Let us place side by side the eye of a vertebrate and that of a mollusc such as the common Pecten. We find the same essential parts in each, composed of analogous elements. The eye of the Pecten presents a retina, a cornea, a lens of cellular structure like our own. There is even that peculiar inversion of retinal elements which is not met with, in general, in the retina of the invertebrates. Now, the origin of molluscs may be a debated question, but, whatever opinion we hold, all are agreed that molluscs and vertebrates separated from their common parent-stem long before the appearance of an eye so complex as that of the Pecten. Whence, then, the structural analogy?
Let us question on this point the two opposed systems of evolutionist explanation in turn—the hypothesis of purely accidental variations, and that of a variation directed in a definite way under the influence of external conditions.
The first, as is well known, is presented to-day in two quite different forms. Darwin spoke of very slight variations being accumulated by natural selection. He was not ignorant of the facts of sudden variation; but he thought these "sports," as he called them, were only monstrosities incapable of perpetuating themselves; and he accounted for the genesis of species by an accumulation of insensible variations.[26] Such is still the opinion of many naturalists. It is tending, however, to give way to the opposite idea that a new species comes into being all at once by the simultaneous appearance of several new characters, all somewhat different from the previous ones. This latter hypothesis, already proposed by various authors, notably by Bateson in a remarkable book,[27] has become deeply significant and acquired great force since the striking experiments of Hugo de Vries. This botanist, working on the Œnothera Lamarckiana, obtained at the end of a few generations a certain number of new species. The theory he deduces from his experiments is of the highest interest. Species pass through alternate periods of stability and transformation. When the period of "mutability" occurs, unexpected forms spring forth in a great number of different directions.[28]—We will not attempt to take sides between this hypothesis and that of insensible variations. Indeed, perhaps both are partly true. We wish merely to point out that if the variations invoked are accidental, they do not, whether small or great, account for a similarity of structure such as we have cited.
Let us assume, to begin with, the Darwinian theory of insensible variations, and suppose the occurrence of small differences due to chance, and continually accumulating. It must not be forgotten that all the parts of an organism are necessarily coördinated. Whether the function be the effect of the organ or its cause, it matters little; one point is certain—the organ will be of no use and will not give selection a hold unless it functions. However the minute structure of the retina may develop, and however complicated it may become, such progress, instead of favoring vision, will probably hinder it if the visual centres do not develop at the same time, as well as several parts of the visual organ itself. If the variations are accidental, how can they ever agree to arise in every part of the organ at the same time, in such way that the organ will continue to perform its function? Darwin quite understood this; it is one of the reasons why he regarded variation as insensible.[29] For a difference which arises accidentally at one point of the visual apparatus, if it be very slight, will not hinder the functioning of the organ; and hence this first accidental variation can, in a sense, wait for complementary variations to accumulate and raise vision to a higher degree of perfection. Granted; but while the insensible variation does not hinder the functioning of the eye, neither does it help it, so long as the variations that are complementary do not occur. How, in that case, can the variation be retained by natural selection? Unwittingly one will reason as if the slight variation were a toothing stone set up by the organism and reserved for a later construction. This hypothesis, so little conformable to the Darwinian principle, is difficult enough to avoid even in the case of an organ which has been developed along one single main line of evolution, e.g. the vertebrate eye. But it is absolutely forced upon us when we observe the likeness of structure of the vertebrate eye and that of the molluscs. How could the same small variations, incalculable in number, have ever occurred in the same order on two independent lines of evolution, if they were purely accidental? And how could they have been preserved by selection and accumulated in both cases, the same in the same order, when each of them, taken separately, was of no use?
Let us turn, then, to the hypothesis of sudden variations, and see whether it will solve the problem. It certainly lessens the difficulty on one point, but it makes it much worse on another. If the eye of the mollusc and that of the vertebrate have both been raised to their present form by a relatively small number of sudden leaps, I have less difficulty in understanding the resemblance of the two organs than if this resemblance were due to an incalculable number of infinitesimal resemblances acquired successively: in both cases it is chance that operates, but in the second case chance is not required to work the miracle it would have to perform in the first. Not only is the number of resemblances to be added somewhat reduced, but I can also understand better how each could be preserved and added to the others; for the elementary variation is now considerable enough to be an advantage to the living being, and so to lend itself to the play of selection. But here there arises another problem, no less formidable, viz., how do all the parts of the visual apparatus, suddenly changed, remain so well coördinated that the eye continues to exercise its function? For the change of one part alone will make vision impossible, unless this change is absolutely infinitesimal. The parts must then all change at once, each consulting the others. I agree that a great number of uncoördinated variations may indeed have arisen in less fortunate individuals, that natural selection may have eliminated these, and that only the combination fit to endure, capable of preserving and improving vision, has survived. Still, this combination had to be produced. And, supposing chance to have granted this favor once, can we admit that it repeats the self-same favor in the course of the history of a species, so as to give rise, every time, all at once, to new complications marvelously regulated with reference to each other, and so related to former complications as to go further on in the same direction? How, especially, can we suppose that by a series of mere "accidents" these sudden variations occur, the same, in the same order,—involving in each case a perfect harmony of elements more and more numerous and complex—along two independent lines of evolution?
The law of correlation will be invoked, of course; Darwin himself appealed to it.[30] It will be alleged that a change is not localized in a single point of the organism, but has its necessary recoil on other points. The examples cited by Darwin remain classic: white cats with blue eyes are generally deaf; hairless dogs have imperfect dentition, etc.—Granted; but let us not play now on the word "correlation." A collective whole of solidary changes is one thing, a system of complementary changes—changes so coördinated as to keep up and even improve the functioning of an organ under more complicated conditions—is another. That an anomaly of the pilous system should be accompanied by an anomaly of dentition is quite conceivable without our having to call for a special principle of explanation; for hair and teeth are similar formations,[31] and the same chemical change of the germ that hinders the formation of hair would probably obstruct that of teeth: it may be for the same sort of reason that white cats with blue eyes are deaf. In these different examples the "correlative" changes are only solidary changes (not to mention the fact that they are really lesions, namely, diminutions or suppressions, and not additions, which makes a great difference). But when we speak of "correlative" changes occurring suddenly in the different parts of the eye, we use the word in an entirely new sense: this time there is a whole set of changes not only simultaneous, not only bound together by community of origin, but so coördinated that the organ keeps on performing the same simple function, and even performs it better. That a change in the germ, which influences the formation of the retina, may affect at the same time also the formation of the cornea, the iris, the lens, the visual centres, etc., I admit, if necessary, although they are formations that differ much more from one another in their original nature than do probably hair and teeth. But that all these simultaneous changes should occur in such a way as to improve or even merely maintain vision, this is what, in the hypothesis of sudden variation, I cannot admit, unless a mysterious principle is to come in, whose duty it is to watch over the interest of the function. But this would be to give up the idea of "accidental" variation. In reality, these two senses of the word "correlation" are often interchanged in the mind of the biologist, just like the two senses of the word "adaptation." And the confusion is almost legitimate in botany, that science in which the theory of the formation of species by sudden variation rests on the firmest experimental basis. In vegetables, function is far less narrowly bound to form than in animals. Even profound morphological differences, such as a change in the form of leaves, have no appreciable influence on the exercise of function, and so do not require a whole system of complementary changes for the plant to remain fit to survive. But it is not so in the animal, especially in the case of an organ like the eye, a very complex structure and very delicate function. Here it is impossible to identify changes that are simply solidary with changes which are also complementary. The two senses of the word "correlation" must be carefully distinguished; it would be a downright paralogism to adopt one of them in the premisses of the reasoning, and the other in the conclusion. And this is just what is done when the principle of correlation is invoked in explanations of detail in order to account for complementary variations, and then correlation in general is spoken of as if it were any group of variations provoked by any variation of the germ. Thus, the notion of correlation is first used in current science as it might be used by an advocate of finality; it is understood that this is only a convenient way of expressing oneself, that one will correct it and fall back on pure mechanism when explaining the nature of the principles and turning from science to philosophy. And one does then come back to pure mechanism, but only by giving a new meaning to the word "correlation"—a meaning which would now make correlation inapplicable to the detail it is called upon to explain.
To sum up, if the accidental variations that bring about evolution are insensible variations, some good genius must be appealed to—the genius of the future species—in order to preserve and accumulate these variations, for selection will not look after this. If, on the other hand, the accidental variations are sudden, then, for the previous function to go on or for a new function to take its place, all the changes that have happened together must be complementary. So we have to fall back on the good genius again, this time to obtain the convergence of simultaneous changes, as before to be assured of the continuity of direction of successive variations. But in neither case can parallel development of the same complex structures on independent lines of evolution be due to a mere accumulation of accidental variations. So we come to the second of the two great hypotheses we have to examine. Suppose the variations are due, not to accidental and inner causes, but to the direct influence of outer circumstances. Let us see what line we should have to take, on this hypothesis, to account for the resemblance of eye-structure in two series that are independent of each other from the phylogenetic point of view.
Though molluscs and vertebrates have evolved separately, both have remained exposed to the influence of light. And light is a physical cause bringing forth certain definite effects. Acting in a continuous way, it has been able to produce a continuous variation in a constant direction. Of course it is unlikely that the eye of the vertebrate and that of the mollusc have been built up by a series of variations due to simple chance. Admitting even that light enters into the case as an instrument of selection, in order to allow only useful variations to persist, there is no possibility that the play of chance, even thus supervised from without, should bring about in both cases the same juxtaposition of elements coördinated in the same way. But it would be different supposing that light acted directly on the organized matter so as to change its structure and somehow adapt this structure to its own form. The resemblance of the two effects would then be explained by the identity of the cause. The more and more complex eye would be something like the deeper and deeper imprint of light on a matter which, being organized, possesses a special aptitude for receiving it.
But can an organic structure be likened to an imprint? We have already called attention to the ambiguity of the term "adaptation." The gradual complication of a form which is being better and better adapted to the mold of outward circumstances is one thing, the increasingly complex structure of an instrument which derives more and more advantage from these circumstances is another. In the former case, the matter merely receives an imprint; in the second, it reacts positively, it solves a problem. Obviously it is this second sense of the word "adapt" that is used when one says that the eye has become better and better adapted to the influence of light. But one passes more or less unconsciously from this sense to the other, and a purely mechanistic biology will strive to make the passive adaptation of an inert matter, which submits to the influence of its environment, mean the same as the active adaptation of an organism which derives from this influence an advantage it can appropriate. It must be owned, indeed, that Nature herself appears to invite our mind to confuse these two kinds of adaptation, for she usually begins by a passive adaptation where, later on, she will build up a mechanism for active response. Thus, in the case before us, it is unquestionable that the first rudiment of the eye is found in the pigment-spot of the lower organisms; this spot may indeed have been produced physically, by the mere action of light, and there are a great number of intermediaries between the simple spot of pigment and a complicated eye like that of the vertebrates.—But, from the fact that we pass from one thing to another by degrees, it does not follow that the two things are of the same nature. From the fact that an orator falls in, at first, with the passions of his audience in order to make himself master of them, it will not be concluded that to follow is the same as to lead. Now, living matter seems to have no other means of turning circumstances to good account than by adapting itself to them passively at the outset. Where it has to direct a movement, it begins by adopting it. Life proceeds by insinuation. The intermediate degrees between a pigment-spot and an eye are nothing to the point: however numerous the degrees, there will still be the same interval between the pigment-spot and the eye as between a photograph and a photographic apparatus. Certainly the photograph has been gradually turned into a photographic apparatus; but could light alone, a physical force, ever have provoked this change, and converted an impression left by it into a machine capable of using it?
It may be claimed that considerations of utility are out of place here; that the eye is not made to see, but that we see because we have eyes; that the organ is what it is, and "utility" is a word by which we designate the functional effects of the structure. But when I say that the eye "makes use of" light, I do not merely mean that the eye is capable of seeing; I allude to the very precise relations that exist between this organ and the apparatus of locomotion. The retina of vertebrates is prolonged in an optic nerve, which, again, is continued by cerebral centres connected with motor mechanisms. Our eye makes use of light in that it enables us to utilize, by movements of reaction, the objects that we see to be advantageous, and to avoid those which we see to be injurious. Now, of course, as light may have produced a pigment-spot by physical means, so it can physically determine the movements of certain organisms; ciliated Infusoria, for instance, react to light. But no one would hold that the influence of light has physically caused the formation of a nervous system, of a muscular system, of an osseous system, all things which are continuous with the apparatus of vision in vertebrate animals. The truth is, when one speaks of the gradual formation of the eye, and, still more, when one takes into account all that is inseparably connected with it, one brings in something entirely different from the direct action of light. One implicitly attributes to organized matter a certain capacity sui generis, the mysterious power of building up very complicated machines to utilize the simple excitation that it undergoes.