# Popular Science Monthly/Volume 43/May 1893/The Inadequacy of Natural Selection II

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* THE INADEQUACY OF "NATURAL SELECTION."
By HERBERT SPENCER.
ALONG with that inadequacy of natural section to explain changes of structure which do not aid life in important ways, alleged in §166 of The Principles of Biology, a further inadequacy was alleged. It was contended that the relative powers of co-operative parts can not be adjusted solely by survival of the fittest; and especially where the parts are numerous and the cooperation complex. In illustration it was pointed out that immensely developed horns, such as those of the extinct Irish elk, weighing over a hundredweight, could not, with the massive skull bearing them, be carried at the extremity of the outstretched neck without many and great modifications of adjacent bones and muscles of the neck and thorax; and that without strengthening of the fore-legs, too, there would be failure alike in fighting and in locomotion. And it was argued that while we can not assume spontaneous increase of all these parts proportionate to the additional strains, we can not suppose them to increase by variation one at once, without supposing the creature to be disadvantaged by the weight and nutrition of parts that were for the time useless—parts, moreover, which would revert to their original sizes before the other needful variations occurred.
When, in reply to me, it was contended that co-operative parts vary together, I named facts conflicting with this assertion—the fact that the blind crabs of the Kentucky caves have lost their eyes but not the foot-stalks carrying them; the fact that the normal proportion between tongue and beak in certain selected varieties of pigeons is lost; the fact that lack of concomitance in ​increase of jaws and teeth in sundry kinds of pet dogs, has caused great crowding of the teeth (The Factors of Organic Evolution, pp. 12, 13). And I then argued that if co-operative parts, small in number and so closely associated as these are, do not vary together, it is unwarrantable to allege that co-operative parts which are very numerous and remote from one another vary together. After making this rejoinder I enforced my argument by a further example—that of the giraffe. Tacitly recognizing the truth that the unusual structure of this creature must have been, in its more conspicuous traits, the result of survival of the fittest (since it is absurd to suppose that efforts to reach a high branch could lengthen the legs), I illustrated afresh the obstacles to co-adaptation. Not dwelling on the objection that increase of any components of the fore-quarters out of adjustment to the others would cause evil rather than good, I went on to argue that the co-adaptation of parts required to make the giraffe's structure useful, is much greater than at first appears. This animal has a grotesque gallop, necessitated by the great difference in length between the fore and the hind limbs. I pointed out that the mode of action of the hind limbs shows that the bones and muscles have all been changed in their proportions and adjustments; and I contended that, difficult as it is to believe that all parts of the fore-quarters have been co-adapted by the appropriate variations now of this part, now of that, it becomes impossible to believe that all the parts in the hind-quarters have been simultaneously co-adapted to one another and to all the parts of the fore-quarters: adding that want of co-adaptation, even in a single muscle, would cause fatal results when high speed had to be maintained while escaping from an enemy.
Since this argument, repeated with this fresh illustration, was published in 1886, I have met with nothing to be called a reply; and might, I think, if convictions usually followed proofs, leave the matter as it stands. It is true that, in his Darwinism, Mr. Wallace has adverted to my renewed objection and, as already said, contended that changes such as those instanced can be effected by natural selection, since such changes can be effected by artificial selection: a contention which, as I have pointed out, assumes a parallelism that does not exist. But now, instead of pursuing the argument further along the same line, let me take a somewhat different line.
If there occurs some change in an organ, say, by increase of its size, which adapts it better to the creature's needs, it is admitted that when, as commonly happens, the use of the organ demands the co-operation of other organs, the change in it will generally be of no service unless the co-operative organs are changed. If, for instance, there takes place such a modification ​of a rodent's tail as that which, by successive increases, produces the trowel-shaped tail of the beaver, no advantage will be derived unless there also take place certain modifications in the bulks and shapes of the adjacent vertebrae and their attached muscles, as well, probably, as in the hind limbs, enabling them to withstand the reactions of the blows given by the tail. And the question is, by what process these many parts, changed in different degrees, are co-adapted to the new requirements—whether variation and natural selection alone can effect the readjustment. There are three conceivable ways in which the parts may simultaneously change: (1) they may all increase or decrease together in like degrees; (2) they may all simultaneously increase or decrease independently, so as not to maintain their previous proportions or assume any other special proportions; (3) they may vary in such ways and degrees as to make them jointly serviceable for the new end. Let us consider closely these several conceivabilities.
And first of all, what are we to understand by co-operative parts? In a general sense, all the organs of the body are co-operative parts, and are respectively liable to be more or less changed by change in any one. In a narrower sense, more directly relevant to the argument, we may, if we choose to multiply difficulties, take the entire framework of bones and muscles as formed of co-operative parts; for these are so related that any considerable change in the actions of some entails change in the actions of most others. It needs only to observe how, when putting out an effort, there goes, along with a deep breath, an expansion of the chest and a bracing up of the abdomen, to see that various muscles beyond those directly concerned are strained along with them. Or, when suffering from lumbago, an effort to lift a chair will cause an acute consciousness that not the arms only are brought into action, but also the muscles of the back. These cases show how the motor organs are so tied together that altered actions of some implicate others quite remote from them.
But without using the advantage which this interpretation of the words would give, let us take as co-operative organs those which are obviously such—the organs of locomotion. What, then, shall we say of the fore and hind limbs of terrestrial mammals, which co-operate closely and perpetually? Do they vary together? If so, how have there been produced such contrasted structures as that of the kangaroo, with its large hind limbs and small fore limbs, and that of the giraffe, in which the hind limbs are small and the fore limbs large—how does it happen that, descending from the same primitive mammal, these creatures have diverged in the proportions of their limbs in opposite directions? Take, again, the articulate animals. Compare one of the lower types, with its rows of almost equal-sized limbs, and one of the higher ​types, as a crab or a lobster, with limbs some very small and some very large. How came this contrast to arise in the course of evolution, if there was the equality of variation supposed?
But now let us narrow the meaning of the phrase still further; giving it a more favorable interpretation. Instead of considering separate limbs as co-operative, let us consider the component parts of the same limb as co-operative, and ask what would result from varying together. It would in that case happen that, though the fore and hind limbs of a mammal might become different in their sizes, they would not become different in their structures. If so, how have there arisen the unlikeness between the hind legs of the kangaroo and those of the elephant? Or if this comparison is objected to, because the creatures belong to the widely different divisions of implacental and placental mammals, take the cases of the rabbit and the elephant, both belonging to the last division. On the hypothesis of evolution these are both derived from the same original form, but the proportions of the parts have become so widely unlike that the corresponding joints are scarcely recognized as such by the unobservant: at what seem corresponding places the legs bend in opposite ways. Equally marked, or more marked, is the parallel fact among the Articulata. Take that limb of the lobster which bears the claw and compare it with the corresponding limb in an inferior articulate animal, or the corresponding limb of its near ally, the crayfish, and it becomes obvious that the component segments of the limb have come to bear to one another in the one case proportions immensely different from those they bear in the other case. Undeniably, then, on contemplating the general facts of organic structure, we see that the concomitant variations in the parts of limbs have not been of a kind to produce equal amounts of change in them, but quite the opposite—have been everywhere producing inequalities. Moreover, we are reminded that this production of inequalities among co-operative parts, is an essential principle of development. Had it not been so, there could not have been that progress from homogeneity of structure to heterogeneity of structure which constitutes evolution.
We pass now to the second supposition:—that the variations in co-operative parts occur irregularly, or in such independent ways that they bear no definite relations to one another—miscellaneously, let us say. This is the supposition which best corresponds with the facts. Glances at the faces around yield conspicuous proofs. Many of the muscles of the face and some of the bones, are distinctly co-operative; and these respectively vary in such ways as to produce in each person a different combination. What we see in the face we have reason to believe holds in the limbs as in all other parts. Indeed, it needs but to compare people whose arms are of the same lengths, and observe how stumpy are ​the fingers of one and how slender those of another; or it needs but to note the unlikeness of gait of passers-by, implying small unlikenesses of structure; to be convinced that the relations among the variations of co-operative parts are anything but fixed. And now, confining our attention to limbs, let us consider what must happen if, by variations taking place miscellaneously, limbs have to be partially changed from fitness for one function to fitness for another function—have to be re-adapted. That the reader may fully comprehend the argument, he must here have patience while a good many anatomical details are set down.
Let us suppose a species of quadruped of which the members have for long past periods been accustomed to locomotion over a relatively even surface, as, for instance, the "prairie dogs" of North America; and let us suppose that increase of numbers has driven part of them into a region full of obstacles to easy locomotion—covered, say, by the decaying stems of fallen trees, such as one sees in portions of primeval forest. Ability to leap must become a useful trait; and, according to the hypothesis we are considering, this ability will be produced by the selection of favorable variations. What are the variations required? A leap is effected chiefly by the bending of the hind limbs so as to make sharp angles at the joints, and then suddenly straightening them; as any one may see on watching a cat leap on to the table. The first required change, then, is increase of the large extensor muscles, by which the hind limbs are straightened. Their increases must be duly proportioned, for if those which straighten one joint become much stronger than those which straighten the other joint, the result must be collapse of the other joint when the muscles are contracted together. But let us make a large admission, and suppose these muscles to vary together; what further muscular change is next required? In a plantigrade mammal the metatarsal bones chiefly bear the reaction of the leap, though the toes may have a share. In a digitigrade mammal, however, the toes form almost exclusively the fulcrum, and if they are to bear the reaction of a higher leap, the flexor muscles which depress and bend them must be proportionately enlarged; if not, the leap will fail from want of a firm point d'appui. Tendons as well as muscles must be modified; and, among others, the many tendons which go to the digits and their phalanges. Stronger muscles and tendons imply greater strains on the joints; and unless these are strengthened, one or other dislocation will be caused by a more powerful spring. Not only the articulations themselves must be so modified as to bear greater stress, but also the numerous ligaments which hold the parts of each in place. Nor can the bodies of the bones remain unstrengthened; for if they have no more than the strengths needed for previous ​movements they will fail to bear more violent movements. Thus, saying nothing of the required changes in the pelvis as well as in the nerves and blood-vessels, there are, counting bones, muscles, tendons, ligaments, at least fifty different parts in each hind leg which have to be enlarged. Moreover, they have to be enlarged in unlike degrees. The muscles and tendons of the outer toes, for example, need not be added to so much as those of the median toes. Now, throughout their successive stages of growth, all these parts have to be kept fairly well balanced; as any one may infer on remembering sundry of the accidents he has known. Among my own friends I could name one who, when playing lawn-tennis, snapped the Achilles tendon; another who, while swinging his children, tore some of the muscular fibers in the calf of his leg; another who, in getting over a fence, tore a ligament of one knee. Such facts, joined with every one's experience of sprains, show that during the extreme exertions to which limbs are now and then subject, there is a giving way of parts not quite up to the required level of strength. How, then, is this balance to be maintained? Suppose the extensor muscles have all varied appropriately; their variations are useless unless the other co-operative parts have also varied appropriately. Worse than this. Saying nothing of the disadvantage caused by extra weight and cost of nutrition, they will be causes of mischief—causes of derangement to the rest by contracting with undue force. And then, how long will it take for the rest to be brought into adjustment? As Mr. Darwin says concerning domestic animals: "Any particular variation would generally be lost by crossing, reversions etc., . . . unless carefully preserved by man." In a state of nature, then, favorable variations of these muscles would disappear again long before one or a few of the co-operative parts could be appropriately varied, much more before all of them could.
With this insurmountable difficulty goes a difficulty still more insurmountable—if the expression may be allowed. It is not a question of increased sizes of parts only, but of altered shapes of parts, too. A glance at the skeletons of mammals shows how unlike are the forms of the corresponding bones of their limbs; and shows that they have been severally remolded in each species to the different requirements entailed by its different habits. The change from the structures of hind limbs fitted only for walking and trotting to hind limbs fitted also for leaping, implies, therefore, that along with strengthenings of bones there must go alterations in their forms. Now the spontaneous alterations of form which may take place in any bone are countless. How long, then, will it be before there takes place that particular alteration which will make the bone fitter for its new action? And what is the ​probability that the many required changes of shape, as well as of size, in bones will each of them be effected before all the others are lost again? If the probabilities against success are incalculable, when we take account only of changes in the size of parts, what shall we say of their incalculableness when differences of form also are taken into account?
"Surely this piling up of difficulties has gone far enough"; the reader will be inclined to say. By no means. There is a difficulty immeasurably transcending those named. We have thus far omitted the second half of the leap, and the provisions to be made for it. After ascent of the animal's body comes descent; and the greater the force with which it is projected up, the greater is the force with which it comes down. Hence, if the supposed creature has undergone such changes in the hind limbs as will enable them to propel it to a greater height, without having undergone any changes in the fore limbs, the result will be that on its descent the fore limbs will give way, and it will come down on its nose. The fore limbs, then, have to be changed simultaneously with the hind. How changed? Contrast the markedly bent hind limbs of a cat with its almost straight fore limbs, or contrast the silence of the upward spring on to the table with the thud which the fore paws make as it jumps off the table. See how unlike the actions of the hind and fore limbs are, and how unlike their structures. In what way, then, is the required co-adaptation to be effected? Even were it a question of relative sizes only, there would be no answer; for facts already given show that we may not assume simultaneous increases of size to take place in the hind and fore limbs; and, indeed, a glance at the various human races, which differ considerably in the ratios of their legs to their arms, shows us this. But it is not simply a question of sizes. To bear the increased shock of descent the fore limbs must be changed throughout in their structures. Like those in the hind limb, the changes must be of many parts in many proportions; and they must be both in sizes and in shapes. More than this. The scapular arch and its attached muscles must also be strengthened and remolded. See, then, the total requirements. We must suppose that by natural selection of miscellaneous variations, the parts of the hind limbs shall be co-adapted to one another, in sizes, shapes, and ratios; that those of the fore limbs shall undergo co-adaptations similar in their complexity, but dissimilar in their kinds; and that the two sets of co-adaptations shall be effected pari passu. If, as may be held, the probabilities are millions to one against the first set of changes being achieved, then it may be held that the probabilities are billions to one against the second being simultaneously achieved, in progressive adjustment to the first.
​There remains only to notice the third conceivable mode of adjustment. It may be imagined that though, by the natural selection of miscellaneous variations, these adjustments can not be effected, they may nevertheless be made to take place appropriately. How made? To suppose them so made is to suppose that the prescribed end is somewhere recognized; and that the changes are step by step simultaneously proportioned for achieving it—is to suppose a designed production of these changes. In such case, then, we have to fall back in part upon the primitive hypothesis; and if we do this in part, we may as well do it wholly—may as well avowedly return to the doctrine of special creation.
What, then, is the only defensible interpretation? If such modifications of structure produced by modifications of function as we see take place in each individual, are in any measure transmissible to descendants, then all these co-adaptations, from the simplest up to the most complex, are accounted for. In some cases this inheritance of acquired characters suffices by itself to explain the facts; and in other cases it suffices when taken in combination with the selection of favorable variations. An example of the first class is furnished by the change just considered; and an example of the second class is furnished by the case before named of development in a deer's horns. If, by some extra massiveness spontaneously arising, or by formation of an additional "point," an advantage is gained either for attack or defense, then, if the increased muscularity and strengthened structure of the neck and thorax, which wielding of these somewhat heavier horns produces, are in a greater or less degree inherited, and in several successive generations, are by this process brought up to the required extra strength, it becomes possible and advantageous for a further increase of the horns to take place, and a further increase in the apparatus for wielding them, and so on continuously. By such processes only, in which each part gains strength in proportion to function, can co-operative parts be kept in adjustment, and be readjusted to meet new requirements. Close contemplation of the facts impresses me more strongly than ever with the two alternatives—either there has been inheritance of acquired characters, or there has been no evolution.—Contemporary Review.
[To be concluded.]