Chapter 3

[11] _Origin of Species_, pp. 230-232; Bates's _Naturalist on the Amazons_. Darwin is "surprised that no one has hitherto advanced the demonstrative case of neuter insects, against the well-known doctrine of inherited habit, as advanced by Lamarck." As he justly observes, "it proves that with animals, as with plants, any amount of modification may be effected by the accumulation of numerous, slight, spontaneous variations, which are in any way profitable, without exercise or habit having been brought into play. For peculiar habits confined to the workers or sterile females, however long they might be followed, could not possibly affect the males and fertile females, which alone leave any descendants." Some slight modification of these remarks, however, may possibly be needed to meet the case of "factitious queens," who (probably through eating particles of the royal food) become capable of producing a few male eggs.

[12] _Descent of Man_, pp. 573, 572, and footnote.

[13] _Contemporary Review_, December, 1875, p. 92.

[14] See _Origin of Species_, pp. 5-8. "Changed conditions induce an almost indefinite amount of fluctuating variability, by which the whole organization is rendered in some degree plastic" (_Descent of Man_, p. 30). It also appears that "the nature of the conditions is of subordinate importance in comparison with the nature of the organism in determining each particular form of variation;--perhaps of not more importance than the nature of the spark, by which a mass of combustible matter is ignited, has in determining the nature of the flames" (_Origin of Species_, p. 8).

DARWIN'S EXAMPLES.

The most formidable cases brought forward by Mr. Spencer are from Darwin. I shall endeavour to show, however, that Darwin was probably wrong in retaining the older explanation of these facts, and that the remains of the Lamarckian theory of use-inheritance need not any longer encumber the great explanation which has superseded that fallacious and unproven theory and has rendered it totally unnecessary. Meanwhile I think it is an excellent sign that Mr. Spencer has to complain that "Nowadays most naturalists are more Darwinian than Mr. Darwin himself"--inasmuch as they are inclined to say that there is "no proof" that the effects of use and disuse are inherited. Other excellent signs are the recent issue of a translation of Weismann's important essays on this and kindred subjects,[15] the strong support given to his views by Wallace in his _Darwinism_, and their adoption by Ray Lankester in his article on Zoology in the latest edition of the _Encyclopaedia Britannica_. So sound and cautious an investigator as Francis Galton had also in 1875 concluded that "acquired modifications are barely, if at all, _inherited_, in the correct sense of that word."

Darwin's belief in the inheritance of acquired characters was more or less hereditary in the family. His grandfather, Erasmus Darwin, anticipated Lamarck's views in his _Zoonomia_, which Darwin at one time "greatly admired." His father was "convinced" of the "inherited evil effects of alcohol," and to this extent at least he strongly impressed the belief in the inheritance of acquired characters upon his children's minds.[16] Darwin must also have been imbued with Lamarckian ideas from other sources, although Dr. Grant's enthusiastic advocacy entirely failed to convert him to a belief in evolution.[17] "Nevertheless," he says, "it is probable that the hearing rather early in life such views maintained and praised may have favoured my upholding them under a different form in my _Origin of Species_"--a remark which refers to Lamarck's views on the general doctrine of evolution, but might also prove equally true if applied to Darwin's partial retention of the Lamarckian explanation of that evolution. Professor Huxley has pointed out that in Darwin's earlier sketch of his theory of evolution (1844) he attached more weight to the inheritance of acquired habits than he does in his _Origin of Species_ published fifteen years later.[18] He appears to have acquired the belief in early life without first questioning and rigorously testing it as he would have done had it originated with himself. In later life it appeared to assist his theory of evolution in minor points, and in particular it appeared absolutely indispensable to him as the _only_ explanation of the diminution of disused parts in cases where, as in domestic animals, economy of growth seemed to be practically powerless. He failed to adequately notice the effect of panmixia, or the withdrawal of selection, in causing or allowing degeneracy and dwindling under disuse; and he hardly attached sufficient importance to the fact that rudimentary organs and other supposed effects of use or disuse are quite as marked features in neuter insects which cannot transmit the effects of use and disuse as they are in the higher animals.

REDUCED WINGS OF BIRDS OF OCEANIC ISLANDS.

Darwin himself has pointed out that the rudimentary wings of island beetles, at first thought to be due to disuse, are mainly brought about by natural selection--the best-winged beetles being most liable to be blown out to sea. But he says that in birds of the oceanic islands "not persecuted by any enemies, the reduction of their wings has probably been caused by disuse." This explanation may be as fallacious as it is acknowledged to have been in the case of the island beetles. According to Darwin's own views, natural selection _must_ at least have played an important part in reducing the wings; for he holds that "natural selection is continually trying to economize every part of the organization." He says: "If under changed conditions of life a structure, before useful, becomes less useful, its diminution will be favoured, for it will profit the individual not to have its nutriment wasted in building up an useless structure.... Thus, as I believe, natural selection will tend in the long run to reduce any part of the organization, as soon as it becomes, through changed habits, superfluous."[19] If, as Darwin powerfully urges (and he here ignores his usual explanation), ostriches' wings are insufficient for flight in consequence of the economy enforced by natural selection,[20] why may not the reduced wings of the dodo, or the penguin, or the apteryx, or of the Cursores generally, be wholly attributed to natural selection in favour of economy of material and adaptation of parts to changed conditions? The great principle of economy is continually at work shaping organisms, as sculptors shape statues, by removing the superfluous parts; and a mere glance at the forms of animals in general will show that it is well-nigh as dominant and universal a principle as is that of the positive development of useful parts. Other causes, moreover besides actual economy, would favour shorter and more convenient wings on oceanic islands. In the first place, birds that were somewhat weak on the wing would be most likely to settle on an island and stay there. Shortened wings would then become advantageous because they would restrain fatal migratory tendencies or useless and perilous flights in which the birds that flew furthest would be most often carried away by storms and adverse winds. Reduced wings would keep the birds near the shelter and the food afforded by the island and its neighbourhood, and in some cases would become adapted to act as fins or flappers for swimming under water in pursuit of fish.

The reduced size of the wings of these island birds is paralleled by the remarkable thinness, &c., of the shell of the "gigantic land-tortoise" of the Galapagos Islands. The changes seen in the carapace can hardly have been brought about by the inherited effects of special disuse. Why then should not the reduction of equally useless, more wasteful, and perhaps positively dangerous wings be also due to an economy which has become advantageous to bird and reptile alike through the absence of the mammalian rivals whose places they are evidently being modified to fill? The _complete_ loss of the wings in neuter ants and termites can scarcely be due to the inherited effects of disuse; and as natural selection has abolished these wings in spite of the opposition of use-inheritance, it must clearly be fully competent to reduce wings without its aid. In considering the rudimentary wings of the apteryx, or of the moa, emu, ostrich, &c., we must not forget the frequent or occasional occurrence of hard seasons, and times of drought and famine, when Nature eliminates redundant, wasteful, and ill-adapted organisms in so severe and wholesale a fashion. Where enemies are absent there would be unrestrained multiplication, and this would greatly increase the severity of the competition for food, and so hasten the elimination of disused and useless parts.

DROOPING EARS AND DETERIORATED INSTINCTS.

Mr. Galton has pointed out that existing races and existing organs are only kept at their present high pitch of organic excellence by the stringent and incessant action of natural or artificial selection; and the simple relaxation or withdrawal of such selective influences will almost necessarily result in a certain amount of deterioration, independently even of the principle of economy.[21] I think that this cessation of a previous selective process will account for the drooping--but _not diminished_--ears of various domesticated animals (human preference and increased weight evidently aiding), and also for the inferior instincts seen in them and in artificially-fed caterpillars of the silk-moth, which now "often commit the strange mistake of devouring the base of the leaf on which they are feeding, and consequently fall down." Anyhow, I fail to see that anything is proved by this latter case, except that natural instinct may be perverted or aborted under unnatural conditions and a changed method of selection which abolishes the powerful corrective formerly supplied by natural selection.

WINGS AND LEGS OF DUCKS AND FOWLS.

The reduced wings and enlarged legs of domesticated ducks and fowls are attributed by Darwin and Spencer to the inheritance of the effects of use and disuse. But the inference by no means follows. Natural selection would usually favour these adaptive changes, and they would also have been aided by an artificial selection which is often unconscious or indirect. Birds with diminished power of flight would be less difficult to keep and manage, and in preserving and multiplying such birds man would be unconsciously bringing about structural changes which would easily be regarded as effects of use and disuse. "About eighteen centuries ago Columella and Varro speak of the necessity of keeping ducks in netted enclosures like other wild fowl, so that at this period there was danger of their flying away."[22] Is it not probable that the best fliers would escape most frequently, or would pine most if kept confined? On the other hand, birds with lessened powers of flight would not be eliminated as under natural conditions, but would be favoured; and natural selection, together with artificial selection of the most flourishing birds, would thicken and strengthen the legs to meet increased demands upon them.

The diminution of the duck's wing is not great even in the birds that "never fly," and from this we must deduct the direct effect of disuse on the individual during its lifetime. As Weismann suggests, the _inherited_ portion of the change could only be ascertained by comparing the bones, &c., of wild and tame ducks _similarly reared_. If individual disuse diminished the weight of the duck's wing-bones by 9 per cent. there would be nothing left to account for.

I suspect that investigation would reveal anomalies inconsistent with the theory of use-inheritance. Thus according to Darwin's tables of comparative weights and measurements[23] the leg-bones of the Penguin duck have slightly diminished in length, although they have increased 39 per cent. in weight. Relatively to the weight of the skeleton, the leg-bones have shortened in the tame breeds of ducks by over 5 per cent. (and in two breeds by over 8 per cent.) although they have increased more than 28 per cent. in proportional weight.[24] How can increased use simultaneously shorten and thicken these bones? If the relative shortening is attributed to a heavier skeleton, then the apparently reduced weight of the wing-bones is fully accounted for by the same circumstance, and disuse has had no inherited effect.

Another strange circumstance is that the wing-bones have diminished _in length only_. The shortening is about 6 per cent. more than in the shortened legs, and it amounts to 11 per cent. as compared with the weight of the skeleton. Such a shortening should represent a reduction of 29 per cent. in weight, whereas the actual reduction in the weight of the wing-bones relatively to the weight of the skeleton is only 9 per cent. even in the breeds that never fly. Independently of shortening, the disused wing-bones have actually thickened or increased in weight. In the Aylesbury duck the disproportion caused by these conflicting changes is so great that the wing-bones are 47 per cent. heavier than they should be if their weight had varied proportionally with their length.[25] The reduction in weight on which Darwin relies seems to be entirely due to the shortening, and this shortening appears to be irrelevant to disuse, since the wings of the Call duck are similarly shortened in their proportions by 12 per cent., although this bird habitually flies to such an extent that Darwin partly attributes the greatly increased weight of its wing-bones to increased use under domestication.

We find that _all_ the changes are in the direction of shorter and thicker bones--a tendency which must be largely dependent upon the suspension of the rigorous elimination which keeps the bones of the wild duck _long and light_. The used leg-bones and the disused wing-bones have alike been shortened and thickened, though in different proportions. Natural or artificial selection might easily thicken legs without lengthening them, or shorten wings without eliminating strong heavy bones, but it can hardly be contended that use-inheritance has acted in such conflicting ways. The thickening of the wing-bones has actually more than kept pace with any increase of weight in the skeleton, in spite of the effect of individual disuse and of the alleged cumulative effect of ancestral disuse for hundreds of generations. The case of the duck deserves special attention as a crucial one, if only from the fact that in this instance, and in this instance only, has Darwin given the weights of the skeletons, thus furnishing the means for a closer examination of his details than is usually possible.

If we ignore such factors as selection, panmixia, correlation, and the effects of use and disuse during lifetime, and still regard the case of the domestic duck as a valid proof of the inheritance of the effects of use and disuse, we must also accept it as an equally valid proof that the effects of use and disuse are _not_ inherited. Nay, we may even have to admit that, in two points out of four, the _inherited_ effect of use and disuse on successive generations is exactly opposite to the immediate effect on the individual.

Among fowls the wing-bones have lost much in weight but little or nothing in length--which is the reverse of what has occurred in ducks, although disuse is alleged to be the common cause in both cases. Some of the fowls which fly least have their wing-bones as long as ever. In the case of the Silk and Frizzled fowls--ancient breeds which "cannot fly at all"--and in that of the Cochins, which "can hardly fly up to a low perch," Darwin observes "how truly the proportions of an organ may be inherited although not fully exercised during many generations."[26] In four out of twelve breeds the wing-bones had become slightly heavier relatively to the leg-bones. Do not these facts tend to show that the changes in fowls' wings are due to fluctuating variability and selective influences rather than to a general law whereby the effects of disuse are cumulatively inherited?

PIGEONS' WINGS.

Concerning pigeons' wings Darwin says: "As fancy pigeons are generally confined in aviaries of moderate size, and as even when not confined they do not search for their own food, they must during many generations have used their wings incomparably less than the wild rock-pigeon ... but when we turn to the wings we find what at first appears a wholly different and unexpected result."[27] This unexpected increase in the spread of the wings from tip to tip is due to the feathers, which have lengthened in spite of disuse. Excluding the feathers, the wings were shorter in seventeen instances, and longer in eight. But as artificial selection has lengthened the wings in some instances, why may it not have shortened them in others? Wings with shortened bones would fold up more neatly than the long wings of the Carrier pigeon for instance, and so might unconsciously be favoured by fanciers. The selection of elegant birds with longer necks or bodies would cause a relative reduction in the wings--as with the Pouter, where the wings have been greatly lengthened but not so much as the body.[28] Slender bodies, too, and the lessened divergence of the furculum,[29] would slightly diminish the spread of the wings, and so would affect the measurements taken. As the wing-bones, moreover, are to some extent correlated with the beak and the feet, the artificial selection of shortened beaks might tend to shorten the wing as well as the feet. Under these circumstances how can we be sure of the actual efficacy of use-inheritance? Surely selection is as fully competent to effect slight changes in the direction of use-inheritance as it undoubtedly is to effect great changes in direct opposition to that alleged factor of evolution.

SHORTENED BREAST-BONE IN PIGEONS.

The shortening of the sternum in pigeons is attributed to disuse of the flight muscles attached to it. The bone is only shortened by a third of an inch, but this represents a very remarkable reduction in proportional length, which Darwin estimates at from one-seventh to one-eighth, or over 13 per cent. This marked reduction, too, quite unlike the slight reduction of the wing-bones to which the other ends of the muscles are attached, was universal in the eleven specimens measured by Darwin; and the bone, though acknowledged to have been modified by artificial selection in some breeds, is not so open to observation as wings or legs. Even, however, if this relative shortening of the sternum remained otherwise inexplicable, it might still be as irrelevant to use and disuse as is the fact that "many breeds" of fancy pigeons have lost a rib, having only seven where the ancestral rock-pigeon has eight.[30] But the excessive reduction in the sternum is far from being inexplicable. In the first place Darwin has somewhat over-estimated it. Instead of comparing the deficiency of length with the increased length which _should_ have been acquired (since the pigeons have increased in average size) he compares it with the length of the breast-bone in the rock-pigeon.[31] By this method if a pigeon had doubled in dimensions while its breast-bone remained unaltered, the reduction would be put down as 100 per cent., whereas obviously the true reduction would be one-half, or 50 per cent. of what the bone _should be_. Avoiding this error and a minor fallacy besides, a sound estimate reduces the supposed reduction of 13 or 14 per cent. to one of 11.7 per cent., which is still of course a considerable diminution.

Part of this reduction must be due to the direct effect of disuse during the lifetime of the individual. Another and perhaps very considerable part of the relative change must be attributed to the lengthening of the neck or body by artificial selection, or to other modifications of shape and proportion effected directly or indirectly by the same cause.[32] The reduction is greatest in the Pouter (18-1/2 per cent.) and in the Pied Scanderoon (17-1/2 per cent.). In the former the body has been greatly elongated by artificial selection and three or four additional vertebrae have been acquired in the hinder part of the body.[33] In the latter a long neck increases the length of the bird, and so causes, or helps to cause, the relative shortening of the breast-bone. In the English Carrier--which experiences the effects of disuse, as it is too valuable to be flown--the relative reduction of 11 per cent. is apparently more than accounted for by the "elongated neck." The Dragon also has a long neck. In the Pouter, although the breast-bone has been shortened by 18-1/2 per cent. relatively to the length of the body, it has _lengthened_ by 20 per cent. relatively to the _bulk_ of the body.[34] Darwin forgot to ask whether allowance must not be made for a frequent, or perhaps general, elongation of the neck and the hinder part of the body, and the relative shortening or the throwing forward of the central portion containing the ribs (frequently one less in number) and the sternum. The whole body of the pigeon is so much under the control of artificial selection, that every precaution must be taken to guard against such possible sources of error.[35]

Under domestication there would be a suspension of the previous elimination of reduced breast-bones by natural selection (Weismann's panmixia), and a diminution of the parts concerned in flying might even be favoured, as lessened powers of _continuous_ flight would prevent pigeons from straying too far, and would fit them for domestication or confinement. Such causes might reduce some of the less observed parts affected by flying, while still leaving the wing of full size for occasional flight, or to suit the requirements of the pigeon-fanciers. A change might thus be commenced like that seen in the rudimentary keel of the sternum in the owl-parrot of New Zealand, which has lost the power of flight although still retaining fairly-developed wings.

SHORTENED FEET IN PIGEONS.

Darwin thinks it highly probable that the short feet of most breeds of pigeons are due to lessened use, though he owns that the effects of correlation with the shortened beak are more plainly shown than the effects of disuse.[36] But why need the inherited effects of disuse be called in to explain an average reduction of some 5 per cent., when Darwin's measurements show that in the breeds where long beaks are favoured the principle of correlation between these parts has lengthened the foot by 13 per cent. in spite of disuse?

SHORTENED LEGS OF RABBITS.

In the case of the domestic rabbit Darwin notices that the bones of the legs have (relatively) become shorter by an inch and a half. But as the leg-bones have _not_ diminished in relative weight,[37] they must clearly have grown _thicker_ or denser. If disuse has shortened them, as Darwin supposes, why has it also thickened them? The ears and the tail have been lengthened in spite of disuse. Why then may not the ungainly hind-legs have been shortened by human preference independently of the inherited effects of disuse? By relying on apparently favourable instances and neglecting the others it would be easy to arrive at all manner of unsound conclusions. We might thus become convinced that vessels tend to sail northwards, or that a pendulum oscillates more often in one direction than in the other. It must not be forgotten that it would be easy to cite an enormous number of cases which are in direct conflict with the supposed law of use-inheritance.

BLIND CAVE-ANIMALS.