The Factors of Organic Evolution · Herbert Spencer
I.
It needs but to remember how, when continuing to walk with a blistered foot, the taking of steps in such a modified way as to diminish pressure on the sore point, soon produces aching of muscles which are called into unusual action, to see that over-straining of any one of the muscles of the giraffe's hind-quarters might quickly incapacitate the animal when putting out all its powers to escape; and to be a few yards behind others would cause death. Hence if we are debarred from assuming that co-operative parts vary together even when adjacent and closely united—if we are still more debarred from assuming that with increased length of fore-legs or of neck, there will go an appropriate change in any one muscle or bone in the hind-quarters; how entirely out of the question it is to assume that there will simultaneously take place the appropriate changes in _all_ those many components of the hind-quarters which severally require re-adjustment. It is useless to reply that an increment of length in the fore-legs or neck might be retained and transmitted to posterity, waiting an appropriate variation in a particular bone or muscle in the hind-quarters, which, being made, would allow of a further increment. For besides the fact that until this secondary variation occurred the primary variation would be a disadvantage often fatal; and besides the fact that before such an appropriate secondary variation might be expected in the course of generations to occur, the primary variation would have died out; there is the fact that the appropriate variation of one bone or muscle in the hind-quarters would be useless without appropriate variations of all the rest—some in this way and some in that—a number of appropriate variations which it is impossible to suppose.
Nor is this all. Far more numerous appropriate variations would be indirectly necessitated. The immense change in the ratio of fore-quarters to hind-quarters would make requisite a corresponding change of ratio in the appliances carrying on the nutrition of the two. The entire vascular system, arterial and veinous, would have to undergo successive unbuildings and rebuildings to make its channels everywhere adequate to the local requirements; since any want of adjustment in the blood-supply in this or that set of muscles, would entail incapacity, failure of speed, and loss of life. Moreover the nerves supplying the various sets of muscles would have to be proportionately changed; as well as the central nervous tracts from which they issued. Can we suppose that all these appropriate changes, too, would be step by step simultaneously made by fortunate spontaneous variations, occurring along with all the other fortunate spontaneous variations? Considering how immense must be the number of these required changes, added to the changes above enumerated, the chances against any adequate re-adjustments fortuitously arising must be infinity to one. If the effects of use and disuse of parts are inheritable, then any change in the fore parts of the giraffe which affects the action of the hind limbs and back, will simultaneously cause, by the greater or less exercise of it, a re-moulding of each component in the hind limbs and back in a way adapted to the new demands; and generation after generation the entire structure of the hind-quarters will be progressively fitted to the changed structure of the fore-quarters: all the appliances for nutrition and innervation being at the same time progressively fitted to both. But in the absence of this inheritance of functionally-produced modifications, there is no seeing how the required re-adjustments can be made.
But there seems no reason to suppose that it will be increased in subsequent generations by natural selection. That it may be thus increased, the individuals not possessing more than average endowments of it, must be more frequently killed off than individuals highly endowed with it; and this can happen only when the attribute is one of greater importance, for the time being, than most of the other attributes. If those members of the species which have but ordinary shares of it, nevertheless survive by virtue of other superiorities which they severally possess; then it is not easy to see how this particular attribute can be developed by natural selection in subsequent generations. The probability seems rather to be, that by gamogenesis, this extra endowment will, on the average, be diminished in posterity—just serving in the long run to compensate the deficient endowments of other individuals, whose special powers lie in other directions; and so to keep up the normal structure of the species. The working out of the process is here somewhat difficult to follow; but it appears to me that as fast as the number of bodily and mental faculties increases, and as fast as the maintenance of life comes to depend less on the amount of any one, and more on the combined action of all; so fast does the production of specialities of character by natural selection alone, become difficult. Particularly does this seem to be so with a species so multitudinous in its powers as mankind; and above all does it seem to be so with such of the human powers as have but minor shares in aiding the struggle for life—the æsthetic faculties, for example.”
Dwelling for a moment on this last illustration of the class of difficulties described, let us ask how we are to interpret the development of the musical faculty. I will not enlarge on the family antecedents of the great composers. I will merely suggest the inquiry whether the greater powers possessed by Beethoven and Mozart, by Weber and Rossini, than by their fathers, were not due in larger measure to the inherited effects of daily exercise of the musical faculty by their fathers, than to inheritance, with increase, of spontaneous variations; and whether the diffused musical powers of the Bach clan, culminating in those of Johann Sebastian, did not result in part from constant practice; but I will raise the more general question—How came there that endowment of musical faculty which characterizes modern Europeans at large, as compared with their remote ancestors. The monotonous chants of low savages cannot be said to show any melodic inspiration; and it is not evident that an individual savage who had a little more musical perception than the rest, would derive any such advantage in the maintenance of life as would secure the spread of his superiority by inheritance of the variation. And then what are we to say of harmony? We cannot suppose that the appreciation of this, which is relatively modern, can have arisen by descent from the men in whom successive variations increased the appreciation of it—the composers and musical performers; for on the whole, these have been men whose worldly prosperity was not such as enabled them to rear many children inheriting their special traits. Even if we count the illegitimate ones, the survivors of these added to the survivors of the legitimate ones, can hardly be held to have yielded more than average numbers of descendants; and those who inherited their special traits have not often been thereby so aided in the struggle for existence as to further the spread of such traits. Rather the tendency seems to have been the reverse.
Since the above passage was written, I have found in the second volume of _Animals and Plants under Domestication_, a remark made by Mr. Darwin, practically implying that among creatures which depend for their lives on the efficiency of numerous powers, the increase of any one by the natural selection of a variation is necessarily difficult. Here it is. “Finally, as indefinite and almost illimitable variability is the usual result of domestication and cultivation, with the same part or organ varying in different individuals in different or even in directly opposite ways; and as the same variation, if strongly pronounced, usually recurs only after long intervals of time, any particular variation would generally be lost by crossing, reversion, and the accidental destruction of the varying individuals, unless carefully preserved by man.”—Vol. ii, 292. Remembering that mankind, subject as they are to this domestication and cultivation, are not, like domesticated animals, under an agency which picks out and preserves particular variations; it results that there must usually be among them, under the influence of natural selection alone, a continual disappearance of any useful variations of particular faculties which may arise. Only in cases of variations which are specially preservative, as for example, great cunning during a relatively barbarous state, can we expect increase from natural selection alone. We cannot suppose that minor traits, exemplified among others by the æsthetic perceptions, can have been evolved by natural selection. But if there is inheritance of functionally-produced modifications of structure, evolution of such minor traits is no longer inexplicable.
There are to be recognized two modes of this effect produced by changed conditions on the reproductive system, and consequently on offspring. Simple arrest of development is one. But beyond the variations of offspring arising from imperfectly developed reproductive systems in parents—variations which must be ordinarily in the nature of imperfections—there are others due to a changed balance of functions caused by changed conditions. The fact noted by Mr. Darwin in the above passage, “that the power of changed conditions accumulates; so that two, three, or more generations must be exposed to new conditions before any effect is visible,” implies that during these generations there is going on some change of constitution consequent on the changed proportions and relations of the functions. I will not dwell on the implication, which seems tolerably clear, that this change must consist of such modifications of organs as adapt them to their changed functions; and that if the influence of changed conditions “accumulates,” it must be through the inheritance of such modifications. Nor will I press the question—What is the nature of the effect registered in the reproductive elements, and which is subsequently manifested by variations?—Is it an effect entirely irrelevant to the new requirements of the variety?—Or is it an effect which makes the variety less fit for the new requirements?—Or is it an effect which makes it more fit for the new requirements? But not pressing these questions, it suffices to point out the necessary implication that changed functions of organs _do_, in some way or other, register themselves in changed proclivities of the reproductive elements. In face of these facts it cannot be denied that the modified action of a part produces an inheritable effect—be the nature of that effect what it may.
The second of the remarks above adverted to as made by Mr. Darwin, is contained in his sections dealing with correlated variations. In the _Origin of Species_, p. 114, he says— “The whole organization is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified.” And a parallel statement contained in _Animals and Plants under Domestication_, vol. ii, p. 320, runs thus— “Correlated variation is an important subject for us; for when one part is modified through continued selection, either by man or under nature, other parts of the organization will be unavoidably modified. From this correlation it apparently follows that, with our domesticated animals and plants, varieties rarely or never differ from each other by some single character alone.”
By what process does a changed part modify other parts? By modifying their functions in some way or degree, seems the necessary answer. It is indeed, imaginable, that where the part changed is some dermal appendage which, becoming larger, has abstracted more of the needful material from the general stock, the effect may consist simply in diminishing the amount of this material available for other dermal appendages, leading to diminution of some or all of them, and may fail to affect in appreciable ways the rest of the organism: save perhaps the blood-vessels near the enlarged appendage. But where the part is an active one—a limb, or viscus, or any organ which constantly demands blood, produces waste matter, secretes, or absorbs—then all the other active organs become implicated in the change. The functions performed by them have to constitute a moving equilibrium; and the function of one cannot, by alteration of the structure performing it, be modified in degree or kind, without modifying the functions of the rest—some appreciably and others inappreciably, according to the directness or indirectness of their relations. Of such inter-dependent changes, the normal ones are naturally inconspicuous; but those which are partially or completely abnormal, sufficiently carry home the general truth.
Thus, unusual cerebral excitement affects the excretion through the kidneys in quantity or quality or both. Strong emotions of disagreeable kinds check or arrest the flow of bile. A considerable obstacle to the circulation offered by some important structure in a diseased or disordered state, throwing more strain upon the heart, causes hypertrophy of its muscular walls; and this change which is, so far as concerns the primary evil, a remedial one, often entails mischiefs in other organs. “Apoplexy and palsy, in a scarcely credible number of cases, are directly dependent on hypertrophic enlargement of the heart.” And in other cases, asthma, dropsy, and epilepsy are caused. Now if a result of this inter-dependence as seen in the individual organism, is that a local modification of one part produces, by changing their functions, correlative modifications of other parts, then the question here to be put is—Are these correlative modifications, when of a kind falling within normal limits, inheritable or not. If they are inheritable, then the fact stated by Mr. Darwin that “when one part is modified through continued selection,” “other parts of the organization will be unavoidably modified” is perfectly intelligible: these entailed secondary modifications are transmitted _pari passu_ with the successive modifications produced by selection. But what if they are not inheritable? Then these secondary modifications caused in the individual, not being transmitted to descendants, the descendants must commence life with organizations out of balance, and with each increment of change in the part affected by selection, their organizations must get more out of balance—must have a larger and larger amounts of re-organization to be made during their lives. Hence the constitution of the variety must become more and more unworkable.
The only imaginable alternative is that the re-adjustments are effected in course of time by natural selection. But, in the first place, as we find no proof of concomitant variation among directly co-operative parts which are closely united, there cannot be assumed any concomitant variation among parts which are both indirectly co-operative and far from one another. And, in the second place, before all the many required re-adjustments could be made, the variety would die out from defective constitution. Even were there no such difficulty, we should still have to entertain a strange group of propositions, which would stand as follows:—1. Change in one part entails, by reaction on the organism, changes, in other parts, the functions of which are necessarily changed. 2. Such changes worked in the individual, affect, in some way, the reproductive elements: these being found to evolve unusual structures when the constitutional balance has been continuously disturbed. 3. But the changes in the reproductive elements thus caused, are not such as represent these functionally-produced changes: the modifications conveyed to offspring are irrelevant to these various modifications functionally produced in the organs of the parents. 4. Nevertheless, while the balance of functions cannot be re-established through inheritance of the effects of disturbed functions on structures, wrought throughout the individual organism; it can be re-established by the inheritance of fortuitous variations which occur in all the affected organs without reference to these changes of function.
Now without saying that acceptance of this group of propositions is impossible, we may certainly say that it is not easy. * * * * * “But where are the direct proofs that inheritance of functionally-produced modifications takes place?” is a question which will be put by those who have committed themselves to the current exclusive interpretation. “Grant that there are difficulties; still, before the transmitted effects of use and disuse can be legitimately assigned in explanation of them, we must have good evidence that the effects of use and disuse _are_ transmitted.” Before dealing directly with this demurrer, let me deal with it indirectly, by pointing out that the lack of recognized evidence may be accounted for without assuming that there is not plenty of it. Inattention and reluctant attention lead to the ignoring of facts which really exist in abundance; as is well illustrated in the case of pre-historic implements. Biassed by the current belief that no traces of man were to be found on the Earth's surface, save in certain superficial formations of very recent date, geologists and anthropologists not only neglected to seek such traces, but for a long time continued to pooh-pooh those who said they had found them. When M. Boucher de Perthes at length succeeded in drawing the eyes of scientific men to the flint implements discovered by him in the quarternary deposits of the Somme valley; and when geologists and anthropologists had thus been convinced that evidences of human existence were to be found in formations of considerable age, and thereafter began to search for them; they found plenty of them all over the world. Or again, to take an instance closely germane to the matter, we may recall the fact that the contemptuous attitude towards the hypothesis of organic evolution which naturalists in general maintained before the publication of Mr. Darwin's work, prevented them from seeing the multitudinous facts by which it is supported. Similarly, it is very possible that their alienation from the belief that there is a transmission of those changes of structure which are produced by changes of action, makes naturalists slight the evidence which supports that belief and refuse to occupy themselves in seeking further evidence.
If it be asked how it happens that there have been recorded multitudinous instances of variations fortuitously arising and re-appearing in offspring, while there have not been recorded instances of the transmission of changes functionally produced, there are three replies. The first is that changes of the one class are many of them conspicuous, while those of the other class are nearly all inconspicuous. If a child is born with six fingers, the anomaly is not simply obvious but so startling as to attract much notice; and if this child, growing up, has six-fingered descendants, everybody in the locality hears of it. A pigeon with specially-coloured feathers, or one distinguished by a broadened and upraised tail, or by a protuberance of the neck, draws attention by its oddness; and if in its young the trait is repeated, occasionally with increase, the fact is remarked, and there follows the thought of establishing the peculiarity by selection. A lamb disabled from leaping by the shortness of its legs, could not fail to be observed; and the fact that its offspring were similarly short-legged, and had a consequent inability to get over fences, would inevitably become widely known. Similarly with plants. That this flower had an extra number of petals, that that was unusually symmetrical, and that another differed considerably in colour from the average of its kind, would be easily seen by an observant gardener; and the suspicion that such anomalies are inheritable having arisen, experiments leading to further proofs that they are so, would frequently be made. But it is not thus with functionally-produced modifications. The seats of these are in nearly all cases the muscular, osseous, and nervous systems, and the viscera—parts which are either entirely hidden or greatly obscured. Modification in a nervous centre is inaccessible to vision; bones may be considerably altered in size or shape without attention being drawn to them; and, covered with thick coats as are most of the animals open to continuous observation, the increases or decreases in muscles must be great before they become externally perceptible.
A further important difference between the two inquiries is that to ascertain whether a fortuitous variation is inheritable, needs merely a little attention to the selection of individuals and the observation of offspring; while to ascertain whether there is inheritance of a functionally-produced modification, it is requisite to make arrangements which demand the greater or smaller exercise of some part or parts; and it is difficult in many cases to find such arrangements, troublesome to maintain them even for one generation, and still more through successive generations. Nor is this all. There exist stimuli to inquiry in the one case which do not exist in the other. The money-interest and the interest of the fancier, acting now separately and now together, have prompted multitudinous individuals to make experiments which have brought out clear evidence that fortuitous variations are inherited. The cattle-breeders who profit by producing certain shapes and qualities; the keepers of pet animals who take pride in the perfections of those they have bred; the florists, professional and amateur, who obtain new varieties and take prizes; form a body of men who furnish naturalists with countless of the required proofs. But there is no such body of men, led either by pecuniary interest or the interest of a hobby, to ascertain by experiments whether the effects of use and disuse are inheritable.
Thus, then, there are amply sufficient reasons why there is a great deal of direct evidence in the one case and but little in the other: such little being that which comes out incidentally. Let us look at what there is of it. * * * * * Considerable weight attaches to a fact which Brown-Séquard discovered, quite by accident, in the course of his researches. He found that certain artificially-produced lesions of the nervous system, so small even as a section of the sciatic nerve, left, after healing, an increasing excitability which ended in liability to epilepsy; and there afterwards came out the unlooked-for result that the offspring of guinea-pigs which had thus acquired an epileptic habit such that a pinch on the neck would produce a fit, inherited an epileptic habit of like kind. It has, indeed, been since alleged that guinea pigs tend to epilepsy, and that phenomena of the kind described, occur where there have been no antecedents like those in Brown-Séquard's case. But considering the improbability that the phenomena observed by him happened to be nothing more than phenomena which occasionally arise naturally, we may, until there is good proof to the contrary, assign some value to his results.
Evidence not of this directly experimental kind, but nevertheless of considerable weight, is furnished by other nervous disorders. There is proof enough that insanity admits of being induced by circumstances which, in one or other way, derange the nervous functions—excesses of this or that kind; and no one questions the accepted belief that insanity is inheritable. Is it alleged that the insanity which is inheritable is that which spontaneously arises, and that the insanity which follows some chronic perversion of functions is not inheritable? This does not seem a very reasonable allegation; and until some warrant for it is forthcoming, we may fairly assume that there is here a further support for belief in the transmission of functionally-produced changes. Moreover, I find among physicians the belief that nervous disorders of a less severe kind are inheritable. Men who have prostrated their nervous systems by prolonged overwork or in some other way, have children more or less prone to nervousness. It matters not what may be the form of inheritance—whether it be of a brain in some way imperfect, or of a deficient blood-supply; it is in any case the inheritance of functionally-modified structures.
Now-a-days most naturalists are more Darwinian than Mr. Darwin himself. I do not mean that their beliefs in organic evolution are more decided; though I shall be supposed to mean this by the mass of readers, who identify Mr. Darwin's great contribution to the theory of organic evolution, with the theory of organic evolution itself, and even with the theory of evolution at large. But I mean that the particular factor which he first recognized as having played so immense a part in organic evolution, has come to be regarded by his followers as the sole factor, though it was not so regarded by him. It is true that he apparently rejected altogether the causal agencies alleged by earlier inquirers. In the Historical Sketch prefixed to the later editions of his _Origin of Species_ (p. xiv, note), he writes:—“It is curious how largely my grandfather, Dr. Erasmus Darwin, anticipated the views and erroneous grounds of opinion of Lamarck in his 'Zoonomia' (vol. i, pp. 500-510), published in 1794.” And since, among the views thus referred to, was the view that changes of structure in organisms arise by the inheritance of functionally-produced changes, Mr. Darwin seems, by the above sentence, to have implied his disbelief in such inheritance. But he did not mean to imply this; for his belief in it as a cause of evolution, if not an important cause, is proved by many passages in his works. In the first chapter of the _Origin of Species_ (p. 11 of the first edition), he says respecting the inherited effects of habit, that “with animals the increased use or disuse of parts has had a marked influence;” and he gives as instances the changed relative weights of the wing bones and leg bones of the wild duck and the domestic duck, “the great and inherited development of the udders in cows and goats,” and the drooping ears of various domestic animals. Here are other passages taken from the latest edition of the work.
Ideas discussed in this chapter
Text from Project Gutenberg, public domain in the United States.