The cases Mr. Spencer adduces as demonstrating the effects of disuse in diminishing the size and strength of organs are, the diminished size of the jaws in the races of civilised men, and the diminution of the muscles used in closing the jaws in the case of pet-dogs fed for generations on soft food. He argues that the minute reduction in any one generation could not possibly have been useful, and, therefore, not the subject of natural selection; and against the theory of correlation of the diminished jaw with increased brain in man, he urges that there are cases of large brain development, accompanied by jaws above the average size. Against the theory of economy of nutrition in the case of the pet-dogs, he places the abundant food of these animals which would render such economy needless.
But neither he nor Mr. Darwin has considered the effects of the withdrawal of the action of natural selection in keeping up the parts in question to their full dimensions, which, of itself, seems to me quite adequate to produce the results observed. Recurring to the evidence, adduced in Chapter III, of the constant variation occurring in all parts of the organism, while selection is constantly acting on these variations in eliminating all that fall below the best working standard, and preserving only those that are fully up to it; and, remembering further, that, of the whole number of the increase produced annually, only a small percentage of the best adapted can be preserved, we shall see that every useful organ will be kept up nearly to its higher limit of size and efficiency. Now Mr. Galton has proved experimentally that, when any part has thus been increased (or diminished) by selection, there is in the offspring a strong tendency to revert to a mean or average size, which tends to check further increase. And this mean appears to be, not the mean of the actual existing individuals but a lower mean, or that from which they had been recently raised by selection.[199] He calls this the law of "Regression towards Mediocrity," and it has been proved by experiments with vegetables and by observations on mankind. This regression, in every generation, takes place even when both parents have been selected for their high development of the organ in question; but when there is no such selection, and crosses are allowed among individuals of every grade of development, the deterioration will be very rapid; and after a time not only will the average size of the part be greatly reduced, but the instances of full development will become very rare. Thus what Weismann terms "panmixia," or free intercrossing, will co-operate with Galton's law of "regression towards mediocrity," and the result will be that, whenever selection ceases to act on any part or organ which has heretofore been kept up to a maximum of size and efficiency, the organ in question will rapidly decrease till it reaches a mean value considerably below the mean of the progeny that has usually been produced each year, and very greatly below the mean of that portion which has survived annually; and this will take place by the general law of heredity, and quite irrespective of any use or disuse of the part in question. Now, no observations have been adduced by Mr. Spencer or others, showing that the average amount of change supposed to be due to disuse is greater than that due to the law of regression towards mediocrity; while even if it were somewhat greater, we can see many possible contributory causes to its production. In the case of civilised man's diminished jaw, there may well be some correlation between the jaw and the brain, seeing that increased mental activity would lead to the withdrawal of blood and of nervous energy from adjacent parts, and might thus lead to diminished growth of those parts in the individual. And in the case of pet-dogs, the selection of small or short-headed individuals would imply the unconscious selection of those with less massive temporal muscles, and thus lead to the concomitant reduction of those muscles. The amount of reduction observed by Darwin in the wing-bones of domestic ducks and poultry, and in the hind legs of tame rabbits, is very small, and is certainly no greater than the above causes will well account for; while so many of the external characters of all our domestic animals have been subject to long-continued artificial selection, and we are so ignorant of the possible correlations of different parts, that the phenomena presented by them seem sufficiently explained without recurrence to the assumption that any changes in the individual, due to disuse, are inherited by the offspring.
Supposed Effects of Disuse among Wild Animals.
It may be urged, however, that among wild animals we have many undoubted results of disuse much more pronounced than those among domestic kinds, results which cannot be explained by the causes already adduced. Such are the reduced size of the wings of many birds on oceanic islands; the abortion of the eyes in many cave animals, and in some which live underground; and the loss of the hind limbs in whales and in some lizards. These cases differ greatly in the amount of the reduction of parts which has taken place, and may be due to different causes. It is remarkable that in some of the birds of oceanic islands the reduction is little if at all greater than in domestic birds, as in the water-hen of Tristan d'Acunha. Now if the reduction of wing were due to the hereditary effects of disuse, we should expect a very much greater effect in a bird inhabiting an oceanic island than in a domestic bird, where the disuse has been in action for an indefinitely shorter period. In the case of many other birds, however—as some of the New Zealand rails and the extinct dodo of Mauritius—the wings have been reduced to a much more rudimentary condition, though it is still obvious that they were once organs of flight; and in these cases we certainly require some other causes than those which have reduced the wings of our domestic fowls. One such cause may have been of the same nature as that which has been so efficient in reducing the wings of the insects of oceanic islands—the destruction of those which, during the occasional use of their wings, were carried out to sea. This form of natural selection may well have acted in the case of birds whose powers of flight were already somewhat reduced, and to whom, there being no enemies to escape from, their use was only a source of danger. We may thus, perhaps, account for the fact that many of these birds retain small but useless wings with which they never fly; for, the wings having been reduced to this functionless condition, no power could reduce them further except correlation of growth or economy of nutrition, causes which only rarely come into play.
The complete loss of eyes in some cave animals may, perhaps, be explained in a somewhat similar way. Whenever, owing to the total darkness, they became useless, they might also become injurious, on account of their delicacy of organisation and liability to accidents and disease; in which case natural selection would begin to act to reduce, and finally abort them; and this explains why, in some cases, the rudimentary eye remains, although completely covered by a protective outer skin. Whales, like moas and cassowaries, carry us back to a remote past, of whose conditions we know too little for safe speculation. We are quite ignorant of the ancestral forms of either of these groups, and are therefore without the materials needful for determining the steps by which the change took place, or the causes which brought it about.[200]
On a review of the various examples that have been given by Mr. Darwin and others of organs that have been reduced or aborted, there seems too much diversity in the results for all to be due to so direct and uniform a cause as the individual effects of disuse accumulated by heredity. For if that were the only or chief efficient cause, and a cause capable of producing a decided effect during the comparatively short period of the existence of animals in a state of domestication, we should expect to find that, in wild species, all unused parts or organs had been reduced to the smallest rudiments, or had wholly disappeared. Instead of this we find various grades of reduction, indicating the probable result of several distinct causes, sometimes acting separately, sometimes in combination, such as those we have already pointed out.
And if we find no positive evidence of disuse, acting by its direct effect on the individual, being transmitted to the offspring, still less can we find such evidence in the case of the use of organs. For here the very fact of use, in a wild state, implies utility, and utility is the constant subject for the action of natural selection; while among domestic animals those parts which are exceptionally used are so used in the service of man, and have thus become the subjects of artificial selection. Thus "the great and inherited development of the udders in cows and goats," quoted by Spencer from Darwin, really affords no proof of inheritance of the increase due to use, because, from the earliest period of the domestication of these animals, abundant milk-production has been highly esteemed, and has thus been the subject of selection; while there are no cases among wild animals that may not be better explained by variation and natural selection.
Difficulty as to Co-adaptation of Parts by Variation and Selection.
Mr. Spencer again brings forward this difficulty, as he did in his Principles of Biology twenty-five years ago, and urges that all the adjustments of bones, muscles, blood-vessels, and nerves which would be required during, for example, the development of the neck and fore-limbs of the giraffe, could not have been effected by "simultaneous fortunate spontaneous variations." But this difficulty is fully disposed of by the facts of simultaneous variation adduced in our third chapter, and has also been specially considered in Chapter VI, p. 127. The best answer to this objection may, perhaps, be found in the fact that the very thing said to be impossible by variation and natural selection has been again and again effected by variation and artificial selection. During the process of formation of such breeds as the greyhound or the bulldog, of the race-horse and carthorse, of the fantail pigeon or the otter-sheep, many co-ordinate adjustments have been produced; and no difficulty has occurred, whether the change has been effected by a single variation—as in the last case named—or by slow steps, as in all the others. It seems to be forgotten that most animals have such a surplus of vitality and strength for all the ordinary occasions of life that any slight superiority in one part can be at once utilised; while the moment any want of balance occurs, variations in the insufficiently developed parts will be selected to bring back the harmony of the whole organisation. The fact that, in all domestic animals, variations do occur, rendering them swifter or stronger, larger or smaller, stouter or slenderer, and that such variations can be separately selected and accumulated for man's purposes, is sufficient to render it certain that similar or even greater changes may be effected by natural selection, which, as Darwin well remarks, "acts on every internal organ, on every shade of constitututional difference, on the whole machinery of life." The difficulty as to co-adaptation of parts by variation and natural selection appears to me, therefore, to be a wholly imaginary difficulty which has no place whatever in the operations of nature.
Direct Action of the Environment.