# The Principles of Biology Vol. I/Chapter III.12

CHAPTER XII.
INDIRECT EQUILIBRATION.
§ 164. Besides those perturbations produced in any organism by special disturbing forces, there are ever going on many others—the reverberating effects of disturbing forces previously experienced by the individual, or by ancestors; and the multiplied deviations of function so caused imply multiplied deviations of structure. In § 155 there was re-illustrated the truth, set forth at length when treating of Adaptation ( § 69), that an organism in a state of moving equilibrium, cannot have extra function thrown on any organ, and extra growth produced in such organ, without correlative changes being entailed throughout all other functions, and eventually throughout all other organs. And when treating of Variation ( § 90), we saw that individuals which have been made, by their different circumstances, to deviate functionally and structurally from the average type in different directions, will bequeath to their joint offspring, compound perturbations of function and compound deviations of structure, endlessly varied in their kinds and amounts.
Now if the individuals of a species are thus necessarily made unlike in countless ways and degrees—if in one individual the amount of energy in a particular direction is greater than in any other individual, or if here a peculiar combination gives a resulting action which is not found elsewhere; then, among all the individuals, some will be less liable than others to have their equilibria overthrown by ​a particular incident force previously unexperienced. Unless the change in the environment is so violent as to be universally fatal to the species, it must affect more or less differently the slightly-different moving equilibria which the members of the species present. Inevitably some will be more stable than others when exposed to this new or altered factor. That is to say, those individuals whose functions are most out of equilibrium with the modified aggregate of external forces, will be those to die; and those will survive whose functions happen to be most nearly in equilibrium with the modified aggregate of external forces.
But this survival of the fittest implies multiplication of the fittest. Out of the fittest thus multiplied there will, as before, be an overthrowing of the moving equilibrium wherever it presents the least opposing force to the new incident force. And by the continual destruction of the individuals least capable of maintaining their equilibria in presence of this new incident force, there must eventually be reached an altered type completely in equilibrium with the altered conditions.
§ 165. This survival of the fittest, which I have here sought to express in mechanical terms, is that which Mr. Darwin has called "natural selection, or the preservation of ​favoured races in the struggle for life." That there goes on a process of this kind throughout the organic world, Mr. Darwin's great work on the Origin of Species has shown to the satisfaction of nearly all naturalists. Indeed, when once enunciated, the truth of his hypothesis is so obvious as scarcely to need proof. Though evidence may be required to show that natural selection accounts for everything ascribed to it, yet no evidence is required to show that natural selection has always been going on, is going on now, and must ever continue to go on. Recognizing this as an à priori certainty, let us contemplate it under its two distinct aspects.
That organisms which live, thereby prove themselves fit for living, in so far as they have been tried, while organisms which die, thereby prove themselves in some respects unfitted for living, are facts no less manifest than is the fact that this self-purification of a species must tend ever to insure adaptation between it and its environment. This adaptation may be either so maintained or so produced. Doubtless many who have looked at Nature with philosophic eyes, have observed that death of the worst and multiplication of the best, tends towards maintenance of a constitution in harmony with surrounding circumstances. That the average vigour of any race would be diminished did the diseased and feeble habitually survive and propagate; and that the destruction of such, through failure to fulfil some of the conditions to life, leaves behind those which are able to fulfil the conditions to life, and thus keeps up the average fitness to the conditions of life; are almost self-evident truths. But to recognize "Natural Selection" as a means of preserving an already-established balance between the powers of a species and the forces to which it is subject, is to recognize only its simplest and most general mode of action. It is the more special mode of action with which we are here concerned. This more special mode of action, Mr. Darwin has been the first to recognize as an all-important factor, though, besides his co-discoverer Mr. A. R. Wallace, some others have perceived ​that such a factor is at work. To him we owe due appreciation of the fact that natural selection is capable of producing fitness between organisms and their circumstances. He has worked up an enormous mass of evidence showing that this "preservation of favoured races in the struggle for life," is an ever-acting cause of divergence among organic forms. He has traced out the involved results of the process with marvellous subtlety. He has shown how hosts of otherwise inexplicable facts, are accounted for by it. In brief, he has proved that the cause he alleges is a true cause; that it is a cause which we see habitually in action; and that the results to be inferred from it are in harmony with the phenomena which the Organic Creation presents, both as a whole and in its details. Let us glance at a few of the more important interpretations which the hypothesis furnishes.
A soil possessing some ingredient in unusual quantity, may supply to a plant an excess of the matter required for certain of its tissues; and may cause all the parts formed of such tissues to be abnormally developed. Suppose that among these are the hairs clothing its surfaces, including those which grow on its seeds. Thus furnished with somewhat longer fibres, its seeds, when shed, are carried a little further by the wind before they fall to the ground. The plants growing from them, being rather more widely dispersed than those produced by other individuals of the same species, will be less liable to smother one another; and a greater number may therefore reach maturity and fructify. Supposing the next generation subject to the same peculiarity of nutrition, some of the seeds borne by its members will not simply inherit this increased development of hairs, but will carry it further; and these, still more advantaged in the same way as before, will, on the average, have still more numerous chances of continuing the race. Thus, by the survival, generation after generation, of those possessing these longer hairs, and the inheritance of successive increments of growth in the hairs, there may result a seed deviating greatly ​from the original. Other individuals of the same species, subject to the different physical conditions of other localities, may develop somewhat thicker or harder coatings to their seeds: so rendering their seeds less digestible by the birds which devour them. Such thicker-coated seeds, by escaping undigested more frequently than thinner-coated ones, will have additional chances of growing and leaving offspring; and this process, acting in a cumulative manner season after season, will produce a seed diverging in another direction from the ancestral type. Again, elsewhere, some modification in the physiologic actions of the plant may lead to an unusual secretion of an essential oil in the seeds; rendering them unpalatable to creatures which would otherwise feed on them: so giving an advantage to the variety in its rate of multiplication. This incidental peculiarity, proving a preservative, will, as before, be increased by natural selection until it constitutes another divergence. Now in such cases, we see that plants may become better adapted, or re-adapted, to the aggregate of surrounding agencies, not through any direct action of such agencies on them, but through their indirect action—through the destruction by them of the individuals least congruous with them, and the survival of those most congruous with them. All these slight variations of function and structure, arising among the members of a species, serve as so many experiments; the great majority of which fail, but a few of which succeed. Just as each plant bears a multitude of seeds, out of which some two or three happen to fulfil all the conditions required for reaching maturity and continuing the race; so each species is ever producing numerous slightly-modified forms, deviating in all directions from the average, out of which most fit the surrounding conditions no better than their parents, or not so well, but some few of which fit the conditions better; and, doing so, are enabled the better to preserve themselves, and to produce offspring similarly capable of preserving themselves. Among animals the like process results in ​the like development of various structures which cannot have been affected by the performance of functions—their functions being purely passive. The thick shell of a mollusk cannot have arisen from direct reactions of the organism against the external actions to which it is exposed; but it is quite explicable as an effect of the survival, generation after generation, of individuals whose thicker coverings protected them against enemies. Similarly with such dermal structure as that of the tortoise. Though we have evidence that the skin, where it is continually exposed to pressure and friction, may thicken, and so re-establish the equilibrium by opposing a greater inner force to a greater outer force; yet we have no evidence that a coat of armour like that of the tortoise can be so produced. Nor, indeed, are the conditions under which alone its production in such a manner could be accounted for, fulfilled; since the surface of the tortoise is not exposed to greater pressure and friction than the surfaces of other creatures. This massive carapace, and the strangely-adapted osseous frame-work which supports it, are inexplicable as results of evolution, unless through the process of natural selection. So, too, is it with the formation of odoriferous glands in some mammals, or the growth of such excrescences as those of the camel. Thus, in short, is it with all those organs of animals which do not play active parts.
Besides giving us explanations of structural characters that are otherwise unaccountable, Mr. Darwin shows how natural selection explains peculiar relations between individuals in certain species. Such facts as the dimorphism of the primrose and other flowers, he proves to be in harmony with his hypothesis, though stumbling-blocks to all other hypotheses. The various differences which accompany difference of sex, sometimes slight, sometimes very great, are similarly accounted for. As before suggested ( § 79), natural selection appears capable of producing and maintaining the right proportion of the sexes in each species; and it requires but to contemplate the bearings of the argument, to see that ​the formation of different sexes may itself have been determined in the same way.
To convey here an adequate idea of Mr. Darwin's doctrine, throughout the immense range of its applications, is of course impossible. The few illustrations just given, are intended simply to remind the reader what Mr. Darwin's hypothesis is, and what are the else insoluble problems which it solves for us.
§ 166. But now, though it seems to me that we are thus supplied with a key to phenomena which are multitudinous and varied beyond all conception; it also seems to me that there is a moiety of the phenomena which this key will not unlock. Mr. Darwin himself recognizes use and disuse of parts, as causes of modifications in organisms; and does this, indeed, to a greater extent than do some who accept his general conclusion. But I conceive that he does not recognize them to a sufficient extent. While he shows that the inheritance of changes of structure caused by changes of function, is utterly insufficient to explain a great mass—probably the greater mass—of morphological phenomena; I think he leaves unconsidered a mass of morphological phenomena which are explicable as results of functionally-produced modifications, and are not explicable as results of natural selection.
By induction, as well as by inference from the hypothesis of natural selection, we know that there exists a balance among the powers of organs which habitually act together—such proportions among them that no one has any considerable excess of efficiency. We see, for example, that throughout the vascular system there is maintained an equilibrium of the component parts: in some cases, under continued excess of exertion, the heart gives way, and we have enlargement; in other cases the large arteries give way, and we have aneurisms; in other cases the minute blood-vessels give way—now bursting, now becoming chronically congested. ​That is to say, in the average constitution, no superfluous strength is possessed by any of the appliances for circulating the blood. Take, again, a set of motor organs. Great strain here causes the fibres of a muscle to tear. There the muscle does not yield but the tendon snaps. Elsewhere neither muscle nor tendon is damaged, but the bone breaks. Joining with these instances the general fact that, under the same adverse conditions, different individuals show their slight differences of constitution by going wrong some in one way and some in another; and that even in the same individual, similar adverse conditions will now affect one viscus and now another; it becomes manifest that though there cannot be maintained an accurate balance among the powers of the organs composing an organism, yet their excesses and deficiencies of power are extremely slight. That they must be extremely slight, is, as before said, a deduction from the hypothesis of natural selection. Mr. Darwin himself argues "that natural selection is continually trying to economise in every part of the organization. If under changed conditions of life a structure before useful becomes less useful, any diminution, however slight, in its development, will be seized on by natural selection, for it will profit the individual not to have its nutriment wasted in building up an useless structure." In other words, if any muscle has more fibres than are required, or if a bone is stronger than needful, no advantage results but rather a disadvantage—a disadvantage which will decrease the chances of survival. Hence it follows that among any organs which habitually act in concert, an increase of one can be of no service unless there is a concomitant increase of the rest. The co-operative parts must vary together; otherwise variation will be detrimental. A stronger muscle must have a stronger bone to resist its contractions; must have stronger correlated muscles and ligaments to secure the neighbouring articulations; must have larger blood-vessels to bring it supplies; must have a more massive nerve to transmit stimulus, and some extra ​development of a nervous centre to supply the extra stimulus. The question arises, then,—do variations of the appropriate kinds occur simultaneously in all these co-operative parts? Have we any reason to think that the parts spontaneously increase or decrease together? The assumption that they do seems to me untenable; and its untenability will, I think, become conspicuous if we take a case, and observe how extremely numerous and involved are the variations which must be supposed to occur together. In illustration of another point, we have already considered the modifications required to accompany increased weight of the head ( § 155). Instead of the bison, the moose deer, or the extinct Irish elk, will here best serve our purpose. In this last species the male has enormously-developed horns, used for purposes of offence and defence. These horns, weighing upwards of a hundred-weight, are carried at great mechanical disadvantage: supported as they are, along with the massive skull which bears them, at the extremity of the outstretched neck. Further, that these heavy horns may be of use in fighting, the supporting bones and muscles must be strong enough, not simply to carry them, but to put them in motion with the rapidity needed for giving blows. Let us, then, ask how, by natural selection, this complex apparatus of bones and muscles can have been developed, pari passu with the horns? If we suppose the horns to have been originally of like size with those borne by other kinds of deer; and if we suppose that in some individual they became larger by spontaneous variation; what would be the concomitant changes required to render their greater size useful? Other things equal, the blow given by a larger horn would be a blow given by a heavier mass moving at a smaller velocity: the momentum would be the same as before; and the area of contact with the body struck being somewhat increased, while the velocity was decreased, the injury done would be less. That horns may become better weapons, the whole apparatus concerned in moving them must be so ​strengthened as to impress more force on them, and to bear the more violent reactions of the blows given. The bones of the skull on which the horns are seated must be thickened; otherwise they will break. The vertebræ of the neck must be further developed; and unless the ligaments which hold together these vertebræ, and the muscles which move them, are also enlarged, nothing will be gained. Again the upper dorsal vertebræ and their spines must be strengthened, that they may withstand the stronger contractions of the neck-muscles; and like changes must be made on the scapular arch. Still more must there be required a simultaneous development of the bones and muscles of the fore-legs; since these extra growths in the horns, in the skull, in the neck, in the shoulders, add to the burden they have to bear; and without they are strengthened the creature will not only suffer from loss of speed but will fail in fight. Hence, to make larger horns of use, additional sizes must be acquired by numerous bones, muscles, and ligaments, as well as by the blood-vessels and nerves on which their actions depend. On calling to mind how the spraining of a single small muscle in the foot incapacitates for walking, or how permanent weakness in a knee-ligament will diminish the power of the leg, it will be seen that unless all these many changes are simultaneously made, they may as well be none of them made—or rather, they would better be none of them made; since the enlargements of some parts, by putting greater strains on connected parts, would render them relatively weaker if they remained unenlarged. Can we with any propriety assume that these many enlargements duly proportioned will be simultaneously effected by spontaneous variations? I think not. It would be a strong supposition that the vertebræ and muscles of the neck suddenly became bigger at the same time as the horns. It would be a still stronger supposition that the upper dorsal vertebræ not only at the same time became more massive, but appropriately altered their proportions, by the development of their immense neural ​spines. And it would be an assumption still more straining our powers of belief, that along with heavier horns there should spontaneously take place the required strengthenings in the bones, muscles, arteries, and nerves of the scapular and the fore-legs.
Besides the multiplicity of directly-coöperative organs, the multiplicity of organs which do not coöperate, save in the degree implied by their combination in the same organism, seems to me a further hindrance to the development of special structures by natural selection alone. Where the life is simple, or where circumstances render some one function supremely important, survival of the fittest may readily bring about the appropriate structural change, without aid from the transmission of functionally-caused modifications. But in proportion as the life grows complex—in proportion as a healthy existence cannot be secured by a large endowment of some one power, but demands many powers; in the same proportion do there arise obstacles to the increase of any particular power by "the preservation of favoured races in the struggle for life." As fast as the faculties are multiplied, so fast does it become possible for the several members of a species to have various kinds of superiorities over one another. While one saves its life by higher speed, another does the like by clearer vision, another by keener scent, another by quicker hearing, another by greater strength, another by unusual power of enduring cold or hunger, another by special sagacity, another by special timidity, another by special courage; and others by other bodily and mental attributes. Conditions being alike, each of these life-saving attributes is likely to be transmitted to posterity. But we may not assume that it will be increased in subsequent generations by natural selection. Increase of it can result only if individuals possessing average endowments of it are more frequently killed off than individuals highly endowed with it; and this can happen only when the attribute is one of greater importance, for the time being, than ​most of the other attributes. If those members of the species which have but ordinary shares of it, nevertheless survive by virtue of other superiorities which they severally possess; then it is not easy to see how this particular attribute can be developed by natural selection in subsequent generations. The probability seems rather to be that, by gamogenesis, this extra endowment will, on the average, be diminished in posterity—just serving in the long run to make up for the deficient endowments of those whose special powers lie in other directions; and so to keep up the normal structure of the species. As fast as the number of bodily and mental faculties increases, and as fast as maintenance of life comes to depend less on the amount of any one and more on the combined actions of all; so fast does the production of specialities of character by natural selection alone, become difficult. Particularly does this seem to be so with a species so multitudinous in its powers as mankind; and above all does it seem to be so with such of the human powers as have but minor shares in aiding the struggle for life—the æsthetic faculties, for example.
It by no means follows, however, that in cases of this kind, and cases of the preceding kind, natural selection plays no part. Wherever it is not the chief agent in working organic changes, it is still, very generally, a secondary agent. The survival of the fittest must nearly always further the production of modifications which produce fitness, whether they be incidental modifications, or modifications caused by direct adaptation. Evidently, those individuals whose constitutions have facilitated the production in them of any structural change consequent on any functional change demanded by some new external condition, will be the individuals most likely to live and to leave descendants. There must be a natural selection of functionally-acquired peculiarities, as well as of spontaneously-acquired peculiarities; and hence such structural changes in a species as result from changes of habit necessitated by changed circumstances, natural selection will render more rapid than they would otherwise be.
​There are, however, some modifications in the sizes and forms of parts, which cannot have been aided by natural selection; but which must have resulted wholly from the inheritance of functionally-caused alterations. The dwindling of organs of which the undue sizes entail no appreciable evils, furnishes the best evidence of this. Take, for an example, that diminution of the jaws and teeth which characterizes the civilized races, as contrasted with the savage races. How can the civilized races have been ​benefited in the struggle for life, by the slight decrease in these comparatively-small bones? No functional superiority possessed by a small jaw over a large jaw in civilized life, can be named as having caused the more frequent survival of small-jawed individuals. The only advantage accompanying smallness of jaw, is the advantage of economized nutrition; and this cannot be great enough to further the preservation of those distinguished by it. The decrease of weight in the jaw and co-operative parts, which has arisen in the course of thousands of years, does not amount to more than a few ounces. This decrease has to be divided among the many generations which have lived and died in the interval. Let us admit that the weight of these parts diminished to the extent of an ounce in a single generation (which is a large admission); it still cannot be contended that the having to carry an ounce less in weight, and to keep in repair an ounce less of tissue, could sensibly affect any man's fate. And if it never did this—nay, if it did not cause a frequent survival of small-jawed individuals where large-jawed individuals died; natural selection could neither cause nor aid diminution of the jaw and its appendages. Here, therefore, the decreased action which has accompanied the growth of civilized habits (the use of tools and the disuse of coarse food), must have been the sole cause at work. Through direct equilibration, diminished external stress on these parts has resulted in diminution of the internal forces by which this stress is met. From generation to generation, this lessening of the parts consequent on functional decline has been inherited. And since the survival of individuals must always have been determined by more important structural traits, this trait can have neither been facilitated nor retarded by natural selection.
§ 167. Returning from these extensive classes of facts for ​which Mr. Darwin's hypothesis does not account, to the still more extensive classes of facts for which it does account, and which are unaccountable on any other hypothesis; let us consider in what way this hypothesis is expressible in terms of the general doctrine of evolution. Already it has been pointed out that the evolving of modified types by "natural selection or the preservation of favoured races in the struggle for life," must be a process of equilibration; since it results in the production of organisms which are in equilibrium with their environments. At the outset of this chapter, something was done towards showing how this continual survival of the fittest may be understood as the progressive establishment of a balance between inner and outer forces. Here, however, we must consider the matter more closely.
On previous occasions we have contemplated the assemblage of individuals composing a species, as an aggregate in a state of moving equilibrium. We have seen that its powers of multiplication give it an expansive energy which is antagonized by other energies; and that through the rhythmical variations in these two sets of energies there is maintained an oscillating limit to its habitat, and an oscillating limit to its numbers. On another occasion ( § 96) it was shown that the aggregate of individuals constituting a species, has a kind of general life which, "like the life of an individual, is maintained by the unequal and ever-varying actions of incident forces on its different parts." We saw that "just as, in each organism, incident forces constantly produce divergences from the mean state in various directions, which are constantly balanced by opposite divergences indirectly produced by other incident forces; and just as the combination of rhythmical functions thus maintained, constitutes the life of the organism; so, in a species there is, through gamogenesis, a perpetual neutralization of those contrary deviations from the mean state, which are caused in its different parts by different sets of incident forces; and it is similarly by the rhythmical production and compensation of ​these contrary deviations that the species continues to live." Hence, to understand how a species is affected by causes which destroy some of its units and favour the multiplication of others, we must consider it as a whole whose parts are held together by complex forces that are ever re-balancing themselves—a whole whose moving equilibrium is continually disturbed and continually rectified. Thus much premised, let us next call to mind how moving equilibria in general are changed. In the first place, a new incident force falling on any part of an aggregate with balanced motions, produces a new motion in the direction of least resistance. In the second place, the new incident force is gradually used up in overcoming the opposing forces, and when it is all expended the opposing forces produce a recoil—a reverse deviation which counter-balances the original deviation. Consequently, to consider whether the moving equilibrium of a species is modified in the same way as moving equilibria in general, is to consider whether, when exposed to a new force, a species yields in the direction of least resistance; and whether, by its thus yielding, there is generated in the species a compensating change in the opposite direction. We shall find that it does both these things.
For what, expressed in mechanical terms, is the effect wrought on a species by some previously-unknown enemy, that kills such of its members as fail in defending themselves? The disappearance of those individuals which meet the destroying forces by the smallest preserving forces, is tantamount to the yielding of the species as a whole at the places where the resistances are the least. Or if by some general influence, such as alteration of climate, the members of a species are subject to increase of external actions which are ever tending to overthrow their equilibria, and which they are ever counter-balancing by certain physiological actions, which are the first to die? Those least able to generate the internal energies which antagonize these external energies. If the change be an increase of the ​winter's cold, then such members of the species as have unusual powers of getting food or of digesting food, or such as are by their constitutional aptitude for making fat, furnished with reserve stores of force, available in times of scarcity, or such as have the thickest coats and so lose least heat by radiation, survive; and their survival implies that in each of them the moving equilibrium of functions presents such an adjustment of internal forces, as prevents overthrow by the modified aggregate of external forces. Conversely, the members which die are, other things equal, those deficient in the power of meeting the new action by an equivalent counter-action. Thus, in all cases, a species considered as an aggregate in a state of moving equilibrium, has its state changed by the yielding of its fluctuating mass wherever this mass is weakest in relation to the special forces acting on it. The conclusion is, indeed, a truism. But now what must follow from the destruction of the least-resisting individuals and survival of the most-resisting individuals? On the moving equilibrium of the species as a whole, existing from generation to generation, the effect of this deviation from the mean state is to produce a compensating deviation. For if all such as are deficient of power in a certain direction are destroyed, what must be the effect on posterity? Had they lived and left offspring, the next generation would have had the same average powers as preceding generations: there would have been a like proportion of individuals less endowed with the needful power, and individuals more endowed with it. But the more-endowed individuals being alone left to continue the race, there must result a new generation characterized by a larger average endowment of this power. That is to say, on the moving equilibrium of a species, an action producing change in a given direction is followed, in the next generation, by a reaction producing an opposite change. Observe, too, that these effects correspond in their degrees of violence. If the alteration of some external factor is so ​great that it leaves alive only the few individuals possessing extreme endowments of the power required to antagonize it; then, in succeeding generations, there is a rapid multiplication of individuals similarly possessing extreme endowments of this power—the force impressed calls out an equivalent conflicting force. Moreover, the change is temporary where the cause is temporary, and permanent where the cause is permanent. All that are deficient in the needful attribute having been killed off, and the survivors having the needful attribute in a comparatively high degree, there will descend from them, not only some possessing equal amounts of this attribute with themselves, but also some possessing less amounts of it. If the destructive agency has not continued in action, such less-endowed individuals will multiply; and the species, after sundry oscillations, will return to its previous mean state. But if this agency be a persistent one, such less endowed individuals will be continually killed off, and eventually none but highly-endowed individuals will be produced—a new moving equilibrium, adapted to the new environing conditions, will result.
It may be objected that this mode of expressing the facts does not include the cases in which a species becomes modified in relation to surrounding agencies of a passive kind—cases like that of a plant which acquires hooked seed-vessels, by which it lays hold of the skins of passing animals, and makes them the distributors of its seeds—cases in which the outer agency has no direct tendency at first to affect the species, but in which the species so alters itself as to take advantage of the outer agency. To cases of this kind, however, the same mode of interpretation applies on simply changing the terms. While, in the aggregate of influences amid which a species exists, there are some which tend to overthrow the moving equilibria of its members, there are others which facilitate the maintenance of their moving equilibria, and some which are capable of giving their moving equilibria increased stability: instance the spread into their habitat of ​some new kind of prey, which is abundant at seasons when other prey is scarce. Now what is the process by which the moving equilibrium in any species becomes adapted to some additional external factor furthering its maintenance? Instead of an increased resistance to be met and counterbalanced, there is here a diminished resistance; and the diminished resistance is equilibrated in the same way as the increased resistance. As, in the one case, there is a more frequent survival of individuals whose peculiarities enable them to resist the new adverse factor; so, in the other case, there is a more frequent survival of individuals whose peculiarities enable them to take advantage of the new favourable factor. In each member of the species, the balance of functions and correlated arrangement of structures, differ slightly from those existing in other members. To say that among all its members, one is better fitted than the rest to benefit by some before-unused agency in the environment, is to say that its moving equilibrium is, in so far, more stably adjusted to the sum of surrounding influences. And if, consequently, this individual maintains its moving equilibrium when others fail, and has offspring which do the like—that is, if individuals thus characterized multiply and supplant the rest; there is, as before, a process which effects equilibration between the organism and its environment, not immediately but mediately, through the continuous intercourse between the species as a whole and the environment.
§ 168. Thus we see that indirect equilibration does whatever direct equilibration cannot do. All these processes by which organisms are re-fitted to their ever-changing environments, must be equilibrations of one kind or other. As authority for this conclusion, we have not simply the universal truth that change of every order is towards equilibrium; but we have also the truth that life itself is a moving equilibrium between inner and outer actions—a continuous adjustment of internal relations to external relations; ​or the maintenance of a balance between the forces to which an organism is subject and the forces which it evolves. Hence all changes which enable a species to live under altered conditions, are changes towards equilibrium with the altered conditions; and therefore those which do not come within the class of direct equilibrations, must come within the class of indirect equilibrations.
And now we reach an interpretation of Natural Selection regarded as a part of Evolution at large. As understood in First Principles, Evolution is a continuous redistribution of matter and motion; and a process of evolution which is not expressible in terms of matter and motion has not been reduced to its ultimate form. The conception of Natural Selection is manifestly one not known to physical science: its terms are not of a kind physical science can take cognisance of. But here we have found in what manner it may be brought within the realm of physical science. Rejecting metaphor we see that the process called Natural Selection is literally a survival of the fittest; and the outcome of the above argument is that survival of the fittest is a maintenance of the moving equilibrium of the functions in presence of outer actions: implying the possession of an equilibrium which is relatively stable in contrast with the unstable equilibria of those which do not survive.
* ↑ It will be seen that the argument naturally leads up to this expression—Survival of the Fittest—which was here used for the first time. Two years later (July, 1866) Mr. A. R. Wallace wrote to Mr. Darwin contending that it should be substituted for the expression "Natural Selection." Mr. Darwin demurred to this proposal. Among reasons for retaining his own expression he said that I had myself, in many cases, preferred it—"continually using the words Natural Selection." (Life and Letters, &c., vol. III, pp. 45-6.) Mr. Darwin was quite right in his statement, but not right in the motive he ascribed to me. My reason for frequently using the phrase "Natural Selection," after the date at which the phrase "Survival of the Fittest" was first used above, was that disuse of Mr. Darwin's phrase would have seemed like an endeavour to keep out of sight my own indebtedness to him, and the indebtedness of the world at large. The implied feeling has led me ever since to use the expressions Natural Selection and Survival of the Fittest with something like equal frequency.
* ↑ I am indebted to Mr. [now Sir W.] Flower for the opportunity of examining the many skulls in the Museum of the College of Surgeons for verification of this. Unfortunately the absence, in most cases, of some or many teeth, prevented me from arriving at that specific result which would have been given by weighing a number of the under jaws in each race. Simple inspection, however, disclosed a sufficiently-conspicuous difference. The under jaws of Australians and Negroes, when collated with those of Englishmen, were visibly larger, not only relatively but absolutely. One Australian jaw only seemed about of the same size as an average English jaw; and this (probably the jaw of a woman), belonging as it did to a smaller skull, bore a greater ratio to the whole body of which it formed part, than did an English jaw of the same actual size. In all the other cases, the under jaws of these inferior races (containing larger teeth than our own) were absolutely more massive than our own—often exceeding them in all dimensions; and relatively to their smaller skeletons were much more massive. Let me add that the Australian and Negro jaws are thus strongly contrasted, not with all British jaws, but only with the jaws of the civilized British. An ancient British skull in the collection possesses a jaw almost or quite as massive as those of the Australian skulls. All this is in harmony with the alleged relation between greater size of jaws and greater action of jaws, involved by the habits of savages. [In 1891 Mr. F. Howard Collins carefully investigated this matter: measuring ten Australian, ten Ancient British, and ten recent English skulls in the College of Surgeons Museum. The result proved an absolute difference of the kind above indicated, and a far greater relative difference. To ascertain this last a common standard of comparison was established—an equal size of skull in all the cases; and then when the relative masses or cubic sizes of the jaws were calculated, the result which came out was this:—Australian jaw, 1948; Ancient British jaw, 1135; Recent English jaw, 1030. "Hence," in the words of Mr. Collins, "the mass of the Recent English jaw is, roughly speaking, half that of the Australian relatively to that of the skull, and a ninth less than that of the Ancient British." He adds verifying evidence from witnesses who have no hypothesis to support—members of the Odontological Society. The Vice-President, Mr. Mummery, remarks of the Australians that "the jaw-bones are powerfully developed, and large in proportion to the cranium."]