Are the Effects of Use and Disuse Inherited? - An Examination of the View Held by Spencer and Darwin
by William Platt Ball
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My warmest thanks are due to Mr. Francis Darwin, to Mr. E. B. Poulton (whose interest in the subject here discussed is shown by his share in the translation of Weismann's Essays on Heredity), and to Professor Romanes, for the help afforded by their kindly suggestions and criticisms, and for the advice and recommendation under which this essay is now published. Encouragement from Mr. Francis Darwin is to me the more precious, and the more worthy of grateful recognition, from the fact that my general conclusion that acquired characters are not inherited is at variance with the opinion of his revered father, who aided his great theory by the retention of some remains of Lamarck's doctrine of the inherited effect of habit. I feel as if the son, as representative of his great progenitor, were carrying out the idea of an appreciative editor who writes to me: "We must say that if Darwin were still alive, he would find your arguments of great weight, and undoubtedly would give to them the serious consideration which they deserve." I hope, then, that I may be acquitted of undue presumption in opposing a view sanctioned by the author of the Origin of Species, but already stoutly questioned and firmly rejected by such followers of his as Weismann, Wallace, Poulton, Ray Lankester, and others, to say nothing of its practical rejection by so great an authority on heredity as Francis Galton.

The sociological importance of the subject has already been insisted on in emphatic terms by Mr. Herbert Spencer, and this importance may be even greater than he imagined.

Civilization largely sets aside the harsh but ultimately salutary action of the great law of Natural Selection without providing an efficient substitute for preventing degeneracy. The substitute on which moralists and legislators rely—if they think on the matter at all—is the cumulative inheritance of the beneficial effects of education, training, habits, institutions, and so forth—the inheritance, in short, of acquired characters, or of the effects of use and disuse. If this substitute is but a broken reed, then the deeper thinkers who gradually teach the teachers of the people, and ultimately even influence the legislators and moralists, must found their systems of morality and their criticisms of social and political laws and institutions and customs and ideas on the basis of the Darwinian law rather than on that of Lamarck.

Looking forward to the hope that the human race may become consciously and increasingly master of itself and of its destiny, and recognizing the Darwinian principle of the selection of the fittest as the only means of preventing the moral and physical degeneracy which, like an internal dry rot, has hitherto been the besetting danger of all civilizations, I desire that the thinkers who mould the opinions of mankind shall not be led astray from the true path of enduring progress and happiness by reliance on fallacious beliefs which will not bear examination. Such, at least, is the feeling or motive which has prompted me to devote much time and thought to a difficult but important inquiry in a debatable region of inference and conjecture, where (I am afraid) evidence on either side can never be absolutely conclusive, and where, especially, the absolute demonstration of a universal negative cannot reasonably be expected.











The question whether the effects of use and disuse are inherited, or, in other words, whether acquired characters are hereditary, is of considerable interest to the general student of evolution; but it is, or should be, a matter of far deeper interest to the thoughtful philanthropist who desires to ensure the permanent welfare and happiness of the human race. So profoundly important, in fact, are the moral, social, and political conclusions that depend on the answer to this inquiry, that, as Mr. Herbert Spencer rightly says, it "demands, beyond all other questions whatsoever, the attention of scientific men."

It is obvious that we can produce important changes in the individual. We can, for example, improve his muscles by athletics, and his brain by education. The use of organs enlarges and strengthens them; the disuse of parts or faculties weakens them. And so great is the power of habit that it is proverbially spoken of as "second nature." It is thus certain that we can modify the individual. We can strengthen (or weaken) his body; we can improve (or deteriorate) his intellect, his habits, his morals. But there remains the still more important question which we are about to consider. Will such modifications be inherited by the offspring of the modified individual? Does individual improvement transmit itself to descendants independently of personal teaching and example? Have artificially produced changes of structure or habit any inherent tendency to become congenitally transmissible and to be converted in time into fixed traits of constitution or character? Can the philanthropist rely on such a tendency as a hopeful factor in the evolution of mankind?—the only sound and stable basis of a higher and happier state of things being, as he knows or ought to know, the innate and constitutionally-fixed improvement of the race as a whole. If acquired modifications are impressed on the offspring and on the race, the systematic moral training of individuals will in time produce a constitutionally moral race, and we may hope to improve mankind even in defiance of the unnatural selection by which a spurious but highly popular philanthropy would systematically favour the survival of the unfittest and the rapid multiplication of the worst. But if acquired modifications do not tend to be transmitted, if the use or disuse of organs or faculties does not similarly affect posterity by inheritance, then it is evident that no innate improvement in the race can take place without the aid of natural or artificial selection.

Herbert Spencer maintains that the effects of use and disuse are inherited in kind, and in his Factors of Organic Evolution[1] he has supported his contention with a selection of facts and reasonings which I shall have the temerity to examine and criticize. Darwin also held the same view, though not so strongly. And here, to prevent misunderstanding, I may say that the admiration and reverence and gratitude due to Darwin ought not to be allowed to interfere in the slightest degree with the freest criticism of his conclusions. To perfect his work by the correction of really extraneous errors is as much a sacred duty as to study and apply the great truths he has taught.


[1] Which originally appeared in the Nineteenth Century for April and May, 1886.



Mr. Spencer verified this by comparing English jaws with Australian and Negro jaws at the College of Surgeons.[2] He maintains that the diminution of the jaw in civilized races can only have been brought about by inheritance of the effects of lessened use. But if English jaws are lighter and thinner than those of Australians and Negroes, so too is the rest of the skull. As the diminution in the weight and thickness of the walls of the cranium cannot well be ascribed to disuse, it must be attributed to some other cause; and this cause may have affected the jaw also. Cessation of the process by which natural selection[3] favoured strong thick bones during ages of brutal violence might bring about a change in this direction. Lightness of structure, facilitating agility and being economical of material, would also be favoured by natural selection so far as strength was not too seriously diminished.

Sexual selection powerfully affects the human face, and so must affect the jaws—as is shown by the differences between male and female jaws, and by the relative lightness and smallness of the latter, especially in the higher races. Human preference, both sexual and social, would tend to eliminate huge jaws and ferocious teeth when these were no longer needed as weapons of war or organs of prehension, &c. We can hardly assume that the lower half of the face is specially exempt from the influence of natural and sexual selection; and the effects of these undoubted factors of evolution must be fully considered before we are entitled to call in the aid of a factor whose existence is questioned.

After allowing for lost teeth and the consequent alveolar absorption, and for a reduction proportional to that shown in the rest of the skull, the difference in average weight in fifty European and fourteen Australian male jaws at the College of Surgeons turned out to be less than a fifth of an ounce, or about 5 per cent. This slight reduction may be much more than accounted for by such causes as disuse in the individual, human preference setting back the teeth, and partial transference of the much more marked diminution seen in female jaws. There is apparently no room for accumulated inherited effects of ancestral disuse. The number of jaws is small, indeed; but weighing them is at least more decisive than Mr. Spencer's mere inspection.

The differences between Anglo-Saxon male jaws and Australian and Tasmanian jaws are most easily explained as effects of human preference and natural selection. We can hardly suppose that disuse would maintain or develop the projecting chin, increase its perpendicular height till the jaw is deepest and strongest at its extremity, evolve a side flange, and enlarge the upper jaw-bone to form part of a more prominent nose, while drawing back the savagely obtrusive teeth and lips to a more pleasing and subdued position of retirement and of humanized beauty. If human preference and natural selection caused some of these differences, why are they incompetent to effect changes in the direction of a diminution of the jaw or teeth? And if use and disuse are the sole modifying agents in the case of the human jaw, why should men have any more chin than a gorilla or a dog?

The excessive weight of the West African jaws at the College of Surgeons is partly against Mr. Spencer's contention, unless he assumes that Guinea Negroes use their jaws far more than the Australians, a supposition which seems extremely improbable. The heavier skull and narrower molar teeth point however to other factors than increased use.

The striking variability of the human jaw is strongly opposed to the idea of its being under the direct and dominant control of so uniform a cause as ancestral use and disuse. Mr. Spencer regards a variation of 1 oz. as a large one, but I found that the English jaws in the College of Surgeons varied from 1.9 oz. to 4.3 oz. (or 5 oz. if lost teeth were allowed for); Australian jaws varied from 2 oz. to 4.5 oz. (with no lost teeth to allow for); while in Negro jaws the maximum rose to over 5-1/2 oz.[4] In spite of disuse some European jaws were twice as heavy as the lightest Australian jaw, either absolutely or (in some cases) relatively to the cranium. The uniformity of change relied upon by Mr. Spencer is scarcely borne out by the facts so far as male jaws are concerned. The great reduction in the weight of female jaws and skulls evidently points to sexual selection and to panmixia under male protection.

I think, on the whole, we must conclude that the human jaws do not afford satisfactory proof of the inheritance of the effects of use and disuse, inasmuch as the differences in their weight and shape and size can be more reasonably and consistently accounted for as the result of less disputable causes.


The next example, the reduced biting muscles, &c., of lap-dogs is also unsatisfactory as a proof of the inheritance of the effects of disuse; for the change can readily be accounted for without the introduction of such a factor. The previous natural selection of strong jaws and teeth and muscles is reversed. The conscious or unconscious selection of lap-dogs with the least tendency to bite would easily bring about a general enfeeblement of the whole biting apparatus—weakness of the parts concerned favouring harmlessness. Mr. Spencer maintains that the dwindling of the parts concerned in clenching the jaw is certainly not due to artificial selection because the modifications offer no appreciable external signs. Surely hard biting is sufficiently appreciable by the person bitten without any visual admeasurement of the masseter muscles or the zygomatic arches. Disuse during lifetime would also cause some amount of degeneracy; and I am not sure that Mr. Spencer is right in entirely excluding economy of nutrition from the problem. Breeders would not over-feed these dogs; and the puppies that grew most rapidly would usually be favoured.


The too closely-packed teeth in the "decreasing" jaws of modern men (p. 13)[5] are also suggestive of other causes than use and disuse. Why is there not simultaneous variation in teeth and jaws, if disuse is the governing factor? Are we to suppose that the size of the human teeth is maintained by use at the same time that the jaws are being diminished by disuse? Mr. Spencer acknowledges that the crowding of bull-dogs' and lap-dogs' teeth is caused by the artificial selection of shortened jaws. If a similar change is really occurring in man, could it not be similarly explained by some factor, such as sexual selection, which might affect the outward appearance at the cost of less obvious defects or inconveniences?

Mr. Spencer points to the decay of modern teeth as a sign or result of their being overcrowded through the diminution of the jaw by disuse.[6] But the teeth which are the most frequently overcrowded are the lower incisors. The upper incisors are less overcrowded, being commonly pressed outwards by the lower arc of teeth fitting inside them in biting. The lower incisors are correspondingly pressed inwards and closer together. Yet the upper incisors decay—or at least are extracted—about twenty times as frequently as the closely packed lower incisors.[7] Surely this must indicate that the cause of decay is not overcrowding.

The lateness and irregularity of the wisdom teeth are sometimes supposed to indicate their gradual disappearance through want of room in a diminishing jaw. But a note on Tasmanian skulls in the Catalogue of the College of Surgeons (p. 199) shows that this lateness and irregularity have been common among Tasmanians as well as among civilized races, so that the change can hardly be attributed to the effects of disuse under civilization.


The cave-crabs which have lost their disused eyes but not the disused eye-stalks appear to illustrate the effects of natural selection rather than of disuse. The loss of the exposed, sensitive, and worse-than-useless eye, would be a decided gain, while the disused eye-stalk, being no particular detriment to the crab, would be but slightly affected by natural selection, though open to the cumulative effects of disuse. The disused but better protected eyes of the blind cave-rat are still "of large size" (Origin of Species, p. 110).


It is but fair to add that these instances of the cave-crab's eye-stalk and the closely-packed teeth are put forward by Mr. Spencer with the more immediate object of proving that there is "no concomitant variation in co-operative parts," even when "formed out of the same tissue, like the crab's eye and its peduncle" (pp. 12-14, 23, 33). It escapes his notice, however, that in two out of his three cases it is disuse, or diminished use, which fails to cause concomitant variation or proportionate variation.


Having unwittingly shown that lessened use of closely-connected and co-operative parts does not cause concomitant variation in these parts, Mr. Spencer concludes that the concomitant variation requisite for evolution can only be caused by altered degrees of use or disuse. He elaborately argues that the many co-ordinated modifications of parts necessitated by each important alteration in an animal are so complex that they cannot possibly be brought about except by the inherited effect of the use and disuse of the various parts concerned. He holds, for instance, that natural selection is inadequate to effect the numerous concomitant changes necessitated by such developments as that of the long neck of the giraffe. Darwin, however, on the contrary, holds that natural selection alone "would have sufficed for the production of this remarkable quadruped."[8] He is surprised at Mr. Spencer's view that natural selection can do so little in modifying the higher animals. Thus one of the chief arguments with which Mr. Spencer supports his theory is so poorly founded as to be rejected by a far greater authority on such subjects. All that is needed is that natural selection should preserve the tallest giraffes through times of famine by their being able to reach otherwise inaccessible stores of foliage. The continual variability of all parts of the higher animals gives scope for innumerable changes, and Nature is not in a hurry. Mr. Spencer, however, says that "the chances against any adequate readjustments fortuitously arising must be infinity to one." But he has also shown that altered degree of use does not cause the needed concomitant variation of co-operative parts. So the chances against a beneficial change in an animal must be, at a liberal estimate, infinity to two. Mr. Spencer, if he has proved anything, has proved that it is practically impossible that the giraffe can have acquired a long neck, or the elk its huge horns, or that any species has ever acquired any important modification.

Mr. Wallace, in his Darwinism, answers Mr. Spencer by a collection of facts showing that "variation is the rule," that the range of variation in wild animals and plants is much greater than was supposed, and that "each part varies to a considerable extent independently" of other parts, so that "the materials constantly ready for natural selection to act upon are abundant in quantity and very varied in kind." While co-operative parts would often be more or less correlated, so that they would tend to vary together, coincident variation is not necessary. The lengthened wing might be gained in one generation, and the strengthened muscle at a subsequent period; the bird in the meanwhile drawing upon its surplus energy, aided (as I would suggest) by the strengthening effect of increased use in the individual. Seeing that artificial selection of complicated variations has modified animals in many points either simultaneously or by slow steps, as with otter-sheep, fancy pigeons, &c. (many of the characters thus obtained being clearly independent of use and disuse), natural selection must be credited with similar powers, and Mr. Wallace concludes that Mr. Spencer's insuperable difficulty is "wholly imaginary."

The extract concerning a somewhat similar "class of difficulties," which Mr. Spencer quotes from his Principles of Biology, is faulty in its reasoning,[9] though legitimate in its conclusion concerning the increasing difficulty of evolution in proportion with the increasing number and complexity of faculties to be evolved. But this increasing difficulty of complex evolution is only overcome by some favourably-varying individuals and species—not by all. And as the difficulty increases we find neglect and decay of the less-needed faculties—as with domesticated animals and civilized men, who lose in one direction while they gain in another. The increasing difficulty of complex evolution by natural selection is no proof whatever of use-inheritance[10] except to those who confound difficulty with impossibility.


Mr. Spencer further contends that natural selection, by unduly developing specially advantageous modifications without the necessary but complex secondary modifications, would render the constitution of a variety "unworkable" (p. 23). But this seems hardly feasible, seeing that natural selection must continually favour the most workable constitutions, and will only preserve organisms in proportion as they combine general workableness with the special modification. On the other hand, according to Mr. Spencer himself, use-inheritance must often disturb the balance of the constitution. Thus it tends to make the jaws and teeth unworkable through the overcrowding and decay of the teeth—there being, as his illustrations show, no simultaneous or concomitant or proportional variation in relation to altered degree of use or disuse.


Mr. Spencer also holds that most mental phenomena, especially where complex or social or moral, can only be explained as arising from use-inheritance, which becomes more and more important as a factor of evolution as we advance from the vegetable world and the lower grades of animal life to the more complex activities, tastes, and habits of the higher organizations (preface, and p. 74). But there happens to be a tolerably clear proof that such changes as the evolution of complicated structures and habits and social instincts can take place independently of use-inheritance. The wonderful instincts of the working bees have apparently been evolved (at least in all their later social complications and developments) without the aid of use-inheritance—nay, in spite of its utmost opposition. Working bees, being infertile "neuters," cannot as a rule transmit their own modifications and habits. They are descended from countless generations of queen bees and drones, whose habits have been widely different from those of the workers, and whose structures are dissimilar in various respects. In many species of ants there are two, and in the leaf-cutting ants of Brazil there are three, kinds of neuters which differ from each other and from their male and female ancestors "to an almost incredible degree."[11] The soldier caste is distinguished from the workers by enormously large heads, very powerful mandibles, and "extraordinarily different" instincts. In the driver ant of West Africa one kind of neuter is three times the size of the other, and has jaws nearly five times as long. In another case "the workers of one caste alone carry a wonderful sort of shield on their heads." One of the three neuter classes in the leaf-cutting ants has a single eye in the midst of its forehead. In certain Mexican and Australian ants some of the neuters have huge spherical abdomens, which serve as living reservoirs of honey for the use of the community. In the equally wonderful case of the termites, or so-called "white ants" (which belong, however, to an entirely different order of insect from the ants and bees) the neuters are blind and wingless, and are divided into soldiers and workers, each class possessing the requisite instincts and structures adapting it for its tasks. Seeing that natural selection can form and maintain the various structures and the exceedingly complicated instincts of ants and bees and wasps and termites in direct defiance of the alleged tendency to use-inheritance, surely we may believe that natural selection, unopposed by use-inheritance, is equally competent for the work of complex or social or mental evolution in the many cases where the strong presumptive evidence cannot be rendered almost indisputable by the exceptional exclusion of the modified animal from the work of reproduction.

Ants and bees seem to be capable of altering their habits and methods of action much as men do. Bees taken to Australia cease to store honey after a few years' experience of the mild winters. Whole communities of bees sometimes take to theft, and live by plundering hives, first killing the queen to create dismay among the workers. Slave ants attend devotedly to their captors, and fight against their own species. Forel reared an artificial ant-colony made up of five different and more or less hostile species. Why cannot a much more intelligent animal modify his habits far more rapidly and comprehensively without the aid of a factor which is clearly unnecessary in the case of the more intelligent of the social insects?


The modern development of music and harmony (p. 19) is undeniable, but why could it only have been brought about by the help of the inheritance of the effects of use? Why are we to suppose that "minor traits" such as the "aesthetic perceptions" cannot have been evolved by natural selection (p. 20) or by sexual selection? Darwin holds that our musical faculties were developed by sexual preference long before the acquisition of speech. He believes that the "rhythms and cadences of oratory are derived from previously developed musical powers"—a conclusion "exactly opposite" to that arrived at by Mr. Spencer.[12] The emotional susceptibility to music, and the delicate perceptions needed for the higher branches of art, were apparently the work of natural and sexual selection in the long past. Civilization, with its leisure and wealth and accumulated knowledge, perfects human faculties by artificial cultivation, develops and combines means of enjoyment, and discovers unsuspected sources of interest and pleasure. The sense of harmony, modern as it seems to be, must have been a latent and indirect consequence of the development of the sense of hearing and of melody. Use, at least, could never have called it into existence. Nature favours and develops enjoyments to a certain extent, for they subserve self-preservation and sexual and social preference in innumerable ways. But modern aesthetic advance seems to be almost entirely due to the culture of latent abilities, the formation of complex associations, the selection and encouragement of talent, and the wide diffusion and imitation of the accumulated products of the well-cultivated genius of favourably varying individuals. The fact that uneducated persons do not enjoy the higher tastes, and the rapidity with which such tastes are acquired or professed, ought to be sufficient proof that modern culture is brought about by far swifter and more potent influences than use-inheritance. Neither would this hypothetical factor of evolution materially aid in explaining the many other rapid changes of habit brought about by education, custom, and the changed conditions of civilization generally. Powerful tastes—as is incontestably shown in the cases of alcohol and tobacco—lie latent for ages, and suddenly become manifest when suitable conditions arise. Every discovery, and each step in social and moral evolution, produces its wide-spreading train of consequences. I see no reason why use-inheritance need be credited with any share in the cumulative results of the invention of printing and the steam-engine and gunpowder, or of freedom and security under representative government, or of science and art and the partial emancipation of the mind of man from superstition, or of the innumerable other improvements or changes that take place under modern civilization.

Mr. Spencer suggests an inquiry whether the greater powers possessed by eminent musicians were not mainly due to the inherited effect of the musical practice of their fathers (p. 19). But these great musicians inherited far more than their parents possessed. The excess of their powers beyond their parents' must surely be attributed to spontaneous variation; and who shall say that the rest was in any way due to use-inheritance? If, too, the superiority of geniuses proves use-inheritance, why should not the inferiority of the sons of geniuses prove the existence of a tendency which is the exact opposite of use-inheritance? But nobody collects facts concerning the degenerate branches of musical families. Only the favourably varying branches are noticed, and a general impression of rapid evolution of talent is thus produced. Such cases might be explained, too, by the facts that musical faculty is strong in both sexes, that musical families associate together, and that the more gifted members may intermarry. Great musicians are often astonishingly precocious. Meyerbeer "played brilliantly" at the age of six. Mozart played beautifully at four. Are we to suppose that the effect of the adult practice of parents was inherited at this early age? If use-inheritance was not necessary in the case of Handel, whose father was a surgeon, why is it needed to account for Bach?


The "direct proofs" of use-inheritance are not as plentiful as might be desired, it appears (pp. 24-28). This acknowledged "lack of recognized evidence" is indeed the weakest feature in the case, though Mr. Spencer would fain attribute this lack of direct proof to insufficient investigation and to the inconspicuous nature of the inheritance of the modification. But there is an almost endless abundance of conspicuous examples of the effects of use and disuse in the individual. How is it that the subsequent inheritance of these effects has not been more satisfactorily observed and investigated? Horse-breeders and others could profit by such a tendency, and one cannot help suspecting that the reason they ignore it must be its practical inefficacy, arising probably from its weakness, its obscurity and uncertainty or its non-existence.


Brown-Sequard's discovery that an epileptic tendency artificially produced by mutilating the nervous system of a guinea-pig is occasionally inherited may be a fact of "considerable weight," or on the other hand it may be entirely irrelevant. Cases of this kind strike one as peculiar exceptions rather than as examples of a general rule or law. They seem to show that certain morbid conditions may occasionally affect both the individual and the reproductive elements or transmissible type in a similar manner; but then we also know that such prompt and complete transmission of an artificial modification is widely different from the usual rule. Exceptional cases require exceptional explanations, and are scarcely good examples of the effect of a general tendency which in almost all other cases is so inconspicuous in its immediate effects. Further remarks on this inherited epilepsy can be most conveniently introduced later on in connection with Darwin's explanation of the inherited mutilation which it usually accompanies, but which Mr. Spencer does not mention.


Mr. Spencer infers that, because insanity is usually hereditary, and insanity can be artificially produced by various excesses, therefore this artificially-produced insanity must also be hereditary (p. 28). Direct evidence of this conclusion would be better than a mere inference which may beg the very question at issue. That the liability to insanity commonly runs in families is no proof that strictly non-inherited insanity will subsequently become hereditary. I think that theories should be based on facts rather than facts on theories, especially when those facts are to be the basis or proof of a further theory.

Mr. Spencer also points out that he finds among physicians "the belief that nervous disorders of a less severe kind are inheritable"—a general belief which does not necessarily include the transmission of purely artificially-produced disorders, and so misses the point which is really at issue. He proceeds, however, to state more definitely that "men who have prostrated their nervous systems by prolonged overwork or in some other way, have children more or less prone to nervousness." The following observations will, I think, warrant at least a suspension of judgment concerning this particular form of use-inheritance.

(1) The nervousness is seen in the children at an early age, although the nervous prostration from which it is supposed to be derived obviously occurs in the parent at a much later period of life. This change in time is contrary to the rule of inheritance at corresponding periods; and, together with the unusual promptness and comparative completeness of the inheritance, it may indicate a special injury or deterioration of the reproductive elements rather than true inheritance. The healthy brain of early life has failed to transmit its robust condition. Is use-inheritance, then, only effective for evil? Does it only transfer the newly-acquired weakness, and not the previous long-continued vigour?

(2) Members of nervous families would be liable to suffer from nervous prostration, and by the ordinary law of heredity alone would transmit nervousness to their children.

(3) The shattered nerves or insanity resulting from alcoholic and other excesses, or from overwork or trouble, are evidently signs of a grave constitutional injury which may react upon the reproductive elements nourished and developed in that ruined constitution. The deterioration in parent and child may often display itself in the same organs—those probably which are hereditarily weakest. Acquired diseases or disorders thus appear to be transmitted, when all that was conveyed to the offspring was the exciting cause of a lowered vitality or disordered action, together with the ancestral liability to such diseases under such conditions.

(4) Francis Galton says that "it is hard to find evidence of the power of the personal structure to react upon the sexual elements, that is not open to serious objection." Some of the cases of apparent inheritance he regards as coincidence of effect. Thus "the fact that a drunkard will often have imbecile children, although his offspring previous to his taking to drink were healthy," is an "instance of simultaneous action," and not of true inheritance. "The alcohol pervades his tissues, and, of course, affects the germinal matter in the sexual elements as much as it does that in his own structural cells, which have led to an alteration in the quality of his own nerves. Exactly the same must occur in the case of many constitutional diseases that have been acquired by long-continued irregular habits."[13]


Mr. Spencer finds it hard to believe that the modifications conveyed to offspring are not identical in tendency with the changes effected in the parent by altered use or habit (pp. 23-25, 34). But it is perfectly certain that the two sets of effects do not necessarily correspond. The effect of changed habits or conditions on the individual is often very far from coinciding with the effects on the reproductive elements or the transmissible type. The reproductive system is "extremely sensitive" to very slight changes, and is often powerfully affected by circumstances which otherwise have little effect on the individual (Origin of Species, p. 7). Various animals and plants become sterile when domesticated or supplied with too much nourishment. The native Tasmanians have already become extinct from sterility caused by greatly changed diet and habits. If, as Mr. Spencer teaches, continued culture and brain-work will in time produce lessened fertility or comparative sterility, we may yet have to be careful that intellectual development does not become a species of suicide, and that the culture of the race does not mean its extinction—or at least the extinction of those most susceptible of culture.

The reproductive elements are also disturbed and modified in innumerable minor ways. Changed conditions or habits tend to produce a general "plasticity" of type, the "indefinite variability" thus caused being apparently irrelevant to the change, if any, in the individual.[14] A vast number of variations of structure have certainly arisen independently of similar parental modification as the preliminary. Whatever first caused these "spontaneous" congenital variations affected the reproductive elements quite differently from the individual. "When a new peculiarity first appears we can never predict whether it will be inherited." Many varieties of plants only keep true from shoots, and not from seed, which is by no means acted on in the same way as the individual plant. Seeing that such plants have two reproductive types, both constant, it is evident that these cannot both be modified in the same way as the parent is modified. Many parental modifications of structure and habit are certainly not conveyed to neuter ants and bees; other modifications, which are not seen in the parents, being conveyed instead. Many other circumstances tend to show that the individual and the transmissible type are independent of each other so far as modifications of parts are concerned.

It may seem natural to expect the transmission of an enlarged muscle or a cultivated brain, but, on the other hand, why should it be unreasonable to expect that a modification which was non-congenital in origin should still remain non-congenital? Why should the non-transmission of that which was not transmitted be surprising?

Mr. Spencer thinks that the non-transmission of acquired modifications is incongruous with the great fact of atavism. But the great law of the inheritance of that which is a development of the transmissible type does not necessarily imply the inheritance of modifications acquired by the individual. Because English children may inherit blue eyes and flaxen hair from their Anglo-Saxon ancestors, it by no means follows that an Englishman must inherit his father's sunburnt complexion or smooth-shaven face. Of course atavism ultimately adopts many instances of revolt against its sway. But to assume that these changes of type follow the personal change rather than cause it, is to assume the whole question at issue. That like begets like is true as a broad principle, but it has many exceptions, and the non-heredity of acquired characters may be one of them.


[2] Principles of Biology, Sec. 166, footnote. The English jaws are somewhat lighter than the Australian jaws, though I could not undertake to affirm that they are really shorter and smaller. In the typical skulls depicted on p. 68 of the official guide to the mammalian galleries at South Kensington, the typical Caucasian jaw is very much larger than the Tasmanian jaw, although the repulsively obtrusive teeth of the latter convey the contrary idea to the imagination. Mr. Spencer's assumption that the ancient Britons had large jaws appears to me erroneous. (See Professor Rolleston's Scientific Papers and Addresses, i. p. 250.)

[3] Romanes, Galton, and Weismann have made great use of this principle in explaining the diminution of disused organs. Weismann has given it the name of Panmixia,—all individuals being equally free to survive and commingle their variations, and not merely selected or favoured individuals. See his Essays on Heredity, &c., p. 90 (Clarendon Press).

[4] Inclusive in each case of fixed strengthening wire weighing about a sixteenth of an ounce or less.

[5] References of course are to Factors of Organic Evolution.

[6] P. 13; and Nineteenth Century, February, 1888, p. 211.

[7] Tomes's Dental Surgery, pp. 273-275. Tomes observes that it is as yet uncertain in what way civilization predisposes to caries. But he shows that caries is caused by the lime salts in the teeth being attacked by acids from decomposing food in crevices, from artificial drink such as cyder, from sugar, from medicine, and from vitiated secretions of the mouth. It is evident that in civilized races natural selection cannot so rigorously insist on sound teeth, sound constitutions, and protective alkaline saliva. The reaction of the civilized mouth is often acid, especially when the system is disordered by dyspepsia or other diseases or forms of ill-health common under civilization. The main supply of saliva, which is poured from the cheeks opposite the upper molars, is often acid when in small quantities. But the submaxillary and sub-lingual saliva poured out at the foot of the lower incisors and held in the front part of the jaw as in a spoon, "differs from parotid saliva in being more alkaline" (Foster's Text Book of Physiology, p. 238; Tomes, pp. 284, 685). One observer says that the reaction near the lower incisors is "never acid." Hence (I conclude) the remarkable immunity of the lower incisors and canines from decay, an immunity which extends backwards in a lessening degree to the first and second bicuspids. The close packing of the lower incisors may assist by preventing the retention of decaying fragments of food. Sexual selection may promote caries by favouring white teeth, which are more prone to decay than yellow ones. Acid vitiation of the mucus might account both for caries and (possibly) for the strange infertility of some inferior races under civilization.

[8] Origin of Species, pp. 198-9; Variation of Animals and Plants under Domestication, vol. ii. p. 328 footnote, also p. 206.

[9] Mr. Spencer weakly argues that an advantageous attribute (such as swiftness, keen sight, courage, sagacity, strength, &c.) cannot be increased by natural selection unless it is "of greater importance, for the time being, than most of the other attributes"; and that natural selection cannot develop any one superiority when animals are equally preserved by "other superiorities." But as natural selection will simultaneously eliminate tendencies to slowness, blindness, deafness, stupidity, &c., it must favour and improve many points simultaneously, although no one of them may be of greater importance than the rest. Of course the more complicated the evolution the slower it will be; but time is plentiful, and the amount of elimination is correspondingly vast.

[10] I venture to coin this concise term to signify the direct inheritance of the effects of use and disuse in kind. Having a name for a thing is highly convenient; it facilitates clearness and accuracy in reasoning, and in this particular inquiry it may save some confusion of thought from double or incomplete meanings in the shortened phrases which would otherwise have to be employed to indicate this great but nameless factor of evolution.

[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).


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.


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.


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.


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?


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.


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.


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?


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.


Weak or defective eyesight is by no means rare as a spontaneous variation in animals, "the great French veterinary Huzard going so far as to say that a blind race [of horses] could soon be formed." Natural selection evolves blind races whenever eyes are useless or disadvantageous, as with parasites. This may apparently be done independently of the effects of disuse, for certain neuter ants have eyes which are reduced to a more or less rudimentary condition, and neuter termites are blind as well as wingless. In one species of ant (Eciton vastator) the sockets have disappeared as well as the eyes. In deep caves not only would natural selection cease to maintain good eyesight but it would persistently favour blindness—or the entire removal of the eye when greatly exposed, as in the cave-crab—and as Dr. Ray Lankester has indicated,[38] there would have been a previous selection of animals which through spontaneous weakness, sensitiveness, or other affection of the eye found refuge and preservation in the cave, and a subsequent selection of the descendants whose fitness for relative darkness led them deeper into the cave or prevented them from straying back to the light with its various dangers and severer competition. Panmixia, however, as Weismann has shown, would probably be the most important factor in causing blindness.


Darwin says: "A horse is trained to certain paces, and the colt inherits similar consensual movements."[39] But selection of the constitutional tendency to these paces, and imitation of the mother by the colt, may have been the real causes. The evidence, to be satisfactory, should show that such influences were excluded. Men acquire proficiency in swimming, waltzing, walking, smoking, languages, handicrafts, religious beliefs, &c., but the children only appear to inherit the innate abilities or constitutional proclivities of their parents. Even the songs of birds, including their call-notes, are no more inherited than is language by man (Descent of Man, p. 86). They are learned from the parent. Nestlings which acquire the song of a distinct species, "teach and transmit their new song to their offspring." If use-inheritance has not fixed the song of birds, why should we suppose that in a single generation it has transmitted a newly-taught method of walking or trotting?

It is alleged that dogs inherit the intelligence acquired by association with man, and that retrievers inherit the effects of their training.[40] But selection and imitation are so potent that the additional hypothesis of use-inheritance seems perfectly superfluous. Where intelligence is not highly valued and carefully promoted by selection, the intelligence derivable from association with man does not appear to be inherited. Lap-dogs, for instance, are often remarkably stupid.

Darwin also instances the inheritance of dexterity in seal-catching as a case of use-inheritance.[41] But this is amply explained by the ordinary law of heredity. All that is needed is that the son shall inherit the suitable faculties which the father inherited before him.


Darwin holds that in some cases selection alone has modified the instincts and dispositions of domesticated animals, but that in most cases selection and the inheritance of acquired habits have concurred in effecting the change. "On the other hand," he says, "habit alone in some cases has sufficed; hardly any animal is more difficult to tame than the young of the wild rabbit; scarcely any animal is tamer than the young of the tame rabbit; but I can hardly suppose that domestic rabbits have often been selected for tameness alone; so that we must attribute at least the greater part of the inherited change from extreme wildness to extreme tameness to habit and long-continued close confinement."[42]

But there are strong, and to me irresistible, arguments to the contrary. I think that the following considerations will show that the greater part, if not the whole, of the change must be attributed to selection rather than to the direct inheritance of acquired habit.

(1) For a period which may cover thousands of generations, there has been an entire cessation of the natural selection which maintains the wildness (or excessive fear, caution, activity, &c.) so indispensably essential for preserving defenceless wild rabbits of all ages from the many enemies that prey upon them.

(2) During this same extensive period of time man has usually killed off the wildest and bred from the tamest and most manageable. To some extent he has done this consciously. "It is very conducive to successful breeding to keep only such as are quiet and tractable," says an authority on rabbits,[43] and he enjoins the selection of the handsomest and best-tempered does to serve as breeders. To a still greater extent man has favoured tameness unconsciously and indirectly. He has systematically selected the largest and most prolific animals, and has thus doubled the size and the fertility of the domestic rabbit. In consciously selecting the largest and most flourishing individuals and the best and most prolific mothers, he must have unconsciously selected those rabbits whose relative tameness or placidity of disposition rendered it possible for them to flourish and to produce and rear large and thriving families, instead of fretting and pining as the wilder captives would do. When we consider how exceedingly delicate and easily disturbed yet all-important a function is that of maternity in the continually breeding rabbit, we see that the tamest and the least terrified would be the most successful mothers, and so would continually be selected, although man cared nothing for the tameness in itself. The tamest mothers would also be less liable to neglect or devour their offspring, as rabbits commonly do when their young are handled too soon, or even when merely frightened by mice, &c., or disturbed by changed surroundings.

(3) We must remember the extraordinary fecundity of the rabbit and the excessive amount of elimination that consequently takes place either naturally or artificially. Where nature preserved only the wildest, man has preserved the tamest. If there is any truth in the Darwinian theory, this thorough and long-continued reversal of the selective process must have had a powerful effect. Why should it not be amply sufficient to account for the tameness and mental degeneracy of the rabbit without the aid of a factor which can readily be shown to be far weaker in its normal action than either natural or artificial selection? Why may not the tameness of the rabbit be transferred to the group of cases in which Darwin holds that "habit has done nothing," and selection has done all?

(4) If use-inheritance has tamed the rabbit, why are the bucks still so mischievous and unruly? Why is the Angora breed the only one in which the males show no desire to destroy the young? Why, too, should use-inheritance be so much more powerful in the rabbit than with other animals which are far more easily tamed in the first instance? Wild young rabbits when domesticated "remain unconquerably wild," and, although they may be kept alive, they pine and "rarely come to any good." Yet the animal which acquires least tameness—or apparently, indeed, none at all—inherits most! It appears, in fact, to inherit that which it cannot acquire—a circumstance which indicates the selection of spontaneous variations rather than the inheritance of changed habits. Such variations occasionally occur in animals in a marked degree. Of a litter of wolf-cubs, all brought up in the same way, "one became tame and gentle like a dog, while the others preserved their natural savagery." Is it not probable that permanent domestication was rendered possible by the inevitable selection of spontaneous variations in this direction? The excessive tameness, too, of the young rabbit, while easily explicable as a result of unconscious selection, is not easily explained as a result of acquired habit. No particular care is taken to tame or teach or domesticate rabbits. They are bred for food, or for profit or appearance, and they are left to themselves most of their time. As Sir J. Sebright notices with some surprise, the domestic rabbit "is not often visited, and seldom handled, and yet it is always tame."


Innumerable modifications in accordance with altered use or disuse, such as the enlarged udders of cows and goats, and the diminished lungs and livers in highly bred animals that take little exercise, can be readily and fully explained as depending on selection. As the fittest for the natural or artificial requirements will be favoured, natural or artificial selection may easily enlarge organs that are increasingly used and economize in those that are less needed. I therefore see no necessity whatever for calling in the aid of use-inheritance as Darwin does, to account for enlarged udders, or diminished lungs, or the thick arms and thin legs of canoe Indians, or the enlarged chests of mountaineers, or the diminished eyes of moles, or the lost feet of certain beetles, or the reduced wings of logger-headed ducks, or the prehensile tails of monkeys, or the displaced eyes of soles, or the altered number of teeth in plaice, or the increased fertility of domesticated animals, or the shortened legs and snouts of pigs, or the shortened intestines of tame rabbits, or the lengthened intestines of domestic cats, &c.[44] Changed habits and the requisite change of structure will usually be favoured by natural selection; for habit, as Darwin says, "almost implies that some benefit great or small is thus derived."


Here we perceive a difficulty which will equally trouble those who affirm use-inheritance and those who deny. Broadly speaking, the adaptive effects ascribed to use-inheritance coincide with the effects of natural selection. The individual adaptability (as shown in the thickening of skin, fur, muscle, &c., under the stimulus of friction, cold, use, &c.) is identical in kind and direction with the racial adaptability under natural selection. Consequently the alleged inheritance of the advantageous effects of use and disuse cannot readily be distinguished from the similarly beneficial effects of natural selection. The indisputable fact that natural selection imitates or simulates the beneficial effects ascribed to use-inheritance may be the chief source and explanation of a belief which may prove to be thoroughly fallacious. A similar simulation of course occurs under domestication, where natural selection is partly replaced by artificial selection of the best adapted and therefore most flourishing animals, while in disused parts panmixia or the comparative cessation of selection will aid or replace "economy of growth" in causing diminution.[45]


"The inferiority of Europeans, in comparison with savages, in eyesight and in the other senses," is attributed to "the accumulated and transmitted effect of lessened use during many generations."[46] But why may we not attribute it to the slackened and diverted action of the natural selection which keeps the senses so keen in some savage races?


Darwin notices that watchmakers and engravers are liable to be short-sighted, and that short-sight and long-sight certainly tend to be inherited.[47] But we must be careful not to beg the question at issue by assuming that the frequent heredity of short sight necessarily covers the heredity of artificially-produced short-sight. Elsewhere, however, Darwin states more decisively that "there is ground for believing that it may often originate in causes acting on the individual affected, and may thence-forward become transmissible."[48] This impression may arise (1) from the facts of ordinary heredity—the ancestral liability being excited in father and son by similar artificial habits, such as reading, and viewing objects closely as among watchmakers and engravers—or by constitutional deterioration from indoor life, &c., acting upon a constitutional liability of the eye to the "something like inflammation of the coats, under which they yield" and so cause shortness of sight by altering the spherical shape of the eye-ball. (2) Panmixia, or the suspension of natural selection, together with altered habits, will account for an increase of short-sight among the population generally. (3) Long-sighted people could not work at watchmaking and engraving so comfortably and advantageously as at other occupations, and hence would be less likely to take to such callings.


These are best explained as the result of natural selection and of the diminution of the hand by sexual selection in the gentry. If the larger hands of labourers' infants are really due to the inherited effects of ancestral use, why does the development occur so early in life, instead of only at a corresponding period, as is the rule? During the first few years of its life, at least, the labourer's infant does no more work than the gentleman's child. Why are not the effects of this disuse inherited by the labourer's infant? If the enlargement of the infant's hand illustrates the transference of a character gained later in life, it is evident that the transference must take place in spite of the inherited effects of disuse.


Darwin also attributes the thickened sole in infants, "long before birth," to "the inherited effects of pressure during a long series of generations."[50] But disuse should make the infant's sole thin, and it is this thinness that should be inherited. If we suppose the inheritance of the thickened soles of later life to be transferred to an earlier period, we have the anomaly of the inherited effects of disuse at that earlier period being overpowered by the untimely inheritance of the effects of use at another. On the other hand, it is clear that natural selection would favour thickened soles for walking on, and might also promote an early development which would ensure their being ready in good time for actual use; for variations in the direction of delay would be cut off, while variations in the other direction would be preserved. Anyhow, the mere transference of a character to an earlier period is no proof of use-inheritance. The real question is whether the thickened sole was gained by natural selection or by the inherited effects of pressure, and the mere transference or hastened appearance of the thickening does not in any degree solve this question. It merely excludes the effect of disuse during lifetime, and thus presents a fallacious appearance of being decisive. The thickened sole of the unborn infant, however, like the lanugo or hairy covering, is probably a result of the direct inheritance of ancestral stages of evolution, of which the embryo presents a condensed epitome. While the relative thinness of the infant's sole might be pointed to as the effect of disuse during a long series of generations, its thickness is rather an illustration of atavism still resisting the effects of long-continued disuse. There is nothing to show that the inheritable portion of the full original thickness was not gained by natural selection rather than by the directly inherited effect of use; and the latter, being cumulative and indiscriminative in its action, would apparently have made the sole very much thicker and harder than it is. If natural selection were not supreme in such cases, how could we account for the effects of pressure resulting in hard hoofs in some cases and only soft pads in others?

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