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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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A SOURCE OF MENTAL CONFUSION.

Of course in a certain sense this thickening of the sole has resulted from use. In one sense or other, most—or perhaps all—of the results of natural selection are inherited effects of use or disuse. Natural selection preserves that which is of use and which is used, while it eliminates that which is useless and is not used. The most confident assertions of the effects of use and disuse in modifying the heritable type, appear to rest on this indefeasible basis. Darwin's statements concerning the effects of use and disuse in evolution can frequently be read in two senses. They often command assent as undeniable truisms as they stand, but are of course written in another and more debatable sense. Thus in the case of the shortened wings and thickened legs of the domestic duck, I believe equally with Darwin and Spencer that "no one will dispute that they have resulted from the lessened use of the wings and the increased use of the legs." "Use" is at bottom the determining circumstance in evolution generally. The trunk of the elephant, the fin of the fish, the wing of the bird, the cunning hand of man and his complicated brain—and, in short, all organs and faculties whatsoever—can only have been moulded and developed by use—by usefulness and by using—but not necessarily by use-inheritance, not necessarily by directly inherited effects of use or disuse of parts in the individual. So, too, reduced or rudimentary organs are due to disuse, but it by no means follows that the diminution is caused by any direct tendency to the inheritance of the effects of disuse in the individual. The effects of natural selection are commonly expressible as effects of use and disuse, just as adaptation in nature is expressible in the language of teleology. But use-inheritance is no more proven by one of these necessary coincidences than special design is by the other. The inevitable simulation of use-inheritance may be entirely deceptive.

Darwin thinks that "there can be no doubt that use in our domestic animals has strengthened and enlarged certain parts, and disuse diminished them; and that such modifications are inherited." Undoubtedly "such" or similar modifications have often been inherited, but how can Darwin possibly tell that they are not due to the simulation of use-inheritance by natural or artificial selection acting upon general variability? Of the inevitability of selection and of its generally adaptive tendencies "there can be no doubt," and panmixia would tend to reduce disused parts; so that there must always remain grave doubts of the alleged inheritance of the similar effects of use and disuse, unless we can accomplish the extremely difficult feat of excluding both natural and artificial selection as causes of enlargement, and panmixia and selection as causes of dwindling.

WEAKNESS OF USE-INHERITANCE.

Use-inheritance is normally so weak that it appears to be quite helpless when opposed to any other factor of evolution. Natural selection evolves and maintains the instincts of ants and termites in spite of use-inheritance to a more wonderful degree than it evolves the instincts of almost any other animal with the fullest help of use-inheritance. It develops seldom-used horns or natural armour just as readily as constantly-used hoofs or teeth. Sexual selection evolves elaborate structures like the peacock's tail in spite of disuse and natural selection combined. Artificial selection appears to enlarge or diminish used parts or disused parts with equal facility. The assistance of use-inheritance seems to be as unnecessary as its opposition is ineffective.

The alleged inheritance of the effects of use and disuse in our domestic animals must be very slow and slight.[51] Darwin tells us that "there is no good evidence that this ever follows in the course of a single generation." "Several generations must be subjected to changed habits for any appreciable result."[52] What does this mean? One of two things. Either the tendency is very weak, or it is non-existent. If it is so weak that we cannot detect its alleged effects till several generations have elapsed, during which time the more powerful agency of selection has been at work, how are we to distinguish the effects of the minor factor from that of the major? Are we to conclude that use-inheritance plus selection will modify races, just as Voltaire firmly held that incantations, together with sufficient arsenic, would destroy flocks of sheep? Is it not a significant fact that the alleged instances of use-inheritance so often prove to be self-conflicting in their details?

For satisfactory proof of the prevalence of a law of use-inheritance we require normal instances where selection is clearly inadequate to produce the change, or where it is scarcely allowed time or opportunity to act, as in the immediate offspring of the modified individual. Of the first kind of cases there seems to be a plentiful lack. Of the latter kind, according to Darwin, there appears to be none—a circumstance which contrasts strangely and suspiciously with the many decisive cases in which variation from unknown causes has been inherited most strikingly in the immediate offspring. It must be expected, indeed, that among these innumerable cases some will accidentally mimic the alleged effects of use-inheritance.

If Darwin had felt certain that the effects of habit or use tended in any marked degree to be conveyed directly and cumulatively to succeeding generations, he could hardly have given us such cautious, half-hearted encouragement of good habits as the following:—"It is not improbable that after long practice virtuous tendencies may be inherited." "Habits, moreover followed during many generations probably tend to be inherited."[53] This is probable, independently of use-inheritance. The "many generations" specified or implied, will allow time for the play of selective as well as of cumulatively-educative influences. There must apparently be a constitutional or inheritable predisposition or fitness for the habits spoken of, which otherwise would scarcely be continued for many generations, except by the favourably-varying branches of a family: which again is selection rather than use-inheritance.

Where is the necessity for even the remains of the Lamarckian doctrine of inherited habit? Seeing how powerful the general principle of selection has shown itself in cases where use-inheritance could have given no aid or must even have offered its most strenuous opposition, why should it not equally be able to develop used organs or repress disused organs or faculties without the assistance of a relatively weak ally? Selection evolved the remarkable protective coverings of the armadillo, turtle, crocodile, porcupine, hedgehog, &c.; it formed alike the rose and its thorn, the nut and its shell; it developed the peacock's tail and the deer's antlers, the protective mimicry of various insects and butterflies, and the wonderful instincts of the white ants; it gave the serpent its deadly poison and the violet its grateful odour; it painted the gorgeous plumage of the Impeyan pheasant and the beautiful colours and decorations of countless birds and insects and flowers. These, and a thousand other achievements, it has evidently accomplished without the help of use-inheritance. Why should it be thought incapable of reducing a pigeon's wing or enlarging a duck's leg? Why should it be credited with the help of an officious ally in effecting comparatively slight changes, when great and striking modifications are effected without any such aid?

FOOTNOTES:

[15] Weismann's Essays on Heredity, &c. Clarendon Press, 1889.

[16] Life and Letters, i. p. 16. Darwin's reverence for his father "was boundless and most touching. He would have wished to judge everything else in the world dispassionately, but anything his father had said was received with almost implicit faith; ... he hoped none of his sons would ever believe anything because he said it, unless they were themselves convinced of its truth—a feeling in striking contrast with his own manner of faith" (Life and Letters, i. pp. 10, 11).

[17] Ibid., i. p. 38.

[18] Life and Letters, ii. p. 14.

[19] Origin of Species, pp. 117, 118.

[20] Ibid., p. 180.

[21] Contemporary Review, December, 1875, pp. 89, 93.

[22] Variation of Animals and Plants under Domestication, i. 292.

[23] Variation of Animals and Plants under Domestication, i. 299-301.

[24] To keep pace with this lateral increase in weight, the leg-bones should have lengthened considerably so that their total deficiency in proportional length is 17 per cent.,—a changed proportion which being linear is more excessive than the increase of weight by 28 per cent. So marked is the effect of the combined thickening and shortening that in the Aylesbury breed—which is the most typically representative one—the leg-bones have become 70 per cent. heavier than they should be if their thickness had continued to be proportional to their length.

[25] This excessive thickening under disuse appears to be due partly to a positive lateral enlargement or increase of proportional weight of about 7-1/2 per cent., and partly to a shortening of about 15 per cent. Carefully calculated, the reduction of the weight of the wing-bones in this breed is only 8.3 per cent. relatively to the whole skeleton, or only 5 per cent. relatively to the skeleton minus legs and wings. The latter method is the more correct, since the excessive weight of the leg-bones increases the weight of the skeleton more than the diminished weight of the wing-bones reduces it.

[26] Variation of Animals and Plants under Domestication, i. 284.

[27] Variation of Animals and Plants under Domestication, i. 184, 185.

[28] Ibid., i. 144, 145.

[29] Ibid., i. 185.

[30] Variation of Animals and Plants under Domestication, i. 175.

[31] Variation of Animals and Plants under Domestication, i. 184. I suspect that Darwin was in poor health when he wrote this page. He nods at least four times in it. Twice he speaks of "twelve" breeds where he obviously should have said eleven.

[32] If a prominent breast is admired and selected by fanciers, the sternum might shorten in assuming a more forward and vertical position. If the shortening of the sternum is entirely due to disuse, it seems strange that Darwin has not noticed any similar shortening in the sternum of the duck. But selection has not tended to make the duck elegant, or "pigeon-breasted"; it has enlarged the abdominal sack instead, besides allowing the addition of an extra rib in various cases.

[33] Variation of Animals and Plants under Domestication, 144, 175.

[34] Variation of Animals and Plants under Domestication, i. 179.

[35] In the six largest breeds the shortening of the sternum is nearly twice as great as in the three smaller breeds which remain nearest the rock-pigeon in size. We can hardly suppose that use-inheritance especially affects the eight breeds that have varied most in size. If we exclude these, there is only a total shortening of 7 per cent. to be accounted for.

[36] Variation of Animals and Plants under Domestication, i. 183, 186.

[37] Variation of Animals and Plants under Domestication, i. 130, 135; ii. 288.

[38] Encyclopaedia Britannica, article "Zoology."

[39] Variation of Animals and Plants under Domestication, ii. 367.

[40] Variation of Animals and Plants under Domestication, ii. 367. Why then does the cheetah inherit ancestral habits so inadequately that it is useless for the chase unless it has first learned to hunt for itself before being captured? (ii. 133).

[41] Descent of Man, p. 33.

[42] Origin of Species, pp. 210, 211.

[43] E. S. Delamer on Pigeons and Rabbits, pp. 132, 103. For other points referred to, see pages 133, 102, 100, 95, 131.

[44] Origin of Species, pp. 188, 110; Descent of Man, pp. 32-35; Variation of Animals and Plants under Domestication, ii. 289, 293. Use or disuse during lifetime of course co-operates, and in some cases, as in that of the canoe Indians, may be the principal or even perhaps the sole cause of the change.

[45] For the importance of panmixia as invalidating Darwin's strongest evidence for use-inheritance—namely, that drawn from the effects of disuse in highly-fed domestic animals where there is supposed to be no economy of growth—see Professor Romanes on Panmixia, Nature, April 3, 1890.

[46] Descent of Man, p. 33.

[47] Descent of Man, p. 33.

[48] Variation of Animals and Plants under Domestication, i., 453.

[49] Descent of Man, p. 33.

[50] Descent of Man, p. 33.

[51] Wallace shows that the changes in our domestic animals, if spread over the thousands of years since the animals were first tamed, must be extremely insignificant in each generation, and he concludes that such infinitesimal effects of use and disuse would be swallowed up by the far greater effects of variation and selection (Darwinism, p. 436). Professor Romanes has replied to him in the Contemporary Review (August 1889), showing that this is no disproof of the existence of the minor factor, inasmuch as slight changes in each generation need not necessarily be matters of life and death to the individual, although their cumulative development by use-inheritance might eventually become of much service. But selection would favour spontaneous variations of a similarly serviceable character. The slightest tendency to eliminate the extreme variations in either direction would proportionally modify the average in a breed. Use-inheritance appears to be so relatively weak a factor that probably neither proof nor disproof of its existence can ever be given, owing to the practical impossibility of disentangling its effects (if any) from the effects of admittedly far more powerful factors which often act in unsuspected ways. Thus wild ducklings, which can easily be reared by themselves, invariably "die off" if reared with tame ones (Variation, &c., i. 292, ii. 219). They cannot get their fair share in the competition for food, and are completely eliminated. Professor Romanes fully acknowledges that there is the "gravest possible doubt" as to the transmission of the effects of disuse (Letter on Panmixia, Nature, March 13, 1890).

[52] Variation of Animals and Plants under Domestication, ii. 287-289.

[53] Descent of Man, pp. 612, 131.



INHERITED INJURIES.

INHERITED MUTILATIONS.

The almost universal non-inheritance of mutilations seems to me a far more valid argument against a general law of modification-inheritance than the few doubtful or abnormal cases of such inheritance can furnish in its favour. No inherited effect has been produced by the docking of horses' tails for many generations, or by a well-known mutilation which has been practised by the Hebrew race from time immemorial. As lost or mutilated parts are reproduced in offspring independently of the existence of those parts in the parent, there is the less reason to suppose that the particular condition of parental parts transmits itself, or tends to transmit itself, to the offspring. So unsatisfactory is the argument derivable from inherited mutilations that Mr. Spencer does not mention them at all, and Darwin has to attribute them to a special cause which is independent of any general theory of use-inheritance.[54]

Darwin's most striking case—and to my mind the only case of any importance—is that of Brown-Sequard's epileptic guinea-pigs, which inherited the mutilated condition of parents who had gnawed off their own gangrenous toes when anaesthetic through the sciatic nerve having been divided.[55] Darwin also mentions a cow that lost a horn by accident, followed by suppuration, and subsequently produced three calves which had on the same side of the head, instead of a horn, a bony lump attached merely to the skin. Such cases may seem to prove that mutilation associated with morbid action is occasionally inherited or repeated with a promptitude and thoroughness that contrast most strikingly with the imperceptible nature of the immediate inheritance of the effects of use and disuse; but they by no means prove that mutilation in general is inheritable, and they are absolutely no proof whatever of a normal and non-pathological tendency to the inheritance of acquired characters. Those who accept Darwin's special explanation of the supposed inheritance of mutilations, ought to notice that his explanation applies equally well under a theory which is strongly adverse to use-inheritance—namely, Galton's idea of the sterilization and complete "using up" of otherwise reproductive matter in the growth and maintenance of the personal structure.

Darwin's explanation of inherited mutilations—which, as he notes, occur "especially or perhaps exclusively" when the injury has been followed by disease[56]—is that all the representative gemmules which would develop or repair or reproduce the injured part are attracted to the diseased surface during the reparative process and are there destroyed by the morbid action.[57] Hence they cannot reproduce the part in offspring. This explanation by no means implies that mutilation would usually affect the offspring. On the contrary, in all ordinary cases of mutilation the purely atavistic elements or gemmules would be set free from any modifying influence of the non-existent or mutilated part. The gemmules—as in Galton's theory of heredity and with neuter insects—might be perfectly independent of pangenesis and the normal inheritance of acquired characters. Such self-multiplying gemmules without pangenesis would enable us to understand both the excessive weakness or non-existence of normal use-inheritance, and the excessive strength and abruptness of the effect of their partial destruction under special pathological conditions.

The series of epileptic phenomena that can be excited by tickling a certain part of the cheek and neck of the adult guinea-pig during the growth and rejoining of the ends of the severed nerve, are said to be repeated with striking accuracy of detail in the young who inherit mutilated toes; but as epilepsy is often due to some one exciting cause or morbid condition, the single transmission of a highly morbid condition of the system might easily reproduce the whole chain of consequences and might also have caused the loss of toes.

The particulars of the guinea-pig cases are very inadequately recorded,[58] but the results are so anomalous[59] that Brown-Sequard's own conclusion is that the epilepsy and the inherited injuries are not directly transmitted, but that "what is transmitted is the morbid state of the nervous system." He thinks that the missing toes may "possibly" be exceptions to this conclusion, "but the other facts only imply the transmission of a morbid state of the sympathetic or sciatic nerve or of a part of the medulla oblongata." Until we can tell what is transmitted, we are not in a position to determine whether there is any true inheritance or only an exaggerated simulation of it under peculiar circumstances. When the actual observers believe that the mutilations and epilepsy are not the cause of their own repetition, and when these observers guard themselves by such phrases as, "if any conclusion can at present be drawn from those facts," we who have only incomplete reports to guide us may well be excused if we preserve an even more pronounced attitude of caution and reserve.[60] The morbid state of the system may be wholly due to general injury of the germs rather than to specific inheritance.

Weismann suggests that the morbid condition of the nervous system may be due to some infection such as might arise from microbes, which find a home in the mutilated and disordered nervous system in the parent, and subsequently transmit themselves to the offspring through the reproductive elements, as the infections of various diseases appear to do—the muscardine silkworm disease in particular being known to be conveyed to offspring in this manner.

But whether we can discover the true explanation or not, inherited mutilations can hardly be accounted for as the result of a general tendency to inherit acquired modifications. How could a factor which seems to be totally inoperative in cases of ordinary mutilation, and only infinitesimally operative in transmitting the normal effects of use and disuse, suddenly become so powerful as to completely overthrow atavism, and its own tendency to transmit the non-mutilated type of one of the parents and of the non-mutilated type presented by the injured parent in earlier life? Does not so striking and abrupt an intensification of its usually insignificant power demand an explanation widely different from that which might account for the extremely slow and slight inheritance of the normal effects of use and disuse? Surely it would be better to suspend one's judgment as to the true explanation of highly exceptional and purely pathological cases rather than resort to an hypothesis that creates more difficulties than it solves.

THE MOTMOT'S TAIL.

The narrowing of the long central tail feathers of the motmot is attributed to the inherited effects of habitual mutilation (Descent of Man, pp. 384, 603). But in the specimens at South Kensington[61] the narrowness extends upwards much beyond the habitually denuded part, and the broadened end is the broadest part of the whole feather. If the inherited effect of an inch or two of denudation extends from three to six inches upwards, why has it not also extended two inches downwards so as to narrow the broadened end? The narrowness seems to be a mainly relative or negative effect produced by the broadening out of a long tapering feather at its end under the influence of sexual selection. Several other birds have similarly narrowed or spoon-shaped feathers and do not bite them. Is it not more feasible to suppose that this attractive peculiarity first suggested its artificial intensification, than to suppose that the bird began nibbling without any definite cause? Sexual selection would then encourage the habit. Anyhow, it is as impossible to show that the mutilation preceded the narrowing as it is to show that tonsure preceded baldness.

OTHER INHERITED INJURIES MENTIONED BY DARWIN.

Darwin quotes some cases from Dr. Prosper Lucas's "long" but weak and unsatisfactory "list of inherited injuries."[62] But Lucas was somewhat credulous. One of his cases is that many girls were born in London without mammae through the injurious effect of certain corsets on the mothers. He also gives a long account of a Jew who could read through the thick covers of a book, and whose son inherited this "hyperaesthesia" of the sense of sight in a still more remarkable degree (i. 113-119). Evidently Lucas's cases cannot be accepted without some amount of reserve.

The cases of the three calves which inherited the one-horned condition of the cow, the two sons who inherited a father's crooked finger, and the two sons who were microphthalmic on the same side as their father had lost an eye, may be due to mere coincidence; or an inherited constitutional tendency or liability might lead to somewhat similar results in parent and offspring[63]—just as the tendency to certain fatal diseases or to suicide may produce similar results in father and son, although the artificially-produced hanging or apoplexy obviously cannot be directly transmitted. That more than one of the offspring was affected does not render the chances against coincidence "almost infinitely great," as Darwin mistakenly supposes. It "frequently occurs" that a man's sons or daughters may all exhibit either a latent or a newly-developed congenital peculiarity previously unknown;[64] and the coincidence may merely be that one of the parents accidentally suffered a similar kind of injury—a kind of coincidence which must of course occasionally occur, and which may have been partly caused by a latent tendency. The chances against coincidence are indeed great, but the cases appear to be correspondingly rare.

Darwin acknowledges that many supposed instances of inherited mutilation may be due to coincidence; and there is apparently no more reason for attributing inherited scars, &c., to any special form of heredity than to the effect of the mother's imagination on the unborn babe—a popular but fallacious belief in corroboration of which far more alleged instances could be collected than of the inheritance of injuries.

As an instance of the coincidences that occur, I may mention that a friend of mine has a daughter who was born with a small hole in one ear, just as if it were already pierced for the earring which she has since worn in it. I suppose, however, that no one will venture to claim this as an instance of the inheritance of a mutilation practised by female ancestors, especially as such holes are not altogether unknown or inexplicable, though very rarely occurring low down in the lobe of the ear.[65]

Many cases are known of the inheritance of mutilations or malformations arising congenitally from some abrupt variation in the reproductive elements. In such cases as the one-eared rabbits, the two-legged pigs, the three-legged dogs, the one-horned stags, hornless bulls, earless rabbits, lop-eared rabbits, tailless dogs, &c., if the father or the mother or the embryo had suffered from some accident or disease which might plausibly have been assigned as the cause of the original malformation, these transmitted defects would readily be cited as instances of the inheritance of an accidentally-produced modification.

The inheritance of exostoses on horses' legs may be the inheritance of a constitutional tendency rather than of the effect of the parents' hard travelling. Horses congenitally liable to such formations would transmit the liability,[66] and this might readily be mistaken for inheritance of the results of the liability. An apparent increase in this liability might arise from greater attention being now paid to it, or from increased use of harder roads; or a real increase might be due to panmixia and some obscure forms of correlation.

QUASI-INHERITANCE.

Of course artificially-caused ill-health or weakness in parents will tend in a general way to injure the offspring. But deterioration thus caused is only a form of quasi-inheritance, as I should prefer to call it. Semi-starvation in a new-born babe is not truly inherited from its half-starved mother, but is the direct result of insufficient nourishment. The general welfare of germs—as of parasites—is necessarily bound up with that of the organism which feeds and shelters them, but this is not heredity, and is quite irrelevant to the question whether particular modifications are transmitted or not.

Another form of quasi-inheritance is seen in the communication of certain infections to offspring. Not being transmitted by the action of the organism so much as in defiance of it, such diseases are not truly hereditary, though for convenience' sake they are usually so described.

A perversion or prevention of true inheritance is also seen in the action of alcohol, or excessive overwork, or any other cause which by originating morbid conditions in individuals may also injure the reproductive elements.

These forms of quasi-inheritance are, of course, highly important so far as the improvement of the race is concerned. So, too, is the fact that improved or deteriorated habits and thoughts are transmitted by personal teaching and influence and are cumulative in their effect. But all this must not be confounded with the inheritance of acquired characters. Cases of quasi-inheritance may perhaps be most readily distinguished from cases of true inheritance by the time test. When a modification acquired in adult life is promptly communicated to the child in early life or from birth, it may rightly be suspected that the inheritance, like that of money or title, is not truly congenital, but is extraneous or even anti-congenital in its nature. Judged by such a standard, the inherited injuries in Brown-Sequard's guinea-pigs are only exceptional cases of quasi-inheritance, and are not necessarily indicative of any general rule affecting true inheritance.

FOOTNOTES:

[54] A very able anatomist of my acquaintance denies the inheritance of mutilations and injuries, although he strongly believes in the inheritance of the effects of use and disuse.

[55] Variation of Animals and Plants under Domestication, i. 467-469. Lost toes were only seen by Dr. Dupuy in three young out of two hundred. Obersteiner found that most of the offspring of his epileptic guinea-pigs were injuriously affected, being weakly, small, paralysed in one or more limbs, and so forth. Only two were epileptic, and both were weakly and died early (Weismann's Essays, p. 311). A morbid condition of the spinal cord might affect the hind limbs especially (as in paraplegia) and might occasionally cause loss of toes in the embryo by preventing development or by ulceration. Brown-Sequard does not say that the defective feet were on the same side as in the parents (Lancet, Jan., 1875, pp. 7, 8).

[56] Variation of Animals and Plants under Domestication, ii. 57.

[57] Ibid., ii. 392. Perhaps it might be better to suppose that the best gemmules were sacrificed in repairing the injured nerve, and hence only inferior substitutes were left to take their place, and could only imperfectly reproduce the injured part of the nervous system in offspring.

[58] Hence perhaps Mr. Spencer's error in representing the epileptic liability as permanent and as coming on after healing (Factors of Organic Evolution, p. 27).

[59] It is not claimed that the imperfect foot was on the same side of the body as in the parent, and where parents had lost all the toes of a foot, or the whole foot, the few offspring affected usually had lost only two toes out of the three, or only a part of one or two or three toes. Sometimes the offspring had toes missing on both hind feet, although the parent was only affected in one. One diseased ear and eye in the parent was "generally" or "always" succeeded by two equally affected ears and eyes in the offspring (cf. Pop. Science Monthly, New York, xi. 334). The important law of inheritance at corresponding periods was also set aside. Gangrene or inflammation commenced in both ears and both eyes soon after birth (pointing possibly to infection of some kind); the epileptic period commenced "perhaps two months or more after birth," while the loss of toes had occurred before birth. In no case, as Weismann points out, is the original mutilation of the nervous system ever transmitted. Even where an extirpated ganglion was never regenerated in the parent, the offspring always regained the part in an apparently perfect condition. On the whole the conflicting results ought to be as puzzling to those who may attribute them to a universal tendency to inherit the exact condition of parents as they are to those who, like myself, are sceptical as to the existence of such a law or tendency.

[60] The various results need to be fully and impartially recorded, and they should also be well tested and confirmed in proportion as they appear improbable and contrary to general experience. Professor Romanes has been carrying out the necessary experiments for some time past.

[61] Natural History Museum, central hall, third recess on the left.

[62] Traite de l'Heredite, ii. 489; Variation of Animals and Plants under Domestication, i. 469. If injuries are inherited, why has the repeated rupture of the hymen produced no inherited effect?

[63] Compare the three cases of crooked fingers given in Variation of Animals and Plants under Domestication, ii. 55, 240.

[64] Ibid., i. 460. Thus, where two brothers married two sisters all the seven children were perfect albinos, although none of the parents or their relatives were albinos. In another case the nine children of two sound parents were all born blind (ii. 322).

[65] See pp. 179-182, Evolution and Disease, by J. Bland Sutton, to whom and to our mutual friend Dr. D. Thurston I am indebted for information on various points.

[66] Variation of Animals and Plants under Domestication, ii. 290; i. 454.



MISCELLANEOUS CONSIDERATIONS.

TRUE RELATION OF PARENTS AND OFFSPRING.

It is difficult to entirely free ourselves from the flattering and almost universal idea that parents are true originators or creators of copies of themselves. But the main truth, if not the whole truth, is that they are merely the transmitters of types of which they and their offspring are alike more or less similarly moulded resultants. A parent is a trustee. He transmits, not himself and his own modifications, but the stock, the type, the representative elements, of which he is a product and a custodian in one. It seems probable that he has no more definite or "particulate" influence over the reproductive elements within him than a mother over the embryo or a vessel over its cargo. Parent and offspring are like successive copies of books printed from the same "type." A battered letter in the "type" will display its effects in both earlier and later copies alike, but a purely extraneous or acquired flaw in the first copy is not necessarily repeated in subsequent copies. Unlike printer's type, however, the material source of heredity is of a fluctuating nature, consisting of competing elements derived from two parents and from innumerable ancestors.

Galton compares parent and child to successive pendants on the same chain. Weismann likens them to successive offshoots thrown up by a long underground root or sucker. Such comparisons indicate the improbability of acquired modifications being transmitted to offspring.

That parts are developed in offspring independently of those parts in parents is clear. Mutilated parents transmit parts which they do not possess. The offspring of young parents cannot inherit the later stages of life from parents who have not passed through them. Cases of remote reversion or atavism show that ancestral peculiarities can transmit themselves in a latent or undeveloped condition for hundreds or thousands of generations. Many obvious facts compelled Darwin to suppose that vast numbers of the reproductive gemmules in an individual are not thrown off by his own cells, but are the self-multiplying progeny of ancestral gemmules. Galton restricts the production of gemmules by the personal structure to a few exceptional cases, and would evidently like to dispense with pangenesis altogether, if he could only be sure that acquired characters are never inherited. Weismann entirely rejects pangenesis and the inheritance of acquired characters. This enables him to explain heredity by his theory of the "Continuity of the Germ-plasm."[67] Parent and offspring are alike successive products or offshoots of this persistent germ-substance, which obviously would not be correspondingly affected by modifications of parts in parents, and so would render the transmission of acquired characters impossible.

INVERSE INHERITANCE.

Mr. Galton contends that the reproductive elements become sterile when used in forming and maintaining the individual, and that only a small proportion of them are so used.[68] He holds that the next generation will be formed entirely, or almost entirely, from the residue of undeveloped germs, which, not having been employed in the structure and work of the individual, have been free to multiply and form the reproductive elements whence future individuals are derived. Hence the singular inferiority not infrequently displayed by the children of men of extraordinary genius, especially where the ancestry has been only of a mediocre ability. The valuable germs have been used up in the individual, and rendered sterile in the structure of his person. Hence, too, the "strong tendency to deterioration in the transmission of every exceptionally gifted race." Mr. Galton's hypothesis "explains the fact of certain diseases skipping one or more generations," and it "agrees singularly well with many classes of fact;" and it is strongly opposed to the theory of use-inheritance. The elements which are used die almost universally without germ progeny: the germs which are not used are the great source of posterity. Hence, when the germs or gemmules which achieve development are either better or worse than the residue, the qualities transmitted to offspring will be of an inverse character. If brain-work attracts, develops and sterilizes the best gemmules, the ultimate effect of education on the intellect of posterity may differ from its immediate effect.

EARLY ORIGIN OF THE OVA.

As the ova are formed at as early a period as the rest of the maternal structure, Galton notices that it seems improbable that they would be correspondingly affected by subsequent modifications of parental structure. Of course it is not certain that this is a valid argument. We know that the paternal half of the reproductive elements does not enter the ovum till a comparatively late stage in its history, and it is quite possible that maternal elements or gemmules may also enter the ovum from without. If reproductive elements were confined to one special part or organ, we should be unable to explain the reproduction of lost limbs in salamanders, and the persistent effect of intercrossing on subsequent issue by the same mother, and the propagation of plants from shoots, or of the begonia from minute fragments of leaves, or the development of small pieces of water-worms into complete animals.

MARKED EFFECTS OF USE AND DISUSE ON THE INDIVIDUAL.

These are, to some extent, an argument against the cumulative inheritance of such effects. When a nerve atrophies from disuse, or a duct shrivels, or bone is absorbed, or a muscle becomes small or flabby, it proves, so far, that the average effect of use through enormous ages is not transmitted. When the fibula of a dog's leg thickens by 400 per cent. to a size "equal to or greater than" that of the removed tibia which previously did the work,[69] it shows that in spite of disuse for countless generations, the "almost filiform" bone has retained a potentiality of development which is fully equal to that possessed by the larger one which has been constantly used. When, after being reared on the ailanthus, the caterpillars of the Bombyx hesperus die of hunger rather than return to their natural food, the inherited effect of ancestral habit does not seem to be particularly strong. Neither is there any strongly-inherited effect of long-continued ancestral wildness in many animals which are easily tamed.

WOULD NATURAL SELECTION FAVOUR USE-INHERITANCE?

If use-inheritance is really one of the factors of evolution, it is certainly a subordinate one, and an utterly helpless one, whenever it comes into conflict with the great ruling principle of Selection. Would this dominant cause of evolution have favoured a tendency to use-inheritance if such had appeared, or would it have discouraged and destroyed it? We have already seen that use-inheritance is unnecessary, since natural selection will be far more effective in bringing about advantageous modifications; and if it can be shown that use-inheritance would often be an evil, it then becomes probable that on the whole natural selection would more strongly discourage and eliminate it as a hostile factor than it might occasionally favour such a tendency as a totally unnecessary aid.

USE-INHERITANCE AN EVIL.

Use-inheritance would crudely and indiscriminately proportion parts to actual work done—or rather to the varying nourishment and growth resulting from a multiplicity of causes—and this in its various details would often conflict most seriously with the real necessities of the case, such as occasional passive strength, or appropriate shape, lightness and general adaptation. If its accumulated effects were not corrected by natural or sexual selection, horns and antlers would disappear in favour of enlarged hoofs. The elephant's tusks would become smaller than its teeth. Men would have callosities for sitting on, like certain monkeys, and huge corns or hoofs for walking on. Bones would often be modified disastrously. Thus the condyle of the human jaw would become larger than the body of the jaw, because as the fulcrum of the lever it receives more pressure. Some organs (like the heart, which is always at work) would become inconveniently or unnecessarily large. Other absolutely indispensable organs, which are comparatively passive or are very seldom used, would dwindle until their weakness caused the ruin of the individual or the extinction of the species. In eliminating various evil results of use-inheritance, natural selection would be eliminating use-inheritance itself. The displacement of Lamarck's theory by Darwin's shows that the effects of use-inheritance often differ from those required by natural selection; and it is clear that the latter factor must at least have reduced use-inheritance to the very minor position of comparative feebleness and harmlessness assigned to it by Darwin.

Use-inheritance would be ruinous through causing unequal variation in co-operative parts—of which Mr. Spencer may accept his own instances of the jaws and teeth, and the cave-crab's lost eyes and persistent eye-stalks, as typical examples. That the variation would be unequal seems almost self-evident from the varying rapidity and extent of the effects of use and disuse on different tissues and on different parts of the general structure. The optic nerve may atrophy in a few months from disuse consequent on the loss of the eye. Some of the bones of the rudimentary hind legs of the whale are still in existence after disuse for an enormous period. Evidently use-inheritance could not equally modify the turtle and its shell, or the brain and its skull; and in minor matters there would be the same incongruity of effect. Thus, if the molar teeth lengthened from extra use the incisors could not meet. Unequal and indiscriminate variation would throw the machinery of the organism out of gear in innumerable ways.

Use-inheritance would perpetuate various evils. We are taught, for instance, that it perpetuates short-sight, inferior senses, epilepsy, insanity, nervous disorders, and so forth. It would apparently transmit the evil effects of over-exertion, disuse, hardship, exposure, disease and accident, as well as the defects of age or immaturity.

Would it not be better on the whole if each individual took a fresh start as far as possible on the advantageous typical lines laid down by natural selection? Through the long stages of evolution from primaeval protoplasm upwards, such species as were least affected by use-inheritance would be most free to develop necessary but seldom-used organs, protective coverings such as shells or skulls, and natural weapons, defences, ornaments, special adaptations, and so forth; and this would be an advantage—for survival would obviously depend on the importance of a structure or faculty in deciding the struggle for existence and reproduction, and not on the total amount of its using or nourishment. If natural selection had on the whole favoured this officious ally and frequent enemy, surely we should find better evidence of its existence.

Without laying undue stress upon the evil effects of use-inheritance, a careful examination of them in detail may at least serve to counter-balance the optimistic a priori arguments for belief in that plausible but unproven factor of evolution.

The benefits derivable from use-inheritance are largely illusory. The effects of use, indeed, are generally beneficial up to a certain point; for natural selection has sanctioned or evolved organs which possess the property or potentiality of developing to the right extent under the stimulus of use or nourishment. But use-inheritance would cumulatively alter this individual adaptability, and would tend to fix the size of organs by the average amount of ancestral use or disuse rather than by the actual requirements of the individual. Of course under changed conditions involving increased or lessened use of parts it might become advantageous; but even here it may prove a decided hindrance to adaptive evolution in some respects as well as an unnecessary aid in others. Thus in the case of animals becoming heavier, or walking more, it would lengthen the legs although natural selection might require them to be shortened. In the Aylesbury duck and the Call duck, if use-inheritance has increased the dimensions of the bones and tendons of the leg, natural selection has had to counteract this increase so far as length is concerned, and to effect 8 per cent. of shortening besides. If use-inheritance thickens bones without proportionally lengthening them, it would hinder rather than help the evolution of such structures as the long light wings of birds, or the long legs and neck of the giraffe or crane.

VARIED EFFECTS OF USE AND DISUSE.

The changes which we somewhat roughly and empirically group together as the effects of "use and disuse" are of widely diverse character. Thus bone, as the physiological fact, thickens under alternations of pressure (and the consequent increased flow of nourishment), but atrophies under a steadily continued pressure; so that if the use of a bone involved continuous pressure, the effect of such use would be a partial or total absorption of that bone. Darwin shows that bone lengthens as well as thickens from carrying a greater weight, while tension (as seen in sailors' arms, which are used in pulling) appears to have an equally marked effect in shortening bones (Descent of Man, p. 32). Thus different kinds of use may produce opposite results. The cumulative inheritance of such effects would often be mischievous. The limbs of the sloth and the prehensile tail of the spider monkey would continually grow shorter, while the legs of the evolving elephant or rhinoceros might lengthen to an undesirable extent. Such cumulative tendencies of use-inheritance, if they exist, are obviously well kept under by natural selection.

Although the ultimate effect of use is generally growth or enlargement through increased flow of blood, the first effect usually is a loss of substance, and a consequent diminution of size and strength. When the loss exceeds the growth, use will diminish or deteriorate the part used, while disuse would enlarge or perfect it. Teeth, claws, nails, skin, hair, hoofs, feathers, &c., may thus be worn away faster than they can renew themselves. But this wearing away usually stimulates the repairing process, and so increases the rate of growth; that is, it will increase the size produced, if not the size retained. Which effect of use does use-inheritance transmit in such cases—the increased rate of growth, or the dilapidation of the worn-out parts? We can hardly suppose that both these effects of use will be inherited. Would shaving destroy the beard in time or strengthen it? Will the continued shearing of sheep increase or lessen the growth of wool? What will be the ultimate effect of plucking geese's quills, and of the eider duck's abstraction of the down from her breast? If the mutilated parts grow stronger or more abundantly, why were the motmot's feathers alleged to be narrowed by the inherited effects of ancestral nibbling?

The "use" or "work" or "function" of muscles, nerves, bones, teeth, skin, tendon, glands, ducts, eyes, blood corpuscles, cilia, and the other constituents of the organism, is as widely different as the various parts are from each other, and the effects of their use or disuse are equally varied and complicated.

USE-INHERITANCE IMPLIES PANGENESIS.

How could the transmission of these varied effects to offspring be accounted for? Is it possible to believe, with Mr. Spencer, that the effects of use and disuse on the parts of the personal structure are simultaneously registered in corresponding impressions on the seminal germs? Must we not feel, with Darwin apparently,[70] that the only intelligible explanation of use-inheritance is the hypothesis of Pangenesis, according to which each modified cell, or physiological unit, throws off similarly-modified gemmules or parts of itself, which ultimately reproduce the change in offspring? If we reject pangenesis, it becomes difficult to see how use-inheritance can be possible.

PANGENESIS IMPROBABLE.

The more important and best-known phenomena of heredity do not require any such hypothesis, and leading facts (such as atavism, transmission of lost parts, and the general non-transmission of acquired characters) are so adverse to it that Darwin has to concede that many of the reproductive gemmules are atavistic, and that by continuous self-multiplication they may preserve a practical "continuity of germ-substance," as Weismann would term it. The idea that the relationship of offspring to parent is one of direct descent is, as Galton tells us, "wholly untenable"; and the only reason he admits some supplementary traces of pangenesis into his "Theory of Heredity,"[71] is that he may thus account for the more or less questionable cases of the transmission of acquired characters. But there appears to be no necessity even for this concession. We ought therefore to dispense with the useless and gratuitous hypothesis that cells multiply by throwing off minute self-multiplying gemmules, as well as by the well-known method of self-division. If pangenesis occurs, the transmission of acquired characters ought to be a prominent fact. The size, strength, health and other good or evil qualities of the cells could hardly fail to exercise a marked and corresponding effect upon the size and quality of the reproductive gemmules thrown off by those cells. The direct evidence tends to show that these free gemmules do not exist. Transfusion of blood has failed to affect inheritance in the slightest degree. Pangenesis, with its attraction of gemmules from all parts of the body into the germ-cells, and the free circulation of gemmules in the offspring till they hit upon or are attracted by the particular cell or cells, with which alone they can readily unite, seems a less feasible theory and less in conformity with the whole of the facts than an hypothesis of germ-continuity which supposes that the development of the germ-plasm and of the successive self-dividing cells of the body proceeds from within. Darwin's keen analogy of the fertilization of plants by pollen renders development from without conceivable, but as there are no insects to convey gemmules to their destination, each kind of gemmule would have to be exceedingly numerous and easily attracted from amongst an inconceivable number of other gemmules. Arguments against pangenesis can also be drawn from the case of neuter insects—a fact which seems to have escaped Darwin's notice, although he had seen how strongly that case was opposed to the doctrine which is the essential basis of the theory of pangenesis.

SPENCER'S EXPLANATION OF USE-INHERITANCE.

Mr. Spencer's explanation of the inheritance of the effects of use and disuse (p. 36) is that "while generating a modified consensus of functions and of structures, the activities are at the same time impressing this modified consensus on the sperm-cells and germ-cells whence future individuals are to be produced"—a proposition which reads more like metaphysics than science. Difficult to understand or believe in ordinary instances, such consensus-inheritance seems impossible in cases like that of the hive-bee. Can we suppose that the consensus of the activities of the working bee impresses itself on the sperm-cells of the drones and on the germ-cells of the carefully secluded queen? Buechner thinks so, for he says: "Although the queens and drones do not now work, yet the capacities inherited from earlier times still remain to them, especially to the former, and are kept alive and fresh by the impressions constantly made upon them during life, and they are thus in a position to transmit them to posterity." Surely it is better to abandon a cherished theory than to be compelled to defend it by explanations which are as inconsistent as they are inadequate. New capacities are developed as well as old ones kept fresh. The massacre or expulsion of the drones would have to impress itself on the germ-cells of an onlooking queen, and the imprisonment of the queen on the sperm-cells of the drones—and in such a way, moreover, as to be afterwards developed into action in the neuters only. And use-inheritance all the while is being thoroughly overpowered by impression-inheritance—by the full transmission of that which is merely seen in others! If such a law prevails, one may feel cold because an ancestor thought of the frosty Caucasus. None of this absurdity would arise if it were clearly seen that a parent is only a trustee—that transmission and development are perfectly distinct—that parental modifications are irrelevant to those transmitted to offspring.

FOOTNOTES:

[67] Essays on Heredity, p. 104. Weismann's theory is clear, simple and convenient, but incomplete; for, unlike Darwin's theory of pangenesis, it scarcely attempts any real explanation of the extremely complex potentialities possessed by the reproductive elements. Perhaps we might retain Darwin's self-multiplying gemmules without supposing them to be thrown off by the cells, which will no longer be credited with two modes of multiplication. These minute germs or gemmules may have been evolved by natural selection playing upon the sample germs that achieve development; and they may exist either separately, or (preferably but perhaps not invariably) in aggregates to form Weismann's germ-plasm.

[68] Contemporary Review, Dec., 1875, p. 88.

[69] Variation of Animals and Plants under Domestication, ii. 286.

[70] Variation of Animals and Plants under Domestication, ii. 388, 398, 367; Life and Letters, iii. 44.

[71] Contemporary Review, Dec., 1875, pp. 94, 95.



CONCLUSIONS.

USE-INHERITANCE DISCREDITED AS UNNECESSARY, UNPROVEN, AND IMPROBABLE.

General experience teaches that acquired characters are not usually inherited; and investigation shows that the apparent exceptions to this great rule are probably fallacious. Even the alleged instances of use-inheritance culled by such great and judicious selectors as Darwin and Spencer break down upon examination; for they can be better explained without use-inheritance than with it. On the other hand, the adverse facts and considerations are almost strong enough to prove the actual non-existence of such a law or tendency. There is no need to undertake the apparently impossible task of demonstrating an absolute negative. It will be enough to ask that the Lamarckian factor of use-inheritance shall be removed from the category of accredited factors of evolution to that of unnecessary and improbable hypotheses. The main explanation or source of the fallacy may be found in the fact that natural selection frequently imitates some of the more obvious effects of use and disuse.

MODERN RELIANCE ON USE-INHERITANCE MISPLACED.

Modern philanthropy—so far at least as it ever studies ultimate results—constantly relies on this ill-founded belief as its justification for ignoring the warnings of those who point out the ultimately disastrous results of a systematic defiance or reversal of the great law of natural selection. This reliance finds strong support in Mr. Spencer's latest teachings, for he holds that the inheritance of the effects of use and disuse takes place universally, and that it is now "the chief factor" in the evolution of civilized man (pp. 35, 74, iv)—natural selection being quite inadequate for the work of progressive modification. Practically he abandons the hope of evolution by natural selection, and substitutes the ideal of a nation being "modified en masse by transmission of the effects" of its institutions and habits. Use-inheritance will "mould its members far more rapidly and comprehensively" than can be effected by the survival of the fittest alone.

But could we rely upon the aid of use-inheritance if it really were a universal law and not a mere simulation of one? Let us consider some of the features of this alleged factor of evolution, seeing that it is henceforth to be our principal means of securing the improvement of our species and our continued adaptation to the changing conditions of a progressive civilization.

It is curiously uncertain and irregular in its action. It diminishes or abolishes some structures (such as jaws or eyes) without correspondingly diminishing or abolishing other equally disused and closely related parts (such as teeth, or eye-stalks). It thickens ducks' leg-bones while allowing them to shorten. It shortens the disused wing-bones of ducks and the leg-bones of rabbits while allowing them to thicken; and yet in other cases it greatly reduces the thickness of bones without shortening them. It transmits tameness most powerfully in an animal which usually cannot acquire it. It aids in webbing the feet of water-dogs, but fails to web the feet of the water-hen or to remove the web in the feet of upland geese.[72] It allows the disused fibula to retain a potentiality of development fully equal to that possessed by the long-used tibia. It lengthens legs because they are used in supporting the body, and shortens arms because they are used in pulling. Whether it enlarges brain if used in one way and diminishes it if used in another, we cannot tell; but it must obviously deaden nervous sensibilities in some cases and intensify them in others. It enlarges hands long before they are used, and thickens soles long before the time for walking on them. At the same time, as if by an oversight, it so delays its transmission of the habit of walking on these thickened soles, that the gradual and tedious acquisition of the non-transmitted habit costs the infant much time and trouble and often some pain and danger. Yet where aided by natural selection, as with chickens and foals, it transmits the habit in wonderful perfection and at a remarkably early date. It transmits new paces in horses in a single generation, but fails to perpetuate the songs of birds. It modifies offspring like parents, and yet allows the formation of two reproductive types in plants, and of two or more types widely different from the parents in some of the higher insects. It is said to be indispensable for the co-ordinated development of man and the giraffe and the elk, but appears to be unnecessary for the evolution and the maintenance of wonderful structures and habits and instincts in a thousand species of ants and bees and termites. It is the only possible means of complex evolution and adaptation of co-operative parts, and yet in Mr. Spencer's most representative case it renders such important parts as teeth and jaws unsuited for each other, and is said to ruin the teeth by the consequent overcrowding and decay. It survives amidst a general "lack of recognised evidence," and only seems to act usefully and healthily and regularly in quarters where it can least easily be distinguished from other more powerful and demonstrable factors of evolution. So little does it care to display its powers where they would be easily verifiable as well as useful that practical breeders ignore it. So slight is its independent power that it seems to allow natural selection or sexual selection or artificial selection to modify organisms in sheer defiance of its utmost opposition, just as readily as they modify organisms in other directions with its utmost help. If it partially perpetuates and extends the pecked-out indentations in the motmot's tail feathers, it on the other hand fails to transmit the slightest trace of mutilation in an almost infinite number of ordinary cases, and even where the mutilation is repeated for a hundred generations; and it apparently repairs rather than transmits the ordinary and oft-repeated losses caused by plucking hair, down and feathers, and the wear and tear of claws, teeth, hoofs and skin.

It is often mischievous as well as anomalous in its action. Under civilization with its division of labour, the various functions of mind and body are very unequally exercised. There is overwork or misuse of one part and disuse and neglect of others, leading to the partial breakdown or degeneration of various organs and to general deterioration of health through disturbed balance of the constitution. The brain, or rather particular parts of it, are often over-stimulated, while the body is neglected. In many ways education and civilization foster nervousness and weakness, and undermine the rude natural health and spirits of the human animal. Alcohol, tobacco, tea, coffee, extra brain work, late hours, dissipation, overwork, indoor life, division of labour, preservation of the weak, and many other causes, all help to injure the modern constitution; so that the prospect of cumulative intensification of these evils by the additional influence of use-inheritance is not an encouraging one. It is true that modern progress and prosperity are improving the people in various respects by their direct action; but if use-inheritance has any share in effecting this improvement it must also transmit increased wants and more luxurious habits, together with such evils as have already been referred to. As depicted by its defenders, use-inheritance transmits evils far more powerfully and promptly than benefits. It transmits insanity and shattered nerves rather than the healthy brain which preceded the breakdown. It perpetuates, and cumulatively intensifies, a deterioration in the senses of civilized men, but it fails to perpetuate the rank vigour of various plants when too well nourished, or the flourishing condition of various animals when too fat or when tamed. It already transmits the short-sight caused by so modern an art as watchmaking, but so fails to transmit the long-practised art of seeing (as it does of walking and talking) that vision is worse than useless to a man until he gradually acquires the necessary but non-transmitted associations of sensation and idea by his own experience. In a well-known case, a blind man on gaining his sight by an operation said that "all objects seemed to touch his eyes, as what he felt did his skin"—so little had the universal experience of countless ages impressed itself on his faculties. Under normal healthy conditions use-inheritance is so slow in its action that "several generations" must elapse before it produces any appreciable effect, and then that effect is only precisely what selection might be expected to bring about without its aid. Strong for evil and slow for good, it can convey epilepsy promptly in guinea pigs, but transmits the acquirements of genius so poorly that our best student of the heredity of genius has to account for the frequent and remarkable deterioration of the offspring by a theory which is strongly hostile to use-inheritance. It would tend to make organisms unworkable by the excessive differences in its rate and manner of action on co-operative parts, and by adapting these parts to the total amount of nourishment received rather than to occasional necessity or actual usefulness. It would tend to stereotype habits and convert reason into instinct.

How then can we rely upon use-inheritance for the improvement of the race? Even if it is not a sheer delusion, it may be more detrimental as a positive evil than it is advantageous as an unnecessary benefit; and as a normal modifying agent it is miserably weak and untrustworthy in comparison with the powerful selective influences by which nature and society continually and inevitably affect the species for good or for evil. The effects of use and disuse—rightly directed by education in its widest sense—must of course be called in to secure the highly essential but nevertheless superficial, limited, and partly deceptive improvement of individuals and of social manners and methods; but as this artificial development of already existing potentialities does not directly or readily tend to become congenital, it is evident that some considerable amount of natural or artificial selection of the more favourably varying individuals will still be the only means of securing the race against the constant tendency to degeneration which would ultimately swallow up all the advantages of civilization. The selective influences by which our present high level has been reached and maintained may well be modified, but they must not be abandoned or reversed in the rash expectation that State education, or State feeding of children, or State housing of the poor, or any amount of State socialism or public or private philanthropy, will prove permanently satisfactory substitutes. If ruinous deterioration and other more immediate evils, are to be avoided, the race must still be to the swift and the battle to the strong. The healthy Individualism so earnestly championed by Mr. Spencer must be allowed free play. Open competition, as Darwin teaches, with its survival and multiplication of the fittest, must be allowed to decide the battle of life independently of a foolish benevolence that prefers the elaborate cultivation and multiplication of weeds to the growth of corn and roses. We are trustees for the countless generations of the future. If we are wise we shall trust to the great ruling truths that we assuredly know, rather than to the seductive claims of an alleged factor of evolution for which no satisfactory evidence can be produced.

THE END.

RICHARD CLAY AND SONS, LIMITED, LONDON AND BUNGAY.

FOOTNOTES:

[72] Professor Romanes had casts made of the feet of upland geese, and could not detect any diminution as compared with the web of other geese in relation to the toes.



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