it were the very impress of circumstances, passively received by an indifferent matter.
But let us come to the examples. It would be interesting first to institute here a general comparison between plants and animals. One cannot fail to be struck with the parallel progress which has been accomplished, on both sides, in the direction of sexuality. Not only is fecundation itself the same in higher plants and in animals, since it consists, in both, in the union of two nuclei that differ in their properties and structure before their union and immediately after become equivalent to each other; but the preparation of sexual elements goes on in both under like conditions: it consists essentially in the reduction of the number of chromosomes and the rejection of a certain quantity of chromatic substance.[22] Yet vegetables and animals have evolved on independent lines, favored by unlike circumstances, opposed by unlike obstacles. Here are two great series which have gone on diverging. On either line, thousands and thousands of causes have combined to determine the morphological and functional evolution. Yet these infinitely complicated causes have been consummated, in each series, in the same effect. And this effect, could hardly be called a phenomenon of "adaptation": where is the adaptation, where is the pressure of external circumstances? There is no striking utility in sexual generation; it has been interpreted in the most diverse ways; and some very acute enquirers even regard the sexuality of the plant, at least, as a luxury which nature might have dispensed with.[23] But we do not wish to dwell on facts so disputed. The ambiguity of the term "adaptation," and the necessity of transcending both the point of view of mechanical causality and that of anthropomorphic finality, will stand out more clearly with simpler examples. At all times the doctrine of finality has laid much stress on the marvellous structure of the sense-organs, in order to liken the work of nature to that of an intelligent workman. Now, since these organs are found, in a rudimentary state, in the lower animals, and since nature offers us many intermediaries between the pigment-spot of the simplest organisms and the infinitely complex eye of the vertebrates, it may just as well be alleged that the result has been brought about by natural selection perfecting the organ automatically. In short, if there is a case in which it seems justifiable to invoke adaptation, it is this particular one. For there may be discussion about the function and meaning of such a thing as sexual generation, in so far as it is related to the conditions in which it occurs; but the relation of the eye to light is obvious, and when we call this relation an adaptation, we must know what we mean. If, then, we can show, in this privileged case, the insufficiency of the principles invoked on both sides, our demonstration will at once have reached a high degree of generality.
Let us consider the example on which the advocates of finality have always insisted: the structure of such an organ as the human eye. They have had no difficulty in showing that in this extremely complicated apparatus all the elements are marvelously co-ordinated. In order that vision shall operate, says the author of a well-known book on _Final Causes_, "the sclerotic membrane must become transparent in one point of its surface, so as to enable luminous rays to pierce it;...the cornea must correspond exactly with the opening of the socket;...behind this transparent opening there must be refracting media;...there must be a retina[24] at the extremity of the dark chamber;...perpendicular to the retina there must be an innumerable quantity of transparent cones permitting only the light directed in the line of their axes to reach the nervous membrane,"[25] etc. etc. In reply, the advocate of final causes has been invited to assume the evolutionist hypothesis. Everything is marvelous, indeed, if one consider an eye like ours, in which thousands of elements are coördinated in a single function. But take the function at its origin, in the Infusorian, where it is reduced to the mere impressionability (almost purely chemical) of a pigment-spot to light: this function, possibly only an accidental fact in the beginning, may have brought about a slight complication of the organ, which again induced an improvement of the function. It may have done this either directly, through some unknown mechanism, or indirectly, merely through the effect of the advantages it brought to the living being and the hold it thus offered to natural selection. Thus the progressive formation of an eye as well contrived as ours would be explained by an almost infinite number of actions and reactions between the function and the organ, without the intervention of other than mechanical causes.
The question is hard to decide, indeed, when put directly between the function and the organ, as is done in the doctrine of finality, as also mechanism itself does. For organ and function are terms of different nature, and each conditions the other so closely that it is impossible to say _a priori_ whether in expressing their relation we should begin with the first, as does mechanism, or with the second, as finalism requires. But the discussion would take an entirely different turn, we think, if we began by comparing together two terms of the same nature, an organ with an organ, instead of an organ with its function. In this case, it would be possible to proceed little by little to a solution more and more plausible, and there would be the more chance of a successful issue the more resolutely we assumed the evolutionist hypothesis.
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 _OEnothera 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.
But this is just what is claimed to be unnecessary. Physics and chemistry are said to give us the key to everything. Eimer's great work is instructive in this respect. It is well known what persevering effort this biologist has devoted to demonstrating that transformation is brought about by the influence of the external on the internal, continuously exerted in the same direction, and not, as Darwin held, by accidental variations. His theory rests on observations of the highest interest, of which the starting-point was the study of the course followed by the color variation of the skin in certain lizards. Before this, the already old experiments of Dorfmeister had shown that the same chrysalis, according as it was submitted to cold or heat, gave rise to very different butterflies, which had long been regarded as independent species, _Vanessa levana_ and _Vanessa prorsa_: an intermediate temperature produces an intermediate form. We might class with these facts the important transformations observed in a little crustacean, _Artemia salina_, when the salt of the water it lives in is increased or diminished.[32] In these various experiments the external agent seems to act as a cause of transformation. But what does the word "cause" mean here? Without undertaking an exhaustive analysis of the idea of causality, we will merely remark that three very different meanings of this term are commonly confused. A cause may act by _impelling_, _releasing_, or _unwinding_. The billiard-ball, that strikes another, determines its movement by _impelling_. The spark that explodes the powder acts by _releasing_. The gradual relaxing of the spring, that makes the phonograph turn, _unwinds_ the melody inscribed on the cylinder: if the melody which is played be the effect, and the relaxing of the spring the cause, we must say that the cause acts by _unwinding_. What distinguishes these three cases from each other is the greater or less solidarity between the cause and the effect. In the first, the quantity and quality of the effect vary with the quantity and quality of the cause. In the second, neither quality nor quantity of the effect varies with quality and quantity of the cause: the effect is invariable. In the third, the quantity of the effect depends on the quantity of the cause, but the cause does not influence the quality of the effect: the longer the cylinder turns by the action of the spring, the more of the melody I shall hear, but the nature of the melody, or of the part heard, does not depend on the action of the spring. Only in the first case, really, does cause _explain_ effect; in the others the effect is more or less given in advance, and the antecedent invoked is--in different degrees, of course--its occasion rather than its cause. Now, in saying that the saltness of the water is the cause of the transformations of Artemia, or that the degree of temperature determines the color and marks of the wings which a certain chrysalis will assume on becoming a butterfly, is the word "cause" used in the first sense? Obviously not: causality has here an intermediary sense between those of unwinding and releasing. Such, indeed, seems to be Eimer's own meaning when he speaks of the "kaleidoscopic" character of the variation,[33] or when he says that the variation of organized matter works in a definite way, just as inorganic matter crystallizes in definite directions.[34] And it may be granted, perhaps, that the process is a merely physical and chemical one in the case of the color-changes of the skin. But if this sort of explanation is extended to the case of the gradual formation of the eye of the vertebrate, for instance, it must be supposed that the physico-chemistry of living bodies is such that the influence of light has caused the organism to construct a progressive series of visual apparatus, all extremely complex, yet all capable of seeing, and of seeing better and better.[35] What more could the most confirmed finalist say, in order to mark out so exceptional a physico-chemistry? And will not the position of a mechanistic philosophy become still more difficult, when it is pointed out to it that the egg of a mollusc cannot have the same chemical composition as that of a vertebrate, that the organic substance which evolved toward the first of these two forms could not have been chemically identical with that of the substance which went in the other direction, and that, nevertheless, under the influence of light, the same organ has been constructed in the one case as in the other?
The more we reflect upon it, the more we shall see that this production of the same effect by two different accumulations of an enormous number of small causes is contrary to the principles of mechanistic philosophy. We have concentrated the full force of our discussion upon an example drawn from phylogenesis. But ontogenesis would have furnished us with facts no less cogent. Every moment, right before our eyes, nature arrives at identical results, in sometimes neighboring species, by entirely different embryogenic processes. Observations of "heteroblastia" have multiplied in late years,[36] and it has been necessary to reject the almost classical theory of the specificity of embryonic gills. Still keeping to our comparison between the eye of vertebrates and that of molluscs, we may point out that the retina of the vertebrate is produced by an expansion in the rudimentary brain of the young embryo. It is a regular nervous centre which has moved toward the periphery. In the mollusc, on the contrary, the retina is derived from the ectoderm directly, and not indirectly by means of the embryonic encephalon. Quite different, therefore, are the evolutionary processes which lead, in man and in the Pecten, to the development of a like retina. But, without going so far as to compare two organisms so distant from each other, we might reach the same conclusion simply by looking at certain very curious facts of regeneration in one and the same organism.
If the crystalline lens of a Triton be removed, it is regenerated by the iris.[37] Now, the original lens was built out of the ectoderm, while the iris is of mesodermic origin. What is more, in the _Salamandra maculata_, if the lens be removed and the iris left, the regeneration of the lens takes place at the upper part of the iris; but if this upper part of the iris itself be taken away, the regeneration takes place in the inner or retinal layer of the remaining region.[38] Thus, parts differently situated, differently constituted, meant normally for different functions, are capable of performing the same duties and even of manufacturing, when necessary, the same pieces of the machine. Here we have, indeed, the same effect obtained by different combinations of causes.