Thus we have here the origin of a new character, arising from small beginnings, at least in _S. tiliae_, in which species the coloured stripes are a normal specific character. In the other species, _S.

populi_ and _S. ocellata_, we find the beginnings of the same variation, in one more rarely than in the other, and we can imagine that, in the course of time, in these two species, coloured lines over the oblique stripes will arise. In any case these spots are the elements of variation, out of which coloured lines _may_ be evolved, if they are combined in this direction through the agency of natural selection. In _S. populi_ the spots are often small, but sometimes it seems as though several had united to form large spots. Whether a process of selection in this direction will arise in _S. populi_ and _S. ocellata_, or whether it is now going on cannot be determined, since we cannot tell in advance what biological value the marking might have for these two species. It is conceivable that the spots may have no selection-value as far as these species are concerned, and may therefore disappear again in the course of phylogeny, or, on the other hand, that they may be changed in another direction, for instance towards imitation of the rust-red fungoid patches on poplar and willow leaves. In any case we may regard the smallest spots as the initial stages of variation, the larger as a c.u.mulative summation of these.

Therefore either these initial stages must already possess selection-value, or, as I said before: _There must be some other reason for their c.u.mulative summation_. I should like to give one more example, in which we can infer, though we cannot directly observe, the initial stages.

All the Holothurians or sea-cuc.u.mbers have in the skin calcereous bodies of different forms, usually thick and irregular, which make the skin tough and resistant. In a small group of them--the species of Synapta--the calcareous bodies occur in the form of delicate anchors of microscopic size. Up till 1897 these anchors, like many other delicate microscopic structures, were regarded as curiosities, as natural marvels. But a Swedish observer, Oestergren, has recently shown that they have a biological significance: they serve the footless Synapta as auxiliary organs of locomotion, since, when the body swells up in the act of creeping, they press firmly with their tips, which are embedded in the skin, against the substratum on which the animal creeps, and thus prevent slipping backwards. In other Holothurians this slipping is made impossible by the fixing of the tube-feet. The anchors act automatically, sinking their tips towards the ground when the corresponding part of the body thickens, and returning to the original position at an angle of 45 degrees to the upper surface when the part becomes thin again. The arms of the anchor do not lie in the same plane as the shaft, and thus the curve of the arms forms the outermost part of the anchor, and offers no further resistance to the gliding of the animal. Every detail of the anchor, the curved portion, the little teeth at the head, the arms, etc., can be interpreted in the most beautiful way, above all the form of the anchor itself, for the two arms prevent it from swaying round to the side. The position of the anchors, too, is definite and significant; they lie obliquely to the longitudinal axis of the animal, and therefore they act alike whether the animal is creeping backwards or forwards. Moreover, the tips would pierce through the skin if the anchors lay in the longitudinal direction. Synapta burrows in the sand; it first pushes in the thin anterior end, and thickens this again, thus enlarging the hole, then the anterior tentacles displace more sand, the body is worked in a little farther, and the process begins anew. In the first act the anchors are pa.s.sive, but they begin to take an active share in the forward movement when the body is contracted again. Frequently the animal retains only the posterior end buried in the sand, and then the anchors keep it in position, and make rapid withdrawal possible.

Thus we have in these apparently random forms of the calcereous bodies, complex adaptations in which every little detail as to direction, curve, and pointing is exactly determined. That they have selection-value in their present perfected form is beyond all doubt, since the animals are enabled by means of them to bore rapidly into the ground and so to escape from enemies. We do not know what the initial stages were, but we cannot doubt that the little improvements, which occurred as variations of the originally simple slimy bodies of the Holothurians, were preserved because they already possessed selection-value for the Synaptidae. For such minute microscopic structures whose form is so delicately adapted to the role they have to play in the life of the animal, cannot have arisen suddenly and as a whole, and every new variation of the anchor, that is, in the direction of the development of the two arms, and every curving of the shaft which prevented the tips from projecting at the wrong time, in short, every little adaptation in the modelling of the anchor must have possessed selection-value. And that such minute changes of form fall within the sphere of fluctuating variations, that is to say, _that they occur_ is beyond all doubt.

In many of the Synaptidae the anchors are replaced by calcareous rods bent in the form of an S, which are said to act in the same way.

Others, such as those of the genus Ankyroderma, have anchors which project considerably beyond the skin, and, according to Oestergren, serve "to catch plant-particles and other substances" and so mask the animal. Thus we see that in the Synaptidae the thick and irregular calcareous bodies of the Holothurians have been modified and transformed in various ways in adaptation to the footlessness of these animals, and to the peculiar conditions of their life, and we must conclude that the earlier stages of these changes presented themselves to the processes of selection in the form of microscopic variations.

For it is as impossible to think of any origin other than through selection in this case as in the case of the toughness, and the "drip-tips" of tropical leaves. And as these last could not have been produced directly by the beating of the heavy raindrops upon them, so the calcareous anchors of Synapta cannot have been produced directly by the friction of the sand and mud at the bottom of the sea, and, since they are parts whose function is _pa.s.sive_ the Lamarckian factor of use and disuse does not come into question. The conclusion is unavoidable, that the microscopically small variations of the calcareous bodies in the ancestral forms have been intensified and acc.u.mulated in a particular direction, till they have led to the formation of the anchor. Whether this has taken place by the action of natural selection alone, or whether the laws of variation and the intimate processes within the germ-plasm have cooperated will become clear in the discussion of germinal selection. This whole process of adaptation has obviously taken place within the time that has elapsed since this group of sea-cuc.u.mbers lost their tube-feet, those characteristic organs of locomotion which occur in no group except the Echinoderms, and yet have totally disappeared in the Synaptidae. And after all what would animals that live in sand and mud do with tube-feet?

(_c_) _Coadaptation_

Darwin pointed out that one of the essential differences between artificial and natural selection lies in the fact that the former can modify only a few characters, usually only one at a time, while Nature preserves in the struggle for existence all the variations of a species, at the same time and in a purely mechanical way, if they possess selection-value.

Herbert Spencer, though himself an adherent of the theory of selection, declared in the beginning of the nineties that in his opinion the range of this principle was greatly over-estimated, if the great changes which have taken place in so many organisms in the course of ages are to be interpreted as due to this process of selection alone, since no transformation of any importance can be evolved by itself; it is always accompanied by a host of secondary changes. He gives the familiar example of the Giant Stag of the Irish peat, the enormous antlers of which required not only a much stronger skull cap, but also greater strength of the sinews, muscles, nerves and bones of the whole anterior half of the animal, if their ma.s.s was not to weigh down the animal altogether. It is inconceivable, he says, that so many processes of selection should take place _simultaneously_, and we are therefore forced to fall back on the Lamarckian factor of the use and disuse of functional parts. And how, he asks, could natural selection follow two opposite directions of evolution in different parts of the body at the same time, as for instance in the case of the kangaroo, in which the forelegs must have become shorter, while the hind legs and the tail were becoming longer and stronger?

Spencer's main object was to substantiate the validity of the Lamarckian principle, the cooperation of which with selection had been doubted by many. And it does seem as though this principle, if it operates in nature at all, offers a ready and simple explanation of all such secondary variations. Not only muscles, but nerves, bones, sinews, in short all tissues which function actively, increase in strength in proportion as they are used, and conversely they decrease when the claims on them diminish. All the parts, therefore, which depend on the part that varied first, as for instance the enlarged antlers of the Irish Elk, must have been increased or decreased in strength, in exact proportion to the claims made upon them,--just as is actually the case.

But beautiful as this explanation would be, I regard it as untenable, because it a.s.sumes the _transmissibility of functional modifications_ (so-called "acquired" characters), and this is not only undemonstrable, but is scarcely theoretically conceivable, for the secondary variations which accompany or follow the first as correlative variations, occur also in cases in which the animals concerned are sterile and _therefore cannot transmit anything to their descendants_. This is true of _worker bees_, and particularly of _ants_, and I shall here give a brief survey of the present state of the problem as it appears to me.

Much has been written on both sides of this question since the published controversy on the subject in the nineties between Herbert Spencer and myself. I should like to return to the matter in detail, if the s.p.a.ce at my disposal permitted, because it seems to me that the arguments I advanced at that time are equally cogent to-day, notwithstanding all the objections that have since been urged against them. Moreover, the matter is by no means one of subordinate interest; it is the very kernel of the whole question of the reality and value of the principle of selection. For if selection alone does not suffice to explain "_harmonious adaptation_" as I have called Spencer's _Coadaptation_, and if we require to call in the aid of the Lamarckian factor it would be questionable whether selection would explain any adaptations whatever. In this particular case--of worker bees--the Lamarckian factor may be excluded altogether, for it can be demonstrated that here at any rate the effects of use and disuse cannot be transmitted.

But if it be asked why we are unwilling to admit the cooperation of the Darwinian factor of selection and the Lamarckian factor, since this would afford us an easy and satisfactory explanation of the phenomena, I answer: _Because the Lamarckian principle is fallacious, and because by accepting it we close the way towards deeper insight_.

It is not a spirit of combativeness or a desire for self-vindication that induces me to take the field once more against the Lamarckian principle, it is the conviction that the progress of our knowledge is being obstructed by the acceptance of this fallacious principle, since the facile explanation it apparently affords prevents our seeking after a truer explanation and a deeper a.n.a.lysis.

The workers in the various species of ants are sterile, that is to say, they take no regular part in the reproduction of the species, although individuals among them may occasionally lay eggs. In addition to this they have lost the wings, and the _receptaculum seminis_, and their compound eyes have degenerated to a few facets. How could this last change have come about through disuse, since the eyes of workers are exposed to light in the same way as are those of the s.e.xual insects and thus in this particular case are not liable to "disuse" at all? The same is true of the _receptaculum seminis_, which can only have been disused as far as its glandular portion and its stalk are concerned, and also of the wings, the nerves tracheae and epidermal cells of which could not cease to function until the whole wing had degenerated, for the chitinous skeleton of the wing does not function at all in the active sense.

But, on the other hand, the workers in all species have undergone modifications in a positive direction, as, for instance, the greater development of brain. In many species large workers have evolved,--the so-called _soldiers_, with enormous jaws and teeth, which defend the colony,--and in others there are _small_ workers which have taken over other special functions, such as the rearing of the young Aphides.

This kind of division of the workers into two castes occurs among several tropical species of ants, but it is also present in the Italian species, _Colobopsis truncata_. Beautifully as the size of the jaws could be explained as due to the increased use made of them by the "soldiers," or the enlarged brain as due to the mental activities of the workers, the fact of the infertility of these forms is an insurmountable obstacle to accepting such an explanation. Neither jaws nor brain can have been evolved on the Lamarckian principle.

The problem of coadaptation is no easier in the case of the ant than in the case of the Giant Stag. Darwin himself gave a pretty ill.u.s.tration to show how imposing the difference between the two kinds of workers in one species would seem if we translated it into human terms. In regard to the Driver ants (Anomma) we must picture to ourselves a piece of work, "for instance the building of a house, being carried on by two kinds of workers, of which one group was five feet four inches high, the other sixteen feet high."[39]

Although the ant is a small animal as compared with man or with the Irish Elk, the "soldier" with its relatively enormous jaws is hardly less heavily burdened than the Elk with its antlers, and in the ant's case, too, a strengthening of the skeleton, of the muscles, the nerves of the head, and of the legs must have taken place parallel with the enlargement of the jaws. _Harmonious adaptation_ (coadaptation) has here been active in a high degree, and yet these "soldiers" are sterile! There thus remains nothing for it but to refer all their adaptations, positive and negative alike, to processes of selection which have taken place in the rudiments of the workers within the egg and sperm-cells of their parents. There is no way out of the difficulty except the one Darwin pointed out. He himself did not find the solution of the riddle at once. At first he believed that the case of the workers among social insects presented "the most serious special difficulty" in the way of his theory of natural selection; and it was only after it had become clear to him that it was not the sterile insects themselves but their parents that were selected, according as they produced more or less well adapted workers, that he was able to refer to this very case of the conditions among ants "_in order to show the power of natural selection_."[40] He explains his view by a simple but interesting ill.u.s.tration. Gardeners have produced, by means of long continued artificial selection, a variety of Stock, which bears entirely double, and therefore infertile flowers.[41] Nevertheless the variety continues to be reproduced from seed, because, in addition to the double and infertile flowers, the seeds always produce a certain number of single, fertile blossoms, and these are used to reproduce the double variety. These single and fertile plants correspond "to the males and females of an ant-colony, the infertile plants, which are regularly produced in large numbers, to the neuter workers of the colony."

This ill.u.s.tration is entirely apt, the only difference between the two cases consisting in the fact that the variation in the flower is not a useful, but a disadvantageous one, which can only be preserved by artificial selection on the part of the gardener, while the transformations that have taken place parallel with the sterility of the ants are useful, since they procure for the colony an advantage in the struggle for existence, and they are therefore preserved by natural selection. Even the sterility itself in this case is not disadvantageous, since the fertility of the true females has at the same time considerably increased. We may therefore regard the sterile forms of ants, which have gradually been adapted in several directions to varying functions, _as a certain proof_ that selection really takes place in the germ-cells of the fathers and mothers of the workers, and that _special complexes of primordia_ (_ids_) are present in the workers and in the males and females, and these complexes contain the primordia of the individual parts (_determinants_). But since all living ent.i.ties vary, the determinants must also vary, now in a favourable, now in an unfavourable direction. If a female produces eggs, which contain favourably varying determinants in the worker-ids, then these eggs will give rise to workers modified in the favourable direction, and if this happens with many females, the colony concerned will contain a better kind of worker than other colonies.

I digress here in order to give an account of the intimate processes, which, according to my view, take place within the germ-plasm, and which I have called "_germinal selection_." These processes are of importance since they form the roots of variation, which in its turn is the root of natural selection. I cannot here do more than give a brief outline of the theory in order to show how the Darwin-Wallace theory of selection has gained support from it.

With others, I regard the minimal amount of substance which is contained within the nucleus of the germ-cells, in the form of rods, bands, or granules, as the _germ-substance_ or _germ-plasm_, and I call the individual granules _ids_. There is always a multiplicity of such ids present in the nucleus, either occurring individually, or united in the form of rods or bands (chromosomes). Each id contains the primary const.i.tuents of a _whole_ individual, so that several ids are concerned in the development of a new individual.

In every being of complex structure thousands of primary const.i.tuents must go to make up a single id; these I call _determinants_, and I mean by this name very small individual particles, far below the limits of microscopic visibility, vital units which feed, grow, and multiply by division. These determinants control the parts of the developing embryo,--in what manner need not here concern us. The determinants differ among themselves, those of a muscle are differently const.i.tuted from those of a nerve-cell or a glandular cell, etc., and every determinant is in its turn made up of minute vital units, which I call _bioph.o.r.es_, or the bearers of life.

According to my view, these determinants not only a.s.similate, like every other living unit, but they _vary_ in the course of their growth, as every living unit does; they may vary qualitatively if the elements of which they are composed vary, they may grow and divide more or less rapidly, and their variations give rise to _corresponding_ variations of the organ, cell, or cell-group which they determine. That they are undergoing ceaseless fluctuations in regard to size and quality seems to me the inevitable consequence of their unequal nutrition; for although the germ-cell as a whole usually receives sufficient nutriment, minute fluctuations in the amount carried to different parts within the germ-plasm cannot fail to occur.

Now, if a determinant, for instance of a sensory cell, receives for a considerable time more abundant nutriment than before, it will grow more rapidly--become bigger, and divide more quickly, and, later, when the id concerned develops into an embryo, this sensory cell will become stronger than in the parents, possibly even twice as strong.

This is an instance of a _hereditary individual variation_, arising from the germ.

The nutritive stream which, according to our hypothesis, favours the determinant _N_ by chance, that is, for reasons unknown to us, may remain strong for a considerable time, or may decrease again; but even in the latter case it is conceivable that the ascending movement of the determinant may continue, because the strengthened determinant now _actively_ nourishes itself more abundantly,--that is to say, it attracts the nutriment to itself, and to a certain extent withdraws it from its fellow-determinants. In this way, it may--as it seems to me--get into _permanent upward movement, and attain a degree of strength from which there is no falling back_. Then positive or negative selection sets in, favouring the variations which are advantageous, setting aside those which are disadvantageous.

In a similar manner a _downward_ variation of the determinants may take place, if its progress be started by a diminished flow of nutriment. The determinants which are weakened by this diminished flow will have less affinity for attracting nutriment because of their diminished strength, and they will a.s.similate more feebly and grow more slowly, unless chance streams of nutriment help them to recover themselves. But, as will presently be shown, a change of direction cannot take place at _every_ stage of the degenerative process. If a certain critical stage of downward progress be pa.s.sed, even favourable conditions of food-supply will no longer suffice permanently to change the direction of the variation. Only two cases are conceivable; if the determinant corresponds to a _useful_ organ, only its removal can bring back the germ-plasm to its former level; therefore personal selection removes the id in question, with its determinants, from the germ-plasm, by causing the elimination of the individual in the struggle for existence. But there is another conceivable case; the determinants concerned may be those of an organ which has become _useless_, and they will then continue un.o.bstructed, but with exceeding slowness, along the downward path, until the organ becomes vestigial, and finally disappears altogether.

The fluctuations of the determinants. .h.i.ther and thither may thus be transformed into a lasting ascending or descending movement; and _this is the crucial point of these germinal processes_.

This is not a fantastic a.s.sumption; we can read it in the fact of the degeneration of disused parts. _Useless organs are the only ones which are not helped to ascend again by personal selection, and therefore in their case alone can we form any idea of how the primary const.i.tuents behave, when they are subject solely to intra-germinal forces_.

The whole determinant system of an id, as I conceive it, is in a state of continual fluctuation upwards and downwards. In most cases the fluctuations will counteract one another, because the pa.s.sive streams of nutriment soon change, but in many cases the limit from which a return is possible will be pa.s.sed, and then the determinants concerned will continue to vary in the same direction, till they attain positive or negative selection-value. At this stage personal selection intervenes and sets aside the variation if it is disadvantageous, or favours--that is to say, preserves--it if it is advantageous. Only _the determinant of a useless organ is uninfluenced by personal selection_, and, as experience shows, it sinks downwards; that is, the organ that corresponds to it degenerates very slowly but uninterruptedly till, after what must obviously be an immense stretch of time, it disappears from the germ-plasm altogether.

Thus we find in the fact of the degeneration of disused parts the proof that not all the fluctuations of a determinant return to equilibrium again, but that, when the movement has attained to a certain strength, it continues _in the same direction_. We have entire certainty in regard to this as far as the downward progress is concerned, and we must a.s.sume it also in regard to ascending variations, as the phenomena of artificial selection certainly justify us in doing. If the j.a.panese breeders were able to lengthen the tail-feathers of the c.o.c.k to six feet, it can only have been because the determinants of the tail-feathers in the germ-plasm had already struck out a path of ascending variation, and this movement was taken advantage of by the breeder, who continually selected for reproduction the individuals in which the ascending variation was most marked. For all breeding depends upon the unconscious selection of germinal variations.

Of course these germinal processes cannot be proved mathematically, since we cannot actually see the play of forces of the pa.s.sive fluctuations and their causes. We cannot say how great these fluctuations are, and how quickly or slowly, how regularly or irregularly they change. Nor do we know how far a determinant must be strengthened by the pa.s.sive flow of the nutritive stream if it is to be beyond the danger of unfavourable variations, or how far it must be weakened pa.s.sively before it loses the power of recovering itself by its own strength. It is no more possible to bring forward actual proofs in this case than it was in regard to the selection-value of the initial stages of an adaptation. But if we consider that all heritable variations must have their roots in the germ-plasm, and further, that when personal selection does not intervene, that is to say, in the case of parts which have become useless, a degeneration of the part, and therefore also of its determinant must inevitably take place; then we must conclude that processes such as I have a.s.sumed are running their course within the germ-plasm, and we can do this with as much certainty as we were able to infer, from the phenomena of adaptation, the selection-value of their initial stages. The fact of the degeneration of disused parts seems to me to afford irrefutable proof that the fluctuations within the germ-plasm _are the real root of all hereditary variation_, and the preliminary condition for the occurrence of the Darwin-Wallace factor of selection. Germinal selection supplies the stones out of which personal selection builds her temples and palaces: _adaptations_. The importance for the theory of the process of degeneration of disused parts cannot be over-estimated, especially when it occurs in sterile animal forms, where we are free from the doubt as to the alleged _Lamarckian factor_ which is apt to confuse our ideas in regard to other cases.

If we regard the variation of the many determinants concerned in the transformation of the female into the sterile worker as having come about through the gradual transformation of the ids into worker-ids, we shall see that the germ-plasm of the s.e.xual ants must contain three kinds of ids, male, female, and worker ids, or if the workers have diverged into soldiers and nest-builders, then four kinds. We understand that the worker-ids arose because their determinants struck out a useful path of variation, whether upward or downward, and that they continued in this path until the highest attainable degree of utility of the parts determined was reached. But in addition to the organs of positive or negative selection-value, there were some which were indifferent as far as the success and especially the functional capacity of the workers was concerned: wings, ovarian tubes, _receptaculum seminis_, a number of the facets of the eye, perhaps even the whole eye. As to the ovarian tubes it is is possible that their degeneration was an advantage for the workers, in saving energy, and if so selection would favour the degeneration; but how could the presence of eyes diminish the usefulness of the workers to the colony?

or the minute _receptaculum seminis_, or even the wings? These parts have therefore degenerated _because they were of no further value to the insect_. But if selection did not influence the setting aside of these parts because they were neither of advantage nor of disadvantage to the species, then the Darwinian factor of selection is here confronted with a puzzle which it cannot solve alone, but which at once becomes clear when germinal selection is added. For the determinants of organs that have no further value for the organism, must, as we have already explained, embark on a gradual course of retrograde development.

In ants the degeneration has gone so far that there are no wing-rudiments present in _any_ species, as is the case with so many b.u.t.terflies, flies, and locusts, but in the larvae the imaginable discs of the wings are still laid down. With regard to the ovaries, degeneration has reached different levels in different species of ants, as has been shown by the researches of my former pupil, Elizabeth Bickford. In many species there are twelve ovarian tubes, and they decrease from that number to one; indeed, in one species no ovarian tube at all is present. So much at least is certain from what has been said, that in this case _everything_ depends on the fluctuations of the elements of the germ-plasm. Germinal selection, here as elsewhere, presents the variations of the determinants, and personal selection favours or rejects these, or,--if it be a question of organs which have become useless,--it does not come into play at all, and allows the descending variation free course.

It is obvious that even the problem of _coadaptation in sterile animals_ can thus be satisfactorily explained. If the determinants are oscillating upwards and downwards in continual fluctuation, and varying more p.r.o.nouncedly now in one direction now in the other, useful variations of every determinant will continually present themselves anew, and may, in the course of generations, be combined with one another in various ways. But there is one character of the determinants that greatly facilitates this complex process of selection, that, after a certain limit has been reached, they go on varying in the same direction. From this it follows that development along a path once struck out may proceed without the continual intervention of personal selection. This factor only operates, so to speak, at the beginning, when it selects the determinants which are varying in the right direction, and again at the end, when it is necessary to put a check upon further variation. In addition to this, enormously long periods have been available for all these adaptations, as the very gradual transition stages between females and workers in many species plainly show, and thus this process of transformation loses the marvellous and mysterious character that seemed at the first glance to invest it, and takes rank, without any straining, among the other processes of selection. It seems to me that, from the facts that sterile animal forms can adapt themselves to new vital functions, their superfluous parts degenerate, and the parts more used adapt themselves in an ascending direction, those less used in a descending direction, we must draw the conclusion that harmonious adaptation here comes about _without the cooperation of the Lamarckian principle_.

This conclusion once established, however, we have no reason to refer the thousands of cases of harmonious adaptation, which occur in exactly the same way among other animals or plants, to a principle, the _active intervention of which in the transformation of species is nowhere proved. We do not require it to explain the facts, and therefore we must not a.s.sume it._

The fact of coadaptation, which was supposed to furnish the strongest argument against the principle of selection, in reality yields the clearest evidence in favour of it. We _must_ a.s.sume it, _because no other possibility of explanation is open to us, and because these adaptations actually exist, that is to say, have really taken place_.

With this conviction I attempted, as far back as 1894, when the idea of germinal selection had not yet occurred to me, to make "harmonious adaptation" (coadaptation) more easily intelligible in some way or other, and so I was led to the idea, which was subsequently expounded in detail by Baldwin, and Lloyd Morgan, and also by Osborn, and Gulick as _Organic Selection_. It seemed to me that it was not necessary that all the germinal variations required for secondary variations should have occurred _simultaneously_, since, for instance, in the case of the stag, the bones, muscles, sinews, and nerves would be incited by the increasing heaviness of the antlers to greater activity in _the individual life_, and so would be strengthened. The antlers can only have increased in size by very slow degrees, so that the muscles and bones may have been able to keep pace with their growth in the individual life, until the requisite germinal variations presented themselves. In this way a disharmony between the increasing weight of the antlers and the parts which support and move them would be avoided, since time would be given for the appropriate germinal variations to occur, and so to set agoing the _hereditary_ variation of the muscles, sinews and bones.[42]

I still regard this idea as correct, but I attribute less importance to "organic selection" than I did at that time, in so far that I do not believe that it _alone_ could effect complex harmonious adaptations. Germinal selection now seems to me to play the chief part in bringing about such adaptations. Something the same is true of the principle I have called _Panmixia_. As I became more and more convinced, in the course of years, that the _Lamarckian principle_ ought not to be called in to explain the dwindling of disused parts, I believed that this process might be simply explained as due to the cessation of the conservative effect of natural selection. I said to myself that, from the moment in which a part ceases to be of use, natural selection withdraws its hand from it, and then it must inevitably fall from the height of its adaptiveness, because inferior variants would have as good a chance of persisting as better ones, since all grades of fitness of the part in question would be mingled with one another indiscriminately. This is undoubtedly true, as Romanes pointed out ten years before I did, and this mingling of the bad with the good probably does bring about a deterioration of the part concerned. But it cannot account for the steady diminution, which always occurs when a part is in process of becoming rudimentary, and which goes on until it ultimately disappears altogether. The process of dwindling cannot therefore be explained as due to panmixia alone: we can only find a sufficient explanation in germinal selection.

IV. DERIVATIVES OF THE THEORY OF SELECTION

The impetus in all directions given by Darwin through his theory of selection has been an immeasurable one, and its influence is still felt. It falls within the province of the historian of science to enumerate all the ideas which, in the last quarter of the nineteenth century, grew out of Darwin's theories, in the endeavour to penetrate more deeply into the problem of the evolution of the organic world.

Within the narrow limits to which this paper is restricted, I cannot attempt to discuss any of these.

V. ARGUMENTS FOR THE REALITY OF THE PROCESSES OF SELECTION

(_a_) _s.e.xual Selection_

s.e.xual selection goes hand in hand with natural selection. From the very first I have regarded s.e.xual selection as affording an extremely important and interesting corroboration of natural selection, but, singularly enough, it is precisely against this theory that an adverse judgment has been p.r.o.nounced in so many quarters, and it is only quite recently, and probably in proportion as the wealth of facts in proof of it penetrates into a wider circle, that we seem to be approaching a more general recognition of this side of the problem of adaptations.

Thus Darwin's words in his preface to the second edition (1874) of his book, _The Descent of Man and s.e.xual Selection_, are being justified: "My conviction as to the operation of natural selection remains unshaken," and further, "If naturalists were to become more familiar with the idea of s.e.xual selection, it would, I think, be accepted to a much greater extent, and already it is fully and favourably accepted by many competent judges." Darwin was able to speak thus because he was already acquainted with an immense ma.s.s of facts, which, taken together, yield overwhelming evidence of the validity of the principle of s.e.xual selection.

_Natural selection_ chooses out for reproduction the individuals that are best equipped for the struggle for existence, and it does so at every stage of development; it thus improves the species in all its stages and forms. _s.e.xual selection_ operates only on individuals that are already capable of reproduction, and does so only in relation to the attainment of reproduction. It arises from the rivalry of one s.e.x, usually the male, for the possession of the other, usually the female. Its influence can therefore only _directly_ affect one s.e.x, in that it equips it better for attaining possession of the other. But the effect may extend indirectly to the female s.e.x, and thus the whole species may be modified, without, however, becoming any more capable of resistance in the struggle for existence, for s.e.xual selection only gives rise to adaptations which are likely to give their possessor the victory over rivals in the struggle for possession of the female, and which are therefore peculiar to the wooing s.e.x: the manifold "secondary s.e.xual characters." The diversity of these characters is so great that I cannot here attempt to give anything approaching a complete treatment of them, but I should like to give a sufficient number of examples to make the principle itself, in its various modes of expression, quite clear.

One of the chief preliminary postulates of s.e.xual selection is the unequal number of individuals in the two s.e.xes, for if every male immediately finds his mate there can be no compet.i.tion for the possession of the female. Darwin has shown that, for the most part, the inequality between the s.e.xes is due simply to the fact that there are more males than females, and therefore the males must take some pains to secure a mate. But the inequality does not always depend on the numerical preponderance of the males, it is often due to polygamy; for, if one male claims several females, the number of females in proportion to the rest of the males will be reduced. Since it is almost always the males that are the wooers, we must expect to find the occurrence of secondary s.e.xual characters chiefly among them, and to find it especially frequent in polygamous species. And this is actually the case.

If we were to try to guess--without knowing the facts--what means the male animals make use of to overcome their rivals in the struggle for the possession of the female, we might name many kinds of means, but it would be difficult to suggest any which is not actually employed in some animal group of other. I begin with the mere difference in strength, through which the male of many animals is so sharply distinguished from the female, as, for instance, the lion, walrus, "sea-elephant," and others. Among these the males fight violently for the possession of the female, who falls to the victor in the combat.

In this simple case no one can doubt the operation of selection, and there is just as little room for doubt as to the selection-value of the initial stages of the variation. Differences in bodily strength are apparent even among human beings, although in their case the struggle for the possession of the female is no longer decided by bodily strength alone.

Combats between male animals are often violent and obstinate, and the employment of the natural weapons of the species in this way has led to perfecting of these, e.g. the tusks of the boar, the antlers of the stag, and the enormous, antler-like jaws of the stag-beetle. Here again it is impossible to doubt that variations in these organs presented themselves, and that these were considerable enough to be decisive in combat, and so to lead to the improvement of the weapon.