are no vestiges of limbs, in the python we find very tiny rudiments of the hind-limbs. (Fig. 8.) Now, is it a worthy conception of deity that, while neglecting to maintain his unity of ideal in the case of nearly all the numerous species of snakes, he should have added a tiny rudiment in the case of the python—and even in that case should have maintained his ideal very inefficiently, inasmuch as only two limbs, instead of four, are represented? how much more reasonable is the naturalistic interpretation; for here the very irregularity of their appearance in different species, which constitutes rudimentary structures one of the crowning difficulties to the theory of special design, furnishes the best possible evidence in favour of hereditary descent; seeing that this irregularity then becomes what may be termed the anticipated expression of progressive dwindling due to inutility. Thus, for example, to return to the case of wings, we have already seen that in an extinct genus of bird, dinornis, these organs were reduced to such an extent as to leave it still doubtful whether so much as the tiny rudiment hypothetically supplied to fig. 6 (p. 61) was present in all the species. And here is another well-known case of another genus of still existing bird, which, as was the case with dinornis, occurs only in new zealand. (Fig. 9.) Upon this island there are no four-footed enemies—either existing or extinct—to escape from which the wings of birds would be of any service. Consequently we can understand why on this island we should meet with such a remarkable dwindling away of wings.



Similarly, the logger-headed duck of South America can only flap along the surface of the water, having its wings considerably reduced though less so than the Apteryx of New Zealand. But here the interesting fact is that the young birds are able to fly perfectly well. Now, in accordance with a general law to be considered in a future chapter, the life-history of an individual organism is a kind of condensed recapitulation of the life-history of its species. Consequently, we can understand why the little chickens of the logger-headed duck are able to fly like all other ducks, while their parents are only able to flap along the surface of the water.

Facts analogous to this reduction of wings in birds which have no further use for them, are to be met with also in insects under similar circumstances. Thus, there are on the island of Madeira somewhere between 500 and 600 species of beetles, which are in large part peculiar to that island, though related to other—and therefore presumably parent—species on the neighbouring continent. Now, no less than 200 species—or nearly half the whole number—are so far deficient in wings that they cannot fly. And, if we disregard the species which are not peculiar to the island—that is to say, all the species which likewise occur on the neighbouring continent, and therefore, as evolutionists conclude, have but recently migrated to the island,—we find this very remarkable proportion. There are altogether 29 peculiar genera, and out of these no less than 23 have all their species in this condition.

Similar facts have been recently observed by the Rev. A. E. Eaton with respect to insects inhabiting Kerguelen Island. All the species which he found on the island—viz. a moth, several flies, and numerous beetles—he found to be incapable of flight; and therefore, as Wallace observes, "as these insects could hardly have reached the islands in a wingless state, even if there were any other known land inhabited by them, which there is not, we must assume that, like the Madeiran insects, they were originally winged, and lost their power of flight because its possession was injurious to them"—Kerguelen Island being "one of the stormiest places on the globe," and therefore a place where insects could rarely afford to fly without incurring the danger of being blown out to sea.

Here is another and perhaps an even more suggestive class of facts.

It is now many years ago since the editors of Silliman's Journal requested the late Professor Agassiz to give them his opinion on the following question. In a certain dark subterranean cave, called the Mammoth cave, there are found some peculiar species of blind fishes. Now the editors of Silliman's Journal wished to know whether Prof. Agassiz would hold that these fish had been specially created in these caves, and purposely devoided of eyes which could never be of any use to them; or whether he would allow that these fish had probably descended from other species, but, having got into the dark cave, gradually lost their eyes through disuse. Prof. Agassiz, who was a believer in special creation, allowed that this ought to constitute a crucial test as between the two theories of special design and hereditary descent. "If physical circumstances," he said, "ever modified organized beings, it should be easily ascertained here." And eventually he gave it as his opinion, that these fish "were created under the circumstances in which they now live, within the limits over which they now range, and with the structural peculiarities which now characterise them."

Since then a great deal of attention has been paid to the fauna of this Mammoth cave, and also to the faunas of other dark caverns, not only in the New, but also in the Old World. In the result, the following general facts have been fully established.

(1) Not only fish, but many representatives of other classes, have been found in dark caves.

(2) Wherever the caves are totally dark, all the animals are blind.

(3) If the animals live near enough to the entrance to receive some degree of light, they may have large and lustrous eyes.

(4) In all cases the species of blind animals are closely allied to species inhabiting the district where the caves occur; so that the blind species inhabiting American caves are closely allied to American species, while those inhabiting European caves are closely allied to European species.

(5) In nearly all cases structural remnants of eyes admit of being detected, in various degrees of obsolescence. In the case of some of the crustaceans of the Mammoth cave the foot-stalks of the eyes are present, although the eyes themselves are entirely absent.

Now, it is evident that all these general facts are in full agreement with the theory of evolution, while they offer serious difficulties to the theory of special creation. As Darwin remarks, it is hard to imagine conditions of life more similar than those furnished by deep limestone caverns under nearly the same climate in the two continents of America and Europe; so that, in accordance with the theory of special creation, very close similarity in the organizations of the two sets of faunas might have been expected. But, instead of this, the affinities of these two sets of faunas are with those of their respective continents—as of course they ought to be on the theory of evolution. Again, what would have been the sense of creating useless foot-stalks for the imaginary support of absent eyes, not to mention all the other various grades of degeneration in other cases? So that, upon the whole, if we agree with the late Prof. Agassiz in regarding these cave animals as furnishing a crucial test between the rival theories of creation and evolution, we must further conclude that the whole body of evidence which they now furnish is weighing on the side of evolution.

So much, then, for a few special instances of what Darwin called rudimentary structures, but what may be more descriptively designated—in accordance with the theory of descent—obsolescent or vestigial structures. It is, however, of great importance to add that these structures are of such general occurrence throughout both the vegetable and animal kingdoms, that, as Darwin has observed, it is almost impossible to point to a single species which does not present one or more of them. In other words, it is almost impossible to find a single species which does not in this way bear some record of its own descent from other species; and the more closely the structure of any species is examined anatomically, the more numerous are such records found to be. Thus, for example, of all organisms that of man has been most minutely investigated by anatomists; and therefore I think it will be instructive to conclude this chapter by giving a list of the more noteworthy vestigial structures which are known to occur in the human body. I will take only those which are found in adult man, reserving for the next chapter those which occur in a transitory manner during earlier periods of his life. But, even as thus restricted, the number of obsolescent structures which we all present in our own persons is so remarkable, that their combined testimony to our descent from a quadrumanous ancestry appears to me in itself conclusive. I mean, that even if these structures stood alone, or apart from any more general evidences of our family relationships, they would be sufficient to prove our parentage. Nevertheless, it is desirable to remark that of course these special evidences which I am about to detail do not stand alone. Not only is there the general analogy furnished by the general proof of evolution elsewhere, but there is likewise the more special correspondence between the whole of our anatomy and that of our nearest zoological allies. Now the force of this latter consideration is so enormous, that no one who has not studied human anatomy can be in a position to appreciate it. For without special study it is impossible to form any adequate idea of the intricacy of structure which is presented by the human form. Yet it is found that this enormously intricate organization is repeated in all its details in the bodies of the higher apes. There is no bone, muscle, nerve, or vessel of any importance in the one which is not answered to by the other. Hence there are hundreds of thousands of instances of the most detailed correspondence, without there being any instances to the contrary, if we pay due regard to vestigial characters. The entire corporeal structure of man is an exact anatomical copy of that which we find in the ape.

My object, then, here is to limit attention to those features of our corporeal structure which, having become useless on account of our change in attitude and habits, are in process of becoming obsolete, and therefore occur as mere vestigial records of a former state of things. For example, throughout the vertebrated series, from fish to mammals, there occurs in the inner corner of the eye a semi-transparent eye-lid, which is called the nictitating membrane. The object of this structure is to sweep rapidly, every now and then, over the external surface of the eye, apparently in order to keep the surface clean. But although the membrane occurs in all classes of the sub-kingdom, it is more prevalent in some than in others—e.g. in birds than in mammals. Even, however, where it does not occur of a size and mobility to be of any use, it is usually represented, in animals above fishes, by a functionless rudiment, as here depicted in the case of man. (Fig. 10.)



Now the organization of man presents so many vestigial structures thus referring to various stages of his long ancestral history, that it would be tedious so much as to enumerate them. Therefore I will yet further limit the list of vestigial structures to be given as examples, by not only restricting these to cases which occur in our own organization; but of them I shall mention only such as refer us to the very last stage of our ancestral history—viz. structures which have become obsolescent since the time when our distinctively human branch of the family tree diverged from that of our immediate forefathers, the Quadrumana.

(1) Muscles of the external ear.—These, which are of large size and functional use in quadrupeds, we retain in a dwindled and useless condition (Fig. 11). This is likewise the case in anthropoid apes; but in not a few other Quadrumana (e.g. baboons, macacus, magots, &c.) degeneration has not proceeded so far, and the ears are voluntarily moveable.



(2) Panniculus carnosis.—A large number of the mammalia are able to move their skin by means of sub-cutaneous muscle—as we see, for instance, in a horse, when thus protecting himself against the sucking of flies. We, in common with the Quadrumana, possess an active remnant of such a muscle in the skin of the forehead, whereby we draw up the eyebrows; but we are no longer able to use other considerable remnants of it, in the scalp and elsewhere,—or, more correctly, it is rarely that we meet with persons who can. But most of the Quadrumana (including the anthropoids) are still able to do so. There are also many other vestigial muscles, which occur only in a small percentage of human beings, but which, when they do occur, present unmistakeable homologies with normal muscles in some of the Quadrumana and still lower animals[5].

[5] See especially Mr. John Wood's papers, Proc. R. S., xiii to xvi, and xviii; also Journ. Anat., i and iii. In this connexion Darwin refers to M. Richard, Annls. d. Sc. Nat. Zoolg., tom. xviii, p. 13, 1852.

(3) Feet.—It is observable that in the infant the feet have a strong deflection inwards, so that the soles in considerable measure face one another. This peculiarity, which is even more marked in the embryo than in the infant (see p. 153), and which becomes gradually less and less conspicuous even before the child begins to walk, appears to me a highly suggestive peculiarity. For it plainly refers to the condition of things in the Quadrumana, seeing that in all these animals the feet are similarly curved inwards, to facilitate the grasping of branches. And even when walking on the ground apes and monkeys employ to a great extent the outside edges of their feet, as does also a child when learning to walk. The feet of a young child are also extraordinarily mobile in all directions, as are those of apes. In order to show these points, I here introduce comparative drawings of a young ape and the portrait of a young male child. These drawings, moreover, serve at the same time to illustrate two other vestigial characters, which have often been previously noticed with regard to the infant's foot. I allude to the incurved form of the legs, and the lateral extension of the great toe, whereby it approaches the thumb-like character of this organ in the Quadrumana. As in the case of the incurved position of the legs and feet, so in this case of the lateral extensibility of the great toe, the peculiarity is even more marked in embryonic than in infant life. For, as Prof. Wyman has remarked with regard to the foetus when about an inch in length, "The great toe is shorter than the others; and, instead of being parallel to them, is projected at an angle from the side of the foot, thus corresponding with the permanent condition of this part in the Quadrumana[6]." So that this organ, which, according to Owen, "is perhaps the most characteristic peculiarity in the human structure," when traced back to the early stages of its development, is found to present a notably less degree of peculiarity.

[6] Proc. Nat. Hist. Soc., Boston, 1863.





(4) Hands.—Dr. Louis Robinson has recently observed that the grasping power of the whole human hand is so surprisingly great at birth, and during the first few weeks of infancy, as to be far in excess of present requirements on the part of a young child. Hence he concludes that it refers us to our quadrumanous ancestry—the young of anthropoid apes being endowed with similar powers of grasping, in order to hold on to the hair of the mother when she is using her arms for the purposes of locomotion. This inference appears to me justifiable, inasmuch as no other explanation can be given of the comparatively inordinate muscular force of an infant's grip. For experiments showed that very young babies are able to support their own weight, by holding on to a horizontal bar, for a period varying from one half to more than two minutes[7]. With his kind permission I here reproduce one of Dr. Robinson's instantaneous, and hitherto unpublished, photographs of a very young infant. This photograph was taken after the above paragraph (3) was written, and I introduce it here because it serves to show incidentally—and perhaps even better than the preceding figure—the points there mentioned with regard to the feet and great toes. Again, as Dr. Robinson observes, the attitude, and the disproportionately large development of the arms as compared with the legs, give all the photographs a striking resemblance to a picture of the chimpanzee "Sally" at the Zoological Gardens. For "invariably the thighs are bent nearly at right angles to the body, and in no case did the lower limbs hang down and take the attitude of the erect position." He adds, "In many cases no sign of distress is evinced, and no cry uttered, until the grasp begins to give way."

[7] Nineteenth Century, November, 1891.



(5) Tail.—The absence of a tail in man is popularly supposed to constitute a difficulty against the doctrine of his quadrumanous descent. As a matter of fact, however, the absence of an external tail in man is precisely what this doctrine would expect, seeing that the nearest allies of man in the quadrumanous series are likewise destitute of an external tail. Far, then, from this deficiency in man constituting any difficulty to be accounted for, if the case were not so—i. e. if man did possess an external tail,—the difficulty would be to understand how he had managed to retain an organ which had been renounced by his most recent ancestors. Nevertheless, as the anthropoid apes continue to present the rudimentary vestiges of a tail in a few caudal vertebrae below the integuments, we might well expect to find a similar state of matters in the case of man. And this is just what we do find, as a glance at these two comparative illustrations will show. (Fig. 15.) Moreover, during embryonic life, both of the anthropoid apes and of man, the tail much more closely resembles that of the lower kinds of quadrumanous animals from which these higher representatives of the group have descended. For at a certain stage of embryonic life the tail, both of apes and of human beings, is actually longer than the legs (see Fig. 16). And at this stage of development, also, the tail admits of being moved by muscles which later on dwindle away. Occasionally, however, these muscles persist, and are then described by anatomists as abnormalities. The following illustrations serve to show the muscles in question, when thus found in adult man.







(6) Vermiform Appendix of the Caecum.—This is of large size and functional use in the process of digestion among many herbivorous animals; while in man it is not only too small to serve any such purpose, but is even a source of danger to life—many persons dying every year from inflammation set up by the lodgement in this blind tube of fruit-stones, &c.

In the orang it is longer than in man (Fig. 18), as it is also in the human foetus proportionally compared with the adult. (Fig. 19.) In some of the lower herbivorous animals it is longer than the entire body.

Like vestigial structures in general, however, this one is highly variable. Thus the above cut (Fig. 19) serves to show that it may sometimes be almost as short in the orang as it normally is in man—both the human subjects of this illustration having been normal.





(7) Ear.—Mr. Darwin writes:—

The celebrated sculptor, Mr. Woolner, informs me of one little peculiarity in the external ear, which he has often observed both in men and women.... The peculiarity consists in a little blunt point, projecting from the inwardly folded margin, or helix. When present, it is developed at birth, and, according to Prof. Ludwig Meyer, more frequently in man than in woman. Mr. Woolner made an exact model of one such case, and sent me the accompanying drawing.... The helix obviously consists of the extreme margin of the ear folded inwards; and the folding appears to be in some manner connected with the whole external ear being permanently pressed backwards. In many monkeys, which do not stand high in the order, as baboons and some species of macacus, the upper portion of the ear is slightly pointed, and the margin is not at all folded inwards; but if the margin were to be thus folded, a slight point would necessarily project towards the centre.... The following wood-cut is an accurate copy of a photograph of the foetus of an orang (kindly sent me by Dr. Nitsche), in which it may be seen how different the pointed outline of the ear is at this period from its adult condition, when it bears a close general resemblance to that of man [including even the occasional appearance of the projecting point shown in the preceding woodcut]. It is evident that the folding over of the tip of such an ear, unless it changed greatly during its further development, would give rise to a point projecting inwards[8].

[8] Descent of Man, 2nd ed., pp. 15-16.





The following woodcut serves still further to show vestigial resemblances between the human ear and that of apes. The last two figures illustrate the general resemblance between the normal ear of foetal man and the ear of an adult orang-outang. The other two figures on the lower line are intended to exhibit occasional modifications of the adult human ear, which approximate simian characters somewhat more closely than does the normal type. It will be observed that in their comparatively small lobes these ears resemble those of all the apes; and that while the outer margin of one is not unlike that of the Barbary ape, the outer margin of the other follows those of the chimpanzee and orang. Of course it would be easy to select individual human ears which present either of these characters in a more pronounced degree; but these ears have been chosen as models because they present both characters in conjunction. The upper row of figures likewise shows the close similarity of hair-tracts, and the direction of growth on the part of the hair itself, in cases where the human ear happens to be of an abnormally hirsute character. But this particular instance (which I do not think has been previously noticed) introduces us to the subject of hair, and hair-growth, in general.



(8) Hair.—Adult man presents rudimentary hair over most parts of the body. Wallace has sought to draw a refined distinction between this vestigial coating and the useful coating of quadrumanous animals, in the absence of the former from the human back. But even this refined distinction does not hold. On the one hand, the comparatively hairless chimpanzee which died last year in the Zoological Gardens (T. calvus) was remarkably denuded over the back; and, on the other hand, men who present a considerable development of hair over the rest of their bodies present it also on their backs and shoulders. Again, in all men the rudimentary hair on the upper and lower arm is directed towards the elbow—a peculiarity which occurs nowhere else in the animal kingdom, with the exception of the anthropoid apes and a few American monkeys, where it presumably has to do with arboreal habits. For, when sitting in trees, the orang, as observed by Mr. Wallace, places its hands above its head with its elbows pointing downwards: the disposition of hair on the arms and fore-arms then has the effect of thatch in turning the rain. Again, I find that in all species of apes, monkeys, and baboons which I have examined (and they have been numerous), the hair on the backs of the hands and feet is continued as far as the first row of phalanges; but becomes scanty, or disappears altogether, on the second row; while it is invariably absent on the terminal row. I also find that the same peculiarity occurs in man. We all have rudimentary hair on the first row of phalanges, both of hands and feet: when present at all, it is more scanty on the second row; and in no case have I been able to find any on the terminal row. In all cases these peculiarities are congenital, and the total absence or partial presence of hair on the second phalanges is constant in different species of Quadrumana. For instance, it is entirely absent in all the chimpanzees, which I have examined, while scantily present in all the orangs. As in man, it occurs in a patch midway between the joints.



Besides showing these two features with regard to the disposition of hair on the human arm and hand, the above woodcut illustrates a third. By looking closely at the arm of the very hairy man from whom the drawing was taken, it could be seen that there was a strong tendency towards a whorled arrangement of the hairs on the backs of the wrists. This is likewise, as a general rule, a marked feature in the arrangement of hair on the same places in the gorilla, orang, and chimpanzee. In the specimen of the latter, however, from which the drawing was taken, this characteristic was not well marked. The downward direction of the hair on the backs of the hands is exactly the same in man as it is in all the anthropoid apes. Again, with regard to hair, Darwin notices that occasionally there appears in man a few hairs in the eyebrows much longer than the others; and that they seem to be representative of similarly long and scattered hairs which occur in the chimpanzee, macacus, and baboons.

Lastly, it may be here more conveniently observed than in the next chapter on Embryology, that at about the sixth month the human foetus is often thickly coated with somewhat long dark hair over the entire body, except the soles of the feet and palms of the hands, which are likewise bare in all quadrumanous animals. This covering, which is called the lanugo, and sometimes extends even to the whole forehead, ears, and face, is shed before birth. So that it appears to be useless for any purpose other than that of emphatically declaring man a child of the monkey.

(9) Teeth.—Darwin writes:—

It appears as if the posterior molar or wisdom-teeth were tending to become rudimentary in the more civilized races of man. These teeth are rather smaller than the other molars, as is likewise the case with the corresponding teeth in the chimpanzee and orang; and they have only two separate fangs.... They are also much more liable to vary, both in structure and in the period of their development, than the other teeth. In the Melanian races, on the other hand, the wisdom-teeth are usually furnished with three separate fangs, and are usually sound [i. e. not specially liable to decay]; they also differ from the other molars in size, less than in the Caucasian races.

Now, in addition to these there are other respects in which the dwindling condition of wisdom-teeth is manifested—particularly with regard to the pattern of their crowns. Indeed, in this respect it would seem that even in the anthropoid apes there is the beginning of a tendency to degeneration of the molar teeth from behind forwards. For if we compare the three molars in the lower jaw of the gorilla, orang, and chimpanzee, we find that the gorilla has five well-marked cusps on all three of them; but that in the orang the cusps are not so pronounced, while in the chimpanzee there are only four of them on the third molar. Now in man it is only the first of these three teeth which normally presents five cusps, both the others presenting only four. So that, comparing all these genera together, it appears that the number of cusps is being reduced from behind forwards; the chimpanzee having lost one of them from the third molar, while man has not only lost this, but also one from the second molar,—and, it may be added, likewise partially (or even totally) from the first molar, as a frequent variation among civilized races. But, on the other hand, variations are often met with in the opposite direction, where the second or the third molar of man presents five cusps—in the one case following the chimpanzee, in the other the gorilla. These latter variations, therefore, may fairly be regarded as reversionary. For these facts I am indebted to the kindness of Mr. C. S. Tomes.



(10) Perforations of the humerus.—The peculiarities which we have to notice under this heading are two in number. First, the supra condyloid foramen is a normal feature in some of the lower Quadrumana (Fig. 25), where it gives passage to the great nerve of the fore-arm, and often also to the great artery. In man, however, it is not a normal feature. Yet it occurs in a small percentage of cases—viz., according to Sir W. Turner, in about one per cent., and therefore is regarded by Darwin as a vestigial character. Secondly, there is inter-condyloid foramen, which is also situated near the lower end of the humerus, but more in the middle of the bone. This occurs, but not constantly, in apes, and also in the human species. From the fact that it does so much more frequently in the bones of ancient—and also of some savage—races of mankind (viz. in 20 to 30 per cent. of cases), Darwin is disposed to regard it also as a vestigial feature. On the other hand, Prof. Flower tells me that in his opinion it is but an expression of impoverished nutrition during the growth of the bone.



(11) Flattening of tibia.—In some very ancient human skeletons, there has also been found a lateral flattening of the tibia, which rarely occurs in any existing human beings, but which appears to have been usual among the earliest races of mankind hitherto discovered. According to Broca, the measurements of these fossil human tibiae resemble those of apes. Moreover, the bone is bent and strongly convex forwards, while its angles are so rounded as to present the nearly oval section seen in apes. It is in association with these ape-like human tibiae that perforated humeri of man are found in greatest abundance.

On the other hand, however, there is reason to doubt whether this form of tibia in man is really a survival from his quadrumanous ancestry. For, as Boyd-Dawkins and Hartmann have pointed out, the degree of flattening presented by some of these ancient human bones is greater than that which occurs in any existing species of anthropoid ape. Of course the possibility remains that the unknown species of ape from which man descended may have had its tibia more flattened than is now observable in any of the existing species. Nevertheless, as some doubt attaches to this particular case, I do not press it—and, indeed, only mention it at all in order that the doubt may be expressed.

Similarly, I will conclude by remarking that several other instances of the survival of vestigial structures in man have been alleged, which are of a still more doubtful character. Of such, for example, are the supposed absence of the genial tubercle in the case of a very ancient jaw-bone of man, and the disposition of valves in human veins. From the former it was argued that the possessor of this very ancient jaw-bone was probably speechless, inasmuch as the tubercle in existing man gives attachment to muscles of the tongue. From the latter it has been argued that all the valves in the veins of the human body have reference, in their disposition, to the incidence of blood-pressure when the attitude of the body is horizontal, or quadrupedal. Now, the former case has already broken down, and I find that the latter does not hold. But we can well afford to lose such doubtful and spurious cases, in view of all the foregoing unquestionable and genuine cases of vestigial structures which are to be met with even within the limits of our own organization—and even when these limits are still further limited by selecting only those instances which refer to the very latest chapter of our long ancestral history.



CHAPTER IV.

EMBRYOLOGY.

We will next consider what of late years has become the most important of the lines of evidence, not only in favour of the general fact of evolution, but also of its history: I mean the evidence which has been yielded by the newest of the sciences, the science of Embryology. But here, as in the analogous case of adult morphology, in order to do justice to the mass of evidence which has now been accumulated, a whole volume would be necessary. As in that previous case, therefore, I must restrict myself to giving an outline sketch of the main facts.

First I will display what in the language of Paley we may call "the state of the argument."

It is an observable fact that there is often a close correspondence between developmental changes as revealed by any chronological series of fossils which may happen to have been preserved, and developmental changes which may be observed during the life-history of now existing individuals belonging to the same group of animals. For instance, the successive development of prongs in the horns of deer-like animals, which is so clearly shown in the geological history of this tribe, is closely reproduced in the life-history of existing deer. Or, in other words, the antlers of an existing deer furnish in their development a kind of resume, or recapitulation, of the successive phases whereby the primitive horn was gradually superseded by horns presenting a greater and greater number of prongs in successive species of extinct deer (Fig. 26). Now it must be obvious that such a recapitulation in the life-history of an existing animal of developmental changes successively distinctive of sundry allied, though now extinct species, speaks strongly in favour of evolution. For as it is of the essence of this theory that new forms arise from older forms by way of hereditary descent, we should antecedently expect, if the theory is true, that the phases of development presented by the individual organism would follow, in their main outlines, those phases of development through which their long line of ancestors had passed. The only alternative view is that as species of deer, for instance, were separately created, additional prongs were successively added to their antlers; and yet that, in order to be so added to successive species every individual deer belonging to later species was required to repeat in his own lifetime the process of successive additions which had previously taken place in a remote series of extinct species. Now I do not deny that this view is a possible view; but I do deny that it is a probable one. According to the evolutionary interpretation of such facts, we can see a very good reason why the life-history of the individual is thus a condensed resume of the life-history of its ancestral species. But according to the opposite view no reason can be assigned why such should be the case. In a previous chapter—the chapter on Classification—we have seen that if each species were created separately, no reason can be assigned why they should all have been turned out upon structural patterns so strongly suggestive of hereditary descent with gradual modifications, or slow divergence—the result being group subordinated to group, with the most generalized (or least developed) forms at the bottom, and the highest products of organization at the top. And now we see—or shall immediately see—that this consideration admits of being greatly fortified by a study of the developmental history of every individual organism. If it would be an unaccountable fact that every separately created species should have been created with close structural resemblances to a certain limited number of other species, less close resemblances to certain further species, and so backwards; assuredly it would be a still more unaccountable fact that every individual of every species should exhibit in its own person a history of developmental change, every term of which corresponds with the structural peculiarities of its now extinct predecessors—and this in the exact historical order of their succession in geological time. The more that we think about this antithesis between the naturalistic and the non-naturalistic interpretations, the greater must we feel the contrast in respect of rationality to become; and, therefore, I need not spend time by saying anything further upon the antecedent standing of the two theories in this respect. The evidence, then, which I am about to adduce from the study of development in the life-histories of individual organisms, will be regarded by me as so much unquestionable evidence in favour of similar processes of development in the life-histories of their respective species—in so far, I mean, as the two sets of changes admit of being proved parallel.



In the only illustration hitherto adduced—viz. that of deers' horns—the series of changes from a one-pronged horn to a fully developed arborescent antler, is a series which takes place during the adult life of the animal; for it is only when the breeding age has been attained that horns are required to appear. But seeing that every animal passes through most of the phases of its development, not only before the breeding age has been attained, but even before the time of its own birth, clearly the largest field for the study of individual development is furnished by embryology. For instance, there is a salamander which differs from most other salamanders in being exclusively terrestrial in its habits. Now, the young of this salamander before their birth are found to be furnished with gills, which, however, they are never destined to use. Yet these gills are so perfectly formed, that if the young salamanders be removed from the body of their mother shortly before birth, and be then immediately placed in water, the little animals show themselves quite capable of aquatic respiration, and will merrily swim about in a medium which would quickly drown their own parent. Here, then, we have both morphological and physiological evidence pointing to the possession of gills by the ancestors of the land salamander.

It would be easy to devote the whole of the present chapter to an enumeration of special instances of the kinds thus chosen for purposes of illustration; but as it is desirable to take a deeper, and therefore a more general view of the whole subject, I will begin at the foundation, and gradually work up from the earliest stages of development to the latest. Before starting, however, I ask the reader to bear in mind one consideration, which must reasonably prevent our anticipating that in every case the life-history of an individual organism should present a full recapitulation of the life-history of its ancestral line of species. Supposing the theory of evolution to be true, it must follow that in many cases it would have been more or less disadvantageous to a developing type that it should have been obliged to reproduce in its individual representatives all the phases of development previously undergone by its ancestry—even within the limits of the same family. We can easily understand, for example, that the waste of material required for building up the useless gills of the embryonic salamanders is a waste which, sooner or later, is likely to be done away with; so that the fact of its occurring at all is in itself enough to show that the change from aquatic to terrestrial habits on the part of this species must have been one of comparatively recent occurrence. Now, in as far as it is detrimental to a developing type that it should pass through any particular ancestral phases of development, we may be sure that natural selection—or whatever other adjustive causes we may suppose to have been at work in the adaptation of organisms to their surroundings—will constantly seek to get rid of this necessity, with the result, when successful, of dropping out the detrimental phases. Thus the foreshortening of developmental history which takes place in the individual lifetime may be expected often to take place, not only in the way of condensation, but also in the way of excision. Many pages of ancestral history may be recapitulated in the paragraphs of embryonic development, while others may not be so much as mentioned. And that this is the true explanation of what embryologists term "direct" development—or of a more or less sudden leap from one phase to another, without any appearance of intermediate phases—is proved by the fact that in some cases both direct and indirect development occur within the same group of organisms, some genera or families having dropped out the intermediate phases which other genera or families retain.

* * * * *

The argument from embryology must be taken to begin with the first beginning of individual life in the ovum. And, in order to understand the bearings of the argument in this its first stage, we must consider the phenomena of reproduction in the simplest form which these phenomena are known to present.

The whole of the animal kingdom is divided into two great groups, which are called the Protozoa and the Metazoa. Similarly, the whole of the vegetable kingdom is divided into the Protophyta and the Metaphyta. The characteristic feature of all the Protozoa and Protophyta is that the organism consists of a single physiological cell, while the characteristic of all the Metazoa and Metaphyta is that the organism consists of a plurality of physiological cells, variously modified to subserve different functions in the economy of the animal or plant, as the case may be. For the sake of brevity, I shall hereafter deal only with the case of animals (Protozoa and Metazoa); but it may throughout be understood that everything which is said applies also to the case of plants (Protophyta and Metaphyta).

A Protozooen (like a Protophyton) is a solitary cell, or a "unicellular organism," while a Metazooen (like a Metaphyton) is a society of cells, or a "multicellular organism." Now, it is only in the multicellular organisms that there is any observable distinction of sex. In all the unicellular organisms the phenomena of reproduction appear to be more or less identical with those of growth. Nevertheless, as these phenomena are here in some cases suggestively peculiar, I will consider them more in detail.

A Protozooen is a single corpuscle of protoplasm which in different species of Protozoa varies in size from more than one inch to less than 1/1000 of an inch in diameter. In some species there is an enveloping cortical substance; in other species no such substance can be detected. Again, in most species there is a nucleus, while in other species no such differentiation of structure has hitherto been observed. Nevertheless, from the fact that the nucleus occurs in the majority of Protozoa, coupled with the fact that the demonstration of this body is often a matter of extreme difficulty, not only in some of the Protozoa where it has been but recently detected, but also in the case of certain physiological cells elsewhere,—from these facts it is not unreasonable to suppose that all the Protozoa possess a nucleus, whether or not it admits of being rendered visible by histological methods thus far at our disposal. If this is the case, we should be justified in saying, as I have said, that a Protozooen is an isolated physiological cell, and, like cells in general, multiplies by means of what Spencer and Haeckel have aptly called a process of discontinuous growth. That is to say, when a cell reaches maturity, further growth takes place in the direction of a severance of its substance—the separated portion thus starting anew as a distinct physiological unit. But, notwithstanding the complex changes which have been more recently observed to take place in the nucleus of some Protozoa prior to their division, the process of multiplication by division may still be regarded as a process of growth, which differs from the previous growth of the individual cell in being attended by a severance of continuity. If we take a suspended drop of gum, and gradually add to its size by allowing more and more gum to flow into it, a point will eventually be reached at which the force of gravity will overcome that of cohesion, and a portion of the drop will fall away from the remainder. Here we have a rough physical simile, although of course no true analogy. In virtue of a continuous assimilation of nutriment, the protoplasm of a cell increases in mass, until it reaches the size at which the forces of disruption overcome those of cohesion—or, in other words, the point at which increase of size is no longer compatible with continuity of substance. Nevertheless, it must not be supposed that the process is thus merely a physical one. The phenomena which occur even in the simplest—or so-called "direct"—cell-division, are of themselves enough to prove that the process is vital, or physiological; and this in a high degree of specialization. But so, likewise, are all processes of growth in organic structures; and therefore the simile of the drop of gum is not to be regarded as a true analogy: it serves only to indicate the fact that when cell-growth proceeds beyond a certain point cell-division ensues. The size to which cells may grow before they thus divide is very variable in different kinds of cells; for while some may normally attain a length of ten or twelve inches, others divide before they measure 1/1000 of an inch. This, however, is a matter of detail, and does not affect the general physiological principles on which we are at present engaged.

Now, as we have seen, a Protozooen is a single cell; for even although in some of the higher forms of protozoal life a colony of cells may be bound together in organic connexion, each of these cells is in itself an "individual," capable of self-nourishment, reproduction, and, generally, of independent existence. Consequently, when the growth of a Protozooen ends in a division of its substance, the two parts wander away from each other as separate organisms. (Fig. 27.)



The next point we have to observe is, that in all cases where a cell or a Protozooen multiplies by way of fissiparous division, the process begins in the nucleus. If the nucleus divides into two parts, the whole cell will eventually divide into two parts, each of which retains a portion of the original nucleus, as represented in the above figure. If the nucleus divides into three, four, or even, as happens in the development of some embryonic tissues, into as many as six parts, the cell will subdivide into a corresponding number, each retaining a portion of the nucleus. Therefore, in all cases of fissiparous division, the seat or origin of the process is the nucleus.

Thus far, then, the phenomena of multiplication are identical in all the lowest or unicellular organisms, and in the constituent cells of all the higher or multicellular. And this is the first point which I desire to make apparent. For where the object is to prove a continuity between the phenomena of growth and reproduction, it is of primary importance to show—1st, that there is such a continuity in the case of all the unicellular organisms, and, 2nd, that there are all the above points of resemblance between the multiplication of cells in the unicellular and in the multicellular organisms.

It remains to consider the points of difference, and, if possible, to show that these do not go to disprove the doctrine of continuity which the points of resemblance so forcibly indicate.

The first point of difference obviously is, that in the case of all the multicellular organisms the two or more "daughter-cells," which are produced by division of the "mother-cell," do not wander away from one another; but, as a rule, they continue to be held in more or less close apposition by means of other cells and binding membranes,—with the result of giving rise to those various "tissues," which in turn go to constitute the material of "organs." I cannot suppose, however, that any advocate of discontinuity will care to take his stand at this point. But, if any one were so foolish as to do so, it would be easy to dislodge him by describing the state of matters in some of the Protozoa where a number of unicellular "individuals" are organically united so as to form a "colony." These cases serve to bridge this distinction between Protozoa and Metazoa, of which therefore we may now take leave.

In the second place, there is the no less obvious distinction that the result of cell-division in the Metazoa is not merely to multiply cells all of the same kind: on the contrary, the process here gives rise to as many different kinds of cells as there are different kinds of tissue composing the adult organism. But no one, I should think, is likely to oppose the doctrine of continuity on the ground of this distinction. For the distinction is clearly one which must necessarily arise, if the doctrine of continuity between unicellular and multicellular organisms be true. In other words, it is a distinction which the theory of evolution itself must necessarily pre-suppose, and therefore it is no objection to the theory that its pre-supposition is realized. Moreover, as we shall see better presently, there is no difficulty in understanding why this distinction should have arisen, so soon as it became necessary (or desirable) that individual cells, when composing a "colony," should conform to the economic principle of the division of labour—a principle, indeed, which is already foreshadowed in the constituent parts of a single cell, since the nucleus has one set of functions and its surrounding protoplasm another.

But now, in the third place, we arrive at a more important distinction, and one which lies at the root of the others still remaining to be considered. I refer to sexual propagation. For it is a peculiarity of the multicellular organisms that, although many of them may likewise propagate themselves by other means (Fig. 28), they all propagate themselves by means of sexual congress. Now, in its essence, sexual congress consists in the fusion of two specialized cells (or, as now seems almost certain, of the nuclei thereof), so that it is out of such a combination that the new individual arises by means of successive cell-divisions, which, beginning in the fertilized ovum, eventually build up all the tissues and organs of the body.



This process clearly indicates very high specialization on the part of germ-cells. For we see by it that although these cells when young resemble all other cells in being capable of self-multiplication by binary division (thus reproducing cells exactly like themselves), when older they lose this power; but, at the same time, they acquire an entirely new and very remarkable power of giving rise to a vast succession of many different kinds of cells, all of which are mutually correlated as to their several functions, so as to constitute a hierarchy of cells—or, to speak literally, a multicellular co-organization. Here it is that we touch the really important distinction between the Protozoa and the Metazoa; for although I have said that some of the higher Protozoa foreshadow this state of matters in forming cell-colonies, it must now be noted that the cells composing such colonies are all of the same kind; and, therefore, that the principle of producing different kinds of cells which, by mutual co-adaptation of functions, shall be capable of constructing a multicellular Metazooen,—this great principle of co-organization is but dimly nascent in the cell-colonies of Protozoa. And its marvellous development in the Metazoa appears ultimately to depend upon the highly specialized character of germ-cells. Even in cases where multicellular organisms are capable of reproducing their kind without the need of any preceding process of fertilization (parthenogenesis), and even in the still more numerous cases where complete organisms are budded forth from any part of their parent organism (gemmation, Fig. 28), there is now very good reason to conclude that these powers of a-sexual reproduction on the part of multicellular organisms are all ultimately due to the specialized character of their germ-cells. For in all these cases the tissues of the parent, from which the budding takes place, were ultimately derived from germ-cells—no matter how many generations of budded organisms may have intervened. And that propagation by budding, &c, in multicellular organisms is thus ultimately due to their propagation by sexual methods, seems to be further shown by certain facts which will have to be discussed at some length in my next volume. Here, therefore, I will mention only one of them—and this because it furnishes what appears to be another important distinction between the Protozoa and the Metazoa.

In nearly all cases where a Protozooen multiplies itself by fission, the process begins by a simple division of the nucleus. But when a Metazooen is developed from a germ-cell, although the process likewise begins by a division of the nucleus, this division is not a simple or direct one; on the contrary, it is inaugurated by a series of processes going on within the nucleus, which are so enormously complex, and withal so beautifully ordered, that to my mind they constitute the most wonderful—if not also the most suggestive—which have ever been revealed by microscopical research. It is needless to say that I refer to the phenomena of karyokinesis. A few pages further on they will be described more fully. For our present purposes it is sufficient to give merely a pictorial illustration of their successive phases; for a glance at such a representation serves to reveal the only point to which attention has now to be drawn—namely, the immense complexity of the processes in question, and therefore the contrast which they furnish to the simple (or "direct") division of the nucleus preparatory to cell-division in the unicellular organisms. Here, then (Fig. 29), we see the complex processes of karyokinesis in the first two stages of egg-cell division. But similar processes continue to repeat themselves in subsequent stages; and this, there is now good reason to believe, throughout all the stages of cell-division, whereby the original egg-cell eventually constructs an entire organism. In other words, all the cells composing all the tissues of a multicellular organism, at all stages of its development, are probably originated by these complex processes, which differ so much from the simple process of direct division in the unicellular organisms[9]. In this important respect, therefore, it does at first sight appear that we have a distinction between the Protozoa and the Metazoa of so pronounced a character, as fairly to raise the question whether cell-division is fundamentally identical in unicellular and in multicellular organisms.

[9] I say "probably," because analogy points in this direction. As a matter of fact, in many cases of tissue-formation karyokinesis has not hitherto been detected. But even if in such cases it does not occur—i. e. if failure to detect its occurrence be not due merely to still remaining imperfections of our histological methods,—the large number of cases in which it has been seen to occur in the formation of sundry tissues are of themselves sufficient to indicate some important difference between cells derived from ova (metazoal), and cells which have not been so derived (protozoal). Which is the point now under discussion.



Lastly, the only other distinction of a physiologically significant kind between a single cell when it occurs as a Protozooen and when it does so as the unfertilized ovum of a Metazooen is, that in the latter case the nucleus discharges from its own substance two minute protoplasmic masses ("polar bodies"), which are then eliminated from the cell altogether. This process, which will be more fully described later on, appears to be of invariable occurrence in the case of all egg-cells, while nothing resembling it has ever been observed in any of the Protozoa.

We must now consider these several points of difference seriatim.

First, with regard to sexual propagation, we have already seen that this is by no means the only method of propagation among the multicellular organisms; and it now remains to add that, on the other hand, there is, to say the least, a suggestive foreshadowing of sexual propagation among the unicellular organisms. For although simple binary fission is here the more usual mode of multiplication, very frequently two (rarely three or more) Protozoa of the same species come together, fuse into a single mass, and thus become very literally "one flesh." This process of "conjugation" is usually (though by no means invariably) followed by a period of quiescent "encystation"; after which the contents of the cyst escape in the form of a number of minute particles, or "spores," and these severally develope into the parent type. Obviously this process of conjugation, when it is thus a preliminary to multiplication, appears to be in its essence the same as fertilization. And if it be objected that encystation and spore-formation in the Protozoa are not always preceded by conjugation, the answer would be that neither is oviparous propagation in the Metazoa invariably preceded by fertilization.

Nevertheless, that there are great distinctions between true sexual propagation and this foreshadowing of it in conjugation I do not deny. The question, however, is whether they be so great as to justify any argument against an historical continuity between them. What, then, are these remaining distinctions? Briefly, as we have seen, they are the extrusion from egg-cells of polar bodies, and the occurrence, both in egg-cells and their products (tissue-cells), of the process of karyokinesis. But, as regards the polar bodies, it is surely not difficult to suppose that, whatever their significance may be, it is probably in some way or another connected with the high specialization of the functions which an egg-cell has to discharge. Nor is there any difficulty in further supposing that, whatever purpose is served by getting rid of polar bodies, the process whereby they are got rid of was originally one of utilitarian development—i. e. a process which at its commencement did not betoken any difference of kind, or breach of continuity, between egg-cells and cells of simpler constitution.

Lastly, with respect to karyokinesis, although it is true that the microscope has in comparatively recent years displayed this apparently important distinction between unicellular and multicellular organisms, two considerations have here to be supplied. The first is, that in some of the Protozoa processes very much resembling those of karyokinesis have already been observed taking place in the nucleus preparatory to its division. And although such processes do not present quite the same appearances as are to be met with in egg-cells, neither do the karyokinetic processes in tissue-cells, which in their sundry kinds exhibit great variations in this respect. Moreover, even if such were not the case, the bare fact that nuclear division is not invariably of the simple or direct character in the case of all Protozoa, is sufficient to show that the distinction now before us—like the one last dealt with—is by no means absolute. As in the case of sexual propagation, so in that of karyokinesis, processes which are common to all the Metazoa are not wholly without their foreshadowings in the Protozoa. And seeing how greatly exalted is the office of egg-cells—and even of tissue-cells—as compared with that of their supposed ancestry in protozoal cells, it seems to me scarcely to be wondered at if their specializations of function should be associated with corresponding peculiarities of structure—a general fact which would in no way militate against the doctrine of evolution. Could we know the whole truth, we should probably find that in order to endow the most primitive of egg-cells with its powers of marshalling its products into a living army of cell-battalions, such an egg-cell must have been passed through a course of developmental specialization of so elaborate a kind, that even the complex processes of karyokinesis are but a very inadequate expression thereof.

Probably I have now said enough to show that, remarkable and altogether exceptional as the properties of germ-cells of the multicellular organisms unquestionably show themselves to be, yet when these properties are traced back to their simplest beginnings in the unicellular organisms, they may fairly be regarded as fundamentally identical with the properties of living cells in general. Thus viewed, no line of real demarcation can be drawn between growth and reproduction, even of the sexual kind. The one process is, so to speak, physiologically continuous with the other; and hence, so far as the pre-embryonic stage of life-history is concerned, the facts cannot fairly be regarded as out of keeping with the theory of evolution.

I will now pass on to consider the embryogeny of the Metazoa, beginning at its earliest stage in the fertilization of the ovum. And here it is that the constructive argument in favour of evolution which is derived from embryology may be said properly to commence. For it is surely in itself a most suggestive fact that all the Metazoa begin their life in the same way, or under the same form and conditions. Omne vivum ex ovo. This is a formula which has now been found to apply throughout the whole range of the multicellular organisms. And seeing, as we have just seen, that the ovum is everywhere a single cell, the formula amounts to saying that, physiologically speaking, every Metazooen begins its life as a Protozooen, and every Metaphyton as a Protophyton[10].

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