These are, to some extent, an argument against the cumulative inheritance of such effects. When a nerve atrophies from disuse, or a duct shrivels, or bone is absorbed, or a muscle becomes small or flabby, it proves, so far, that the average effect of use through enormous ages is not transmitted. When the fibula of a dog's leg thickens by 400 per cent. to a size "equal to or greater than" that of the removed tibia which previously did the work,[69] it shows that in spite of disuse for countless generations, the "almost filiform" bone has retained a potentiality of development which is fully equal to that possessed by the larger one which has been constantly used. When, after being reared on the ailanthus, the caterpillars of the Bombyx hesperus die of hunger rather than return to their natural food, the inherited effect of ancestral habit does not seem to be particularly strong. Neither is there any strongly-inherited effect of long-continued ancestral wildness in many animals which are easily tamed.
WOULD NATURAL SELECTION FAVOUR USE-INHERITANCE?
If use-inheritance is really one of the factors of evolution, it is certainly a subordinate one, and an utterly helpless one, whenever it comes into conflict with the great ruling principle of Selection. Would this dominant cause of evolution have favoured a tendency to use-inheritance if such had appeared, or would it have discouraged and destroyed it? We have already seen that use-inheritance is unnecessary, since natural selection will be far more effective in bringing about advantageous modifications; and if it can be shown that use-inheritance would often be an evil, it then becomes probable that on the whole natural selection would more strongly discourage and eliminate it as a hostile factor than it might occasionally favour such a tendency as a totally unnecessary aid.
USE-INHERITANCE AN EVIL.
Use-inheritance would crudely and indiscriminately proportion parts to actual work done—or rather to the varying nourishment and growth resulting from a multiplicity of causes—and this in its various details would often conflict most seriously with the real necessities of the case, such as occasional passive strength, or appropriate shape, lightness and general adaptation. If its accumulated effects were not corrected by natural or sexual selection, horns and antlers would disappear in favour of enlarged hoofs. The elephant's tusks would become smaller than its teeth. Men would have callosities for sitting on, like certain monkeys, and huge corns or hoofs for walking on. Bones would often be modified disastrously. Thus the condyle of the human jaw would become larger than the body of the jaw, because as the fulcrum of the lever it receives more pressure. Some organs (like the heart, which is always at work) would become inconveniently or unnecessarily large. Other absolutely indispensable organs, which are comparatively passive or are very seldom used, would dwindle until their weakness caused the ruin of the individual or the extinction of the species. In eliminating various evil results of use-inheritance, natural selection would be eliminating use-inheritance itself. The displacement of Lamarck's theory by Darwin's shows that the effects of use-inheritance often differ from those required by natural selection; and it is clear that the latter factor must at least have reduced use-inheritance to the very minor position of comparative feebleness and harmlessness assigned to it by Darwin.
Use-inheritance would be ruinous through causing unequal variation in co-operative parts—of which Mr. Spencer may accept his own instances of the jaws and teeth, and the cave-crab's lost eyes and persistent eye-stalks, as typical examples. That the variation would be unequal seems almost self-evident from the varying rapidity and extent of the effects of use and disuse on different tissues and on different parts of the general structure. The optic nerve may atrophy in a few months from disuse consequent on the loss of the eye. Some of the bones of the rudimentary hind legs of the whale are still in existence after disuse for an enormous period. Evidently use-inheritance could not equally modify the turtle and its shell, or the brain and its skull; and in minor matters there would be the same incongruity of effect. Thus, if the molar teeth lengthened from extra use the incisors could not meet. Unequal and indiscriminate variation would throw the machinery of the organism out of gear in innumerable ways.
Use-inheritance would perpetuate various evils. We are taught, for instance, that it perpetuates short-sight, inferior senses, epilepsy, insanity, nervous disorders, and so forth. It would apparently transmit the evil effects of over-exertion, disuse, hardship, exposure, disease and accident, as well as the defects of age or immaturity.
Would it not be better on the whole if each individual took a fresh start as far as possible on the advantageous typical lines laid down by natural selection? Through the long stages of evolution from primæval protoplasm upwards, such species as were least affected by use-inheritance would be most free to develop necessary but seldom-used organs, protective coverings such as shells or skulls, and natural weapons, defences, ornaments, special adaptations, and so forth; and this would be an advantage—for survival would obviously depend on the importance of a structure or faculty in deciding the struggle for existence and reproduction, and not on the total amount of its using or nourishment. If natural selection had on the whole favoured this officious ally and frequent enemy, surely we should find better evidence of its existence.
Without laying undue stress upon the evil effects of use-inheritance, a careful examination of them in detail may at least serve to counter-balance the optimistic a priori arguments for belief in that plausible but unproven factor of evolution.
The benefits derivable from use-inheritance are largely illusory. The effects of use, indeed, are generally beneficial up to a certain point; for natural selection has sanctioned or evolved organs which possess the property or potentiality of developing to the right extent under the stimulus of use or nourishment. But use-inheritance would cumulatively alter this individual adaptability, and would tend to fix the size of organs by the average amount of ancestral use or disuse rather than by the actual requirements of the individual. Of course under changed conditions involving increased or lessened use of parts it might become advantageous; but even here it may prove a decided hindrance to adaptive evolution in some respects as well as an unnecessary aid in others. Thus in the case of animals becoming heavier, or walking more, it would lengthen the legs although natural selection might require them to be shortened. In the Aylesbury duck and the Call duck, if use-inheritance has increased the dimensions of the bones and tendons of the leg, natural selection has had to counteract this increase so far as length is concerned, and to effect 8 per cent. of shortening besides. If use-inheritance thickens bones without proportionally lengthening them, it would hinder rather than help the evolution of such structures as the long light wings of birds, or the long legs and neck of the giraffe or crane.