# Popular Science Monthly/Volume 28/April 1886/The Factors of Organic Evolution I

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* THE FACTORS OF ORGANIC EVOLUTION.
By HERBERT SPENCER.
I.
WITHIN the recollection of men now in middle life, opinion concerning the derivation of animals and plants was in a chaotic state. Among the unthinking there was tacit belief in creation by miracle, which formed an essential part of the creed of Christendom; and among the thinking there were two parties, each of which held an indefensible hypothesis. Immensely the larger of these parties, including nearly all whose scientific culture gave weight to their judgments, though not accepting literally the theologically orthodox doctrine, made a compromise between that doctrine and the doctrines which geologists had established; while opposed to them were some, mostly having no authority in science, who held a doctrine which was heterodox both theologically and scientifically. Professor Huxley, in his lecture on "The Coming of Age of the Origin of Species," remarks concerning the first of these parties as follows:
"One-and-twenty years ago, in spite of the work commenced by Hutton and continued with rare skill and patience by Lyell, the dominant view of the past history of the earth was catastrophic. Great and sudden physical revolutions, wholesale creations and extinctions of living beings, were the ordinary machinery of the geological epic brought into fashion by the misapplied genius of Cuvier. It was gravely maintained and taught that the end of every geological epoch was signalized by a cataclysm, by which every living being on the globe was swept away, to be replaced by a brand new creation when the world returned to quiescence. A scheme of nature which appeared to be modelled on the likeness of a succession of rubbers of whist, at the end of each of which the players upset the table and called for a new pack, did not seem to shock anybody.
Of the party above referred to as not satisfied with this conception described by Professor Huxley, there were two classes. The great majority were admirers of the Vestiges of the Natural History of Creation—a work which, while it sought to show that organic evolution has taken place, contended that the cause of organic evolution is "an impulse" supernaturally "imparted to the forms of life, advancing them. . . through grades of organization." Being nearly all very inadequately acquainted with the facts, those who accepted the view set forth in the Vestiges were ridiculed by the well-instructed for being satisfied with evidence, much of which was either invalid or easily cancelled by counter-evidence, and at the same time they exposed themselves to the ridicule of the more philosophical for being content with a supposed explanation which was in reality no explanation—the alleged "impulse" to advance giving us no more help in understanding the facts than does Nature's alleged "abhorrence of a vacuum" help us to understand the ascent of water in a pump. The remnant, forming the second of these classes, was very small. While rejecting this mere verbal solution, which both Dr. Erasmus Darwin and Lamarck had shadowed forth in other language, there were some few who, rejecting also the hypothesis indicated by both Dr. Darwin and Lamarck, that the promptings of desires or wants produced growths of the parts subserving them, accepted the single vera causa assigned by these writers—the modification of structures resulting from modification of functions. They recognized as the sole process in organic development the adaptation of parts and powers consequent on the effects of use and disuse—that continual moulding and remoulding of organisms to suit their circumstances, which is brought about by direct converse with such circumstances.
But while this cause accepted by these few is a true cause, since unquestionably during the life of the individual organism changes of function produce changes of structure; and while it is a tenable hypothesis that changes of structure so produced are inheritable, yet it was manifest to those not prepossessed, that this cause cannot with reason be assigned for the greater part of the facts. Though in plants there are some characters which may not irrationally be ascribed to the direct effects of modified functions consequent on modified circumstances, yet the majority of the traits presented by plants are not to be thus explained. It is impossible that the thorns by which a briar is in large measure defended against browsing animals, can have been developed and moulded by the continuous exercise of their protective actions; for in the first place, the great majority of the thorns are ​never touched at all, and, in the second place, we have no ground whatever for supposing that those which are touched are thereby made to grow, and to take those shapes which render them efficient. Plants which are rendered uneatable by the thick woolly coatings of their leaves, cannot have had these coatings produced by any process of reaction against the action of enemies; for there is no imaginable reason why, if one part of a plant is eaten, the rest should thereafter begin to develop the hairs on its surface. By what direct effect of function on structure can the shell of a nut have been evolved? Or how can those seeds which contain essential oils, rendering them unpalatable to birds, have been made to secrete such essential oils by these actions of birds which they restrain? Or how can the delicate plumes borne by some seeds, and giving the wind power to waft them to new stations, be due to any immediate influences of surrounding conditions? Clearly in these and in countless other cases, change of structure cannot have been directly caused by change of function. So is it with animals to a large extent, if not to the same extent. Though we have proof that by rough usage the dermal layer may be so excited as to produce a greatly thickened epidermal layer, sometimes quite horny; and though it is a feasible hypothesis that an effect of this kind persistently produced may be inherited; yet no such cause can explain the carapace of the turtle, the armor of the armadillo, or the imbricated covering of the manis. The skins of these animals are no more exposed to habitual hard usage than are those of animals covered by hair. The strange excrescences which distinguish the heads of the hornbills, cannot possibly have arisen from any reaction against the action of surrounding forces; for even were they clearly protective, there is no reason to suppose that the heads of these birds need protection more than the heads of other birds. If, led by the evidence that in animals the amount of covering is in some cases affected by the degree of exposure, it were admitted as imaginable that the development of feathers from preceding dermal growths had resulted from that extra nutrition caused by extra superficial circulation, we should still be without explanation of the structure of a feather. Nor should we have any clue to the specialities of feathers—the crests of various birds, the tails sometimes so enormous, the curiously placed plumes of the bird of paradise, etc. Still more obviously impossible is it to explain as due to use or disuse the colors of animals. No direct adaptation to function could have produced the blue protuberances on a mandril's face, or the striped hide of a tiger, or the gorgeous plumage of a kingfisher, or the eyes in a peacock's tail, or the multitudinous patterns of insects' wings. One single case, that of a deer's horns, might alone have sufficed to show bow insufficient was the assigned cause. During their growth, a deer's horns are not used at all; and when, having been cleared of the dead skin and dried-up blood-vessels covering them, they are ready ​for use, they are nerveless and non-vascular, and hence are incapable of undergoing any changes of structure consequent on changes of function.
Of these few, then, who rejected the belief described by Professor Huxley, and who, espousing the belief in a continuous evolution, had to account for this evolution, it must be said that though the cause assigned was a true cause, yet, even admitting that it operated through successive generations, it left unexplained the greater part of the facts. Obviously the facts that were congruous with the espoused view, monopolized consciousness, and kept out the facts that were incongruous with it—conspicuous though many of them were. The misjudgment was not unnatural. Finding it impossible to accept any doctrine which implied a breach in the uniform course of natural causation, and, by implication, accepting as unquestionable the origin and development of all organic forms by accumulated modifications naturally caused, that which appeared to explain certain classes of these modifications, was supposed to be capable of explaining the rest: the tendency being to assume that these would eventually be similarly accounted for, though it was not clear how.
Returning from this parenthetic remark, we are concerned here chiefly to remember that, as said at the outset, there existed thirty years ago, no tenable theory about the genesis of living things. Of the two alternative beliefs, neither would bear critical examination.
Out of this dead lock we were released—in large measure, though not I believe entirely—by the Origin of Species. That work brought into view a further factor; or, rather, such factor, recognized as in operation by here and there an observer (as pointed out by Mr. Darwin in his introduction to the second edition), was by him for the first time seen to have played so immense a part in the genesis of plants and animals.
Though laying myself open to the charge of telling a thrice-told tale, I feel obliged here to indicate briefly the several great classes of facts which Mr. Darwin's hypothesis explains; because otherwise that which follows would scarcely be understood. And I feel the less hesitation in doing this because the hypothesis which it replaced, not very widely known at any time, has of late so completely dropped into the background, that the majority of readers are scarcely aware of its existence, and do not therefore understand the relation between Mr. Darwin's successful interpretation and the preceding unsuccessful attempt at interpretation. Of these classes of facts, four chief ones may be here distinguished.
In the first place, such adjustments as those exemplified above are made comprehensible. Though it is inconceivable that a structure like that of the pitcher-plant could have been produced by accumulated effects of function on structure; yet it is conceivable that ​successive selections of favorable variations might have produced it; and the like holds of the no less remarkable appliance of the Venus's Flytrap, or the still more astonishing one of that water-plant by which infant-fish are captured. Though it is impossible to imagine how, by direct influence of increased use, such dermal appendages as a porcupine's quills could have been developed; yet, profiting as the members of a species otherwise defenceless might do by the stiffness of their hairs, rendering them unpleasant morsels to eat, it is a feasible supposition that from successive survivals of individuals thus defended in the greatest degrees, and the consequent growth in successive generations of hairs into bristles, bristles into spines, spines into quills (for all these are homologous), this change could have arisen. In like manner, the odd inflatable bag of the bladder-nosed seal, the curious fishing-rod with its worm-like appendage carried on the head of the lophius or angler, the spurs on the wings of certain birds, the weapons of the sword-fish and saw-fish, the wattles of fowls, and numberless such peculiar structures, though by no possibility explicable as due to effects of use or disuse, are explicable as resulting from natural selection operating in one or other way.
In the second place, while showing us how there have arisen countless modifications in the forms, structures, and colors of each part, Mr. Darwin has shown us how, by the establishment of favorable variations, there may arise new parts. Though the first step in the production of horns on the heads of various herbivorous animals, may have been the growth of callosities consequent on the habit of butting—such callosities thus functionally initiated being afterward developed in the most advantageous ways by selection; yet no explanation can be thus given of the sudden appearance of a duplicate set of horns, as occasionally happens in sheep: an addition which, where it proved beneficial, might readily be made a permanent trait by natural selection. Again, the modifications which follow use and disuse can by no possibility account for changes in the numbers of vertebras; but after recognizing spontaneous, or rather fortuitous, variation as a factor, we can see that where an additional vertebra hence resulting (as in some pigeons) proves beneficial, survival of the fittest may make it a constant character; and there may, by further like additions, be produced extremely long strings of vertebrae, such as snakes show us. Similarly with the mammary glands. It is not an unreasonable supposition that by the effects of greater or less function, inherited through successive generations, these may be enlarged or diminished in size; but it is out of the question to allege such a cause for changes in their numbers. There is no imaginable explanation of these save the establishment by inheritance of spontaneous variations, such as are known to occur in the human race.
So too, in the third place, with certain alterations in the connexions of parts. According to the greater or smaller demands made on this ​or that limb, the muscles moving it may be augmented or diminished in bulk; and, if there is inheritance of changes so wrought, the limb may, in course of generations, be rendered larger or smaller. But changes in the arrangements or attachments of muscles can not be thus accounted for. It is found, especially at the extremities, that the relations of tendons to bones and to one another are not always the same. Variations in their modes of connexion may occasionally prove advantageous, and may thus become established. Here again, then, we have a class of structural changes to which Mr. Darwin's hypothesis gives us the key, and to which there is no other key.
Once more there are the phenomena of mimicry. Perhaps in a more striking way than any others, these show how traits which seem inexplicable are explicable as due to the more frequent survival of individuals that have varied in favorable ways. We are enabled to understand such marvellous simulations as those of the leaf-insect, those of beetles which "resemble glittering dew-drops upon the leaves;" those of caterpillars, which, when asleep, stretch themselves out so as to look like twigs. And we are shown how there have arisen still more astonishing imitations—those of one insect by another. As Mr. Bates has proved, there are cases in which a species of butterfly, rendered so unpalatable to insectivorous birds by its disagreeable taste that they will not catch it, is simulated in its colors and markings by a species which is structurally quite different—so simulated that even a practised entomologist is liable to be deceived: the explanation being that an original slight resemblance, leading to occasional mistakes on the part of birds, was increased generation after generation by the more frequent escape of the most-like individuals, until the likeness became thus great.
But now, recognizing in full this process brought into clear view by Mr. Darwin, and traced out by him with so much care and skill, can we conclude that, taken alone, it accounts for organic evolution? has the natural selection of favorable variations been the sole factor? On critically examining the evidence, we shall find reason to think that it by no means explains all that has to be explained. Omitting for the present any consideration of a factor which may be distinguished as primordial, it may be contended that the above-named factor alleged by Dr. Erasmus Darwin and by Lamarck, must be recognized as a co-operator. Utterly inadequate to explain the major part of the facts as is the hypothesis of the inheritance of functionally produced modifications, yet there is a minor part of the facts, very extensive though less, which must be ascribed to this cause.
When discussing the question more than twenty years ago (Principles of Biology, § 166), I instanced the decreased size of the jaws in the civilized races of mankind, as a change not accounted for by the natural selection of favorable variations; since no one of the ​decrements by which, in thousands of years, this reduction has been effected, could have given to an individual in which it occurred, such advantage as would cause his survival, either through diminished cost of local nutrition or diminished weight to be carried. I did not then exclude, as I might have done, two other imaginable causes. It may be said that there is some organic correlation between increased size of brain and decreased size of jaw: Camper's doctrine of the facial angle being referred to in proof. But this argument may be met by pointing to the many examples of small-jawed people who are also small-brained, and by citing not infrequent cases of individuals remarkable for their mental powers, and at the same time distinguished by jaws not less than the average but greater. Again, if sexual selection be named as a possible cause, there is the reply that, even supposing such slight diminution of jaw as took place in a single generation to have been an attraction, yet the other incentives to choice on the part of men have been too many and great to allow this one to weigh in an adequate degree; while, during the greater portion of the period, choice on the part of women has scarcely operated: in earlier times they were stolen or bought, and in later times mostly coerced by parents. Thus, reconsideration of the facts does not show me the invalidity of the conclusion drawn, that this decrease in size of jaw can have had no other cause than continued inheritance of those diminutions consequent on diminutions of function, implied by the use of selected and well-prepared food. Here, however, my chief purpose is to add an instance showing, even more clearly, the connexion between change of function and change of structure. This instance, allied in nature to the other, is presented by those varieties, or rather sub-varieties, of dogs, which, having been household pets, and habitually fed on soft food, have not been called on to use their jaws in tearing and crunching, and have been but rarely allowed to use them in catching prey and in fighting. No inference can be drawn from the sizes of the jaws themselves, which, in these dogs, have probably been shortened mainly by selection. To get direct proof of the decrease of the muscles concerned in closing the jaws or biting, would require a series of observations very difficult to make. But it is not difficult to get indirect proof of this decrease by looking at the bony structures with which these muscles are connected. Examination of the skulls of sundry indoor dogs contained in the Museum of the College of Surgeons, proves the relative smallness of such parts. The only pug dog's skull is that of an individual not perfectly adult; and though its traits are quite to the point they cannot with safety be taken as evidence. The skull of a toy-terrier has much restricted areas of insertion for the temporal muscles; has weak zygomatic arches; and has extremely small attachments for the masseter muscles. Still more significant is the evidence furnished by the skull of a King Charles's spaniel, which, if we allow three years to a generation, and bear in ​mind that the variety must have existed before Charles the Second's reign, we may assume belongs to something approaching to the hundredth generation of these household pets. The relative breadth between the outer surfaces of the zygomatic arches is conspicuously small; the narrowness of the temporal fossæ is also striking; the zygomata are very slender; the temporal muscles have left no marks whatever, either by limiting lines or by the character of the surfaces covered; and the places of attachment for the masseter muscles are very feebly developed. At the Museum of Natural History, among skulls of dogs there is one which, though unnamed, is shown by its small size and by its teeth, to have belonged to one variety or other of lap-dogs, and which has the same traits in an equal degree with the skull just described. Here, then, we have two if not three kinds of dogs which, similarly leading protected and pampered lives, show that in the course of generations the parts concerned in clenching the jaws have dwindled. To what cause must this decrease be ascribed? Certainly not to artificial selection; for most of the modifications named make no appreciable external signs: the width across the zygomata could alone be perceived. Neither can natural selection have had anything to do with it; for even were there any struggle for existence among such dogs, it cannot be contended that any advantage in the struggle could be gained by an individual in which a decrease took place. Economy of nutrition, too, is excluded. Abundantly fed as such dogs are, the constitutional tendency is to find places where excess of absorbed nutriment may be conveniently deposited, rather than to find places where some cutting down of the supplies is practicable. Nor again can there be alleged a possible correlation between these diminutions and that shortening of the jaws which has probably resulted from selection; for in the bull-dog, which has also relatively short jaws, these structures concerned in closing them are unusually large. Thus there remains as the only conceivable cause, the diminution of size which results from diminished use. The dwindling of a little exercised part has, by inheritance, been made more and more marked in successive generations.
Difficulties of another class may next be exemplified—those which present themselves when we ask how there can be effected by the selection of favorable variations, such changes of structure as adapt an organism to some useful action in which many different parts cooperate. None can fail to see how a simple part may, in course of generations, be greatly enlarged, if each enlargement furthers, in some decided way, maintenance of the species. It is easy to understand, too, how a complex part, as an entire limb, may be increased as a whole by the simultaneous due increase of its co-operative parts; since if, while it is growing, the channels of supply bring to the limb an unusual quantity of blood, there will naturally result a ​proportionately greater size of all its components—bones, muscles, arteries, veins, etc. But though in cases like this, the co-operative parts forming some large complex part may be expected to vary together, nothing implies that they necessarily do so; and we have proof that in various cases, even when closely united, they do not do so. An example is furnished by those blind crabs named in the Origin of Species which inhabit certain dark caves of Kentucky, and which, though they have lost their eyes, have not lost the foot-stalks which carried their eyes. In describing the varieties which have been produced by pigeon-fanciers, Mr. Darwin notes the fact that along with changes in length of beak produced by selection, there have not gone proportionate changes in length of tongue. Take again the case of teeth and jaws. In mankind these have not varied together. During civilization the jaws have decreased, but the teeth have not decreased in proportion; and hence that prevalent crowding of them, often remedied in childhood by extraction of some, and in other cases causing that imperfect development which is followed by early decay. But the absence of proportionate variation in co-operative parts that are close together, and are even bound up in the same mass, is best seen in those varieties of dogs named above as illustrating the inherited effects of disuse. We see in them, as we see in the human race, that diminution in the jaws has not been accompanied by corresponding diminution in the teeth. In the catalogue of the College of Surgeons Museum, there are appended to the entry which identifies a Blenheim Spaniel's skull, the words—"the teeth are closely crowded together," and to the entry concerning the skull of a King Charles's Spaniel the words—"the teeth are closely packed, p. 3, is placed quite transversely to the axis of the skull." It is further noteworthy that in a case where there is no diminished use of the jaws, but where they have been shortened by selection, a like want of concomitant variation is manifested: the case being that of the bull-dog, in the upper jaw of which also, "the premolars. . . are excessively crowded, and placed obliquely or even transversely to the long axis of the skull."
If, then, in cases where we can test it, we find no concomitant variation in co-operative parts that are near together—if we do not find it in parts which, though belonging to different tissues, are so closely united as teeth and jaws—if we do not find it even when the co-operative parts are not only closely united, but are formed out of the same tissue, like the crab's eye and its peduncle; what shall we say of co-operative parts which, besides being composed of different tissues, are remote from one another? Not only are we forbidden to ​assume that they vary together, but we are warranted in asserting that they can have no tendency to vary together. And what are the implications in cases where increase of a structure can be of no service unless there is concomitant increase in many distant structures, which have to join it in performing the action for which it is useful?
As far back as 1804 (Principles of Biology, § 166) I named in illustration an animal carrying heavy horns—the extinct Irish elk; and indicated the many changes in bones, muscles, blood-vessels, nerves, composing the fore-part of the body, which would be required to make an increment of size in such horns advantageous. Here let me take another instance—that of the giraffe: an instance which I take partly because, in the sixth edition of the Origin of Species, issued in 1872, Mr. Darwin has referred to this animal when effectually disposing of certain arguments urged against his hypothesis. He there says:
And in the summary of the chapter, he remarks concerning the adjustments in the same quadruped, that "the prolonged use of all the parts together with inheritance will have aided in an important manner in their co-ordination" (p. 199): a remark probably having reference chiefly to the increased massiveness of the lower part of the neck; the increased size and strength of the thorax required to bear the additional burden; and the increased strength of the fore-legs required to carry the greater weight of both. But now I think that further consideration suggests the belief that the entailed modifications are much more numerous and remote than at first appears; and that the greater part of these are such as cannot be ascribed in any degree to the selection of favorable variations, but must be ascribed exclusively to the inherited effects of changed functions. Whoever has seen a giraffe gallop will long remember the sight as a ludicrous one. The reason for the strangeness of the motions is obvious. Though the fore-limbs and the hind-limbs differ so much in length, yet in galloping they have to keep pace—must take equal strides. The result is that at each stride, the angle which the hind-limbs describe round their centre of motion is much larger than the angle described by the fore-limbs. And beyond this, as an aid in equalizing the strides, the hind part of the back is at each stride bent very much downward and forward. Hence the hind-quarters appear to be doing nearly all the work. Now a moment's observation shows that the bones and muscles composing the hind-quarters of the giraffe, perform actions differing in one or other way and degree, from the actions performed by the homologous bones and muscles in a mammal of ordinary proportions, ​and from those in the ancestral mammal which gave origin to the giraffe. Each further stage of that growth which produced the large fore-quarters and neck, entailed some adapted change in sundry of the numerous parts composing the hind-quarters; since any failure in the adjustment of their respective strengths would entail some defect in speed and consequent loss of life when chased. 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 forequarters 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.
Yet a third class of difficulties stands in the way of the belief that the natural selection of useful variations is the sole factor of organic evolution. This class of difficulties, already pointed out in § 166 of the Principles of Biology, I cannot more clearly set forth than in the words there used. Hence I may perhaps be excused for here quoting them:
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:
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.
Two remarks made by Mr. Darwin have implications from which the same general conclusion must, I think, be drawn. Speaking of the variability of animals and plants under domestication, he says:
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:
And a parallel statement contained in Animals and Plants under Domestication, vol. ii, p. 320, runs thus:
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 How 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 hypertropic 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 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 quaternary 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 toward 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 reappearing 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-colored 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 color 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 afterward 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.
​Verification of the reasons above given for the paucity of this direct evidence is yielded by contemplation of it, for it is observable that the cases named are cases which, from one or other cause, have thrust themselves on observation. They justify the suspicion that it is not because such cases are rare that many of them cannot be cited, but simply because they are mostly unobtrusive, and to be found only by that deliberate search which nobody makes. I say nobody, but I am wrong. Successful search has been made by one whose competence as an observer is beyond question, and whose testimony is less liable than that of all others to any bias toward the conclusion that such inheritance takes place. I refer to the author of the Origin of Species.
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.
In his subsequent work, The Variation of Animals and Plants under Domestication, where he goes into full detail, Mr. Darwin gives more numerous illustrations of the inherited effects of use and disuse. The following are some of the cases, quoted from volume i of the first edition:
Here are some instances of like meaning from volume ii.
But Mr. Darwin's admission, or rather his assertion, that the inheritance of functionally-produced modifications has been a factor in organic evolution, is made clear not by these passages alone and by kindred ones. It is made clearer still by a passage in the preface to the second edition of his Descent of Man. lie there protests against that current version of his views in which this factor makes no appearance. The passage is as follows:
Nor is this all. There is evidence that Mr. Darwin's belief in the efficiency of this factor, became stronger as he grew older and accumulated more evidence. The first of the extracts above given, taken from the sixth edition of the Origin of Species, runs thus:
Now on turning to the first edition, p. 134, it will be found that instead of the words—"I think there can be no doubt," the words originally used were—"I think there can be little doubt." That this deliberate erasure of a qualifying word and substitution of a word implying unqualified belief, was due to a more decided recognition of a factor originally under-estimated, is clearly implied by the wording of the above-quoted passage from the preface to the Descent of Man; where he says that "even in the first edition of the 'Origin of Species,'" etc.: the implication being that much more in subsequent editions, and ​subsequent works, had he insisted on this factor. The change thus indicated is especially significant as having occurred at a time of life when the natural tendency is toward fixity of opinion.
During that earlier period when he was discovering the multitudinous cases in which his own hypothesis afforded solutions, and simultaneously observing how utterly futile in these multitudinous cases was the hypothesis propounded by his grandfather and Lamarck, Mr. Darwin was, not unnaturally, almost betrayed into the belief that the one is all-sufficient and the other inoperative. But in the mind of one so candid and ever open to more evidence, there naturally came a reaction. The inheritance of functionally-produced modifications, which, judging by the passage quoted above concerning the views of these earlier inquirers, would seem to have been at one time denied, but which as we have seen was always to some extent recognized, came to be recognized more and more, and deliberately included as a factor of importance.
Of this reaction displayed in the later writings of Mr. Darwin, let us now ask—Has it not to be carried further? Was the share in organic evolution which Mr. Darwin latterly assigned to the transmission of modifications caused by use and disuse, its due share? Consideration of the groups of evidences given above, will, I think, lead us to believe that its share has been much larger than he supposed even in his later days.
There is first the implication yielded by extensive classes of phenomena which remain inexplicable in the absence of this factor. If, as we see, co-operative parts do not vary together, even when few and close together, and may not therefore be assumed to do so when many and remote, we cannot account for those innumerable changes in organization which are implied when, for advantageous use of some modified part, many other parts which join it in action have to be modified.
Further, as increasing complexity of structure, accompanying increasing complexity of life, implies increasing number of faculties, of which each one conduces to preservation of self or descendants; and as the various individuals of a species, severally requiring something like the normal amounts of all these, may individually profit, here by an unusual amount of one, and there by an unusual amount of another; it follows that as the number of faculties becomes greater, it becomes more difficult for any one to be further developed by natural selection. Only where increase of some one is predominantly advantageous does the means seem adequate to the end. Especially in the case of powers which do not subserve self-preservation in appreciable degrees, does development by natural selection appear impracticable.
It is a fact recognized by Mr. Darwin, that where, by selection through successive generations, a part has been increased or decreased, ​its reaction upon other parts entails changes in them. This reaction is effected through the changes of function involved. If the changes of structure produced by such changes of function, are inheritable, then the re-adjustment of parts throughout the organism, taking place generation after generation, maintains an approximate balance; but if not, then generation after generation the organism must get more and more out of gear, and tend to become unworkable.
Further, as it is proved that change in the balance of functions registers its effects on the reproductive elements, we have to choose between the alternatives that the registered effects are irrelevant to the particular modifications which the organism has undergone, or that they are such as tend to produce repetitions of these modifications. The last of these alternatives makes the facts comprehensible; but the first of them not only leaves us with several unsolved problems, but is incongruous with the general truth that by reproduction, ancestral traits, down to minute details, are transmitted.
Though, in the absence of pecuniary interests and the interests in hobbies, no such special experiments as those which have established the inheritance or fortuitous variations have been made to ascertain whether functionally-produced modifications are inherited; yet certain apparent instances of such inheritance have forced themselves on observation without being sought for. In addition to other indications of a less conspicuous kind, is the one I have given above—the fact that the apparatus for tearing and mastication has decreased with decrease of its function, alike in civilized man and in some varieties of dogs which lead protected and pampered lives. Of the numerous cases named by Mr. Darwin, it is observable that they are yielded not by one class of parts only, but by most if not all classes—by the dermal system, the muscular system, the osseous system, the nervous system, the viscera; and that among parts liable to be functionally modified, the most numerous observed cases of inheritance are furnished by those which admit of preservation and easy comparison—the bones: these cases, moreover, being specially significant as showing how, in sundry unallied species, parallel changes of structure have occurred along with parallel changes of habit.
What, then, shall we say of the general implication? Are we to stop short with the admission that inheritance of functionally-produced modifications takes place only in cases in which there is evidence of it? May we properly assume that these many instances of changes of structure caused by changes of function, occurring in various tissues and various organs, are merely special and exceptional instances having no general significance? Shall we suppose that though the evidence which already exists has come to light without aid from a body of inquirers, there would be no great increase were due attention devoted to the collection of evidence? This is, I think, not a reasonable supposition. To me the ensemble of the facts suggests the belief, ​scarcely to be resisted, that the inheritance of functionally-produced modifications takes place universally. Looking at physiological phenomena as conforming to physical principles, it is difficult to conceive that a changed play of organic forces which in many cases of different kinds produces an inherited change of structure, does not do this in all cases. The implication, very strong I think, is that the action of every organ produces on it a reaction which, usually not altering its rate of nutrition, sometimes leaves it with diminished nutrition consequent on diminished action, and at other times increases its nutrition in proportion to its increased action; 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; and that in ways mostly too small to be identified, but occasionally in more conspicuous ways and in the course of generations, the resulting modifications of one or other kind show themselves. Further, it seems to me that as there are certain extensive classes of phenomena which are inexplicable if we assume the inheritance of fortuitous variations to be the sole factor, but which become at once explicable if we admit the inheritance of functionally-produced changes, we are justified in concluding that this inheritance of functionally-produced changes has been not simply a co-operating factor in organic evolution, but has been a co-operating factor without which organic evolution, in its higher forms at any rate, could never have taken place.
Be this or be it not a warrantable conclusion, there is, I think, good reason for a provisional acceptance of the hypothesis that the effects of use and disuse are inheritable; and for a methodic pursuit of inquiries with the view of either establishing it or disproving it. It seems scarcely reasonable to accept without clear demonstration, the belief that while a trivial difference of structure arising spontaneously is transmissible, a massive difference of structure, maintained generation after generation by change of function, leaves no trace in posterity. Considering that unquestionably the modification of structure by function is a vera causa, in so far as concerns the individual; and considering the number of facts which so competent an observer as Mr. Darwin regarded as evidence that transmission of such modifications takes place in particular cases; the hypothesis that such transmission takes place in conformity with a general law, holding of all active structures, should, I think, be regarded as at least a good working hypothesis.
But now supposing the broad conclusion above drawn to be granted—supposing all to agree that from the beginning, along with inheritance of useful variations fortuitously arising, there has been inheritance of effects produced by use and disuse; do there remain no classes of organic phenomena unaccounted for? To this question I ​think it must be replied that there Jo remain classes of organic phenomena unaccounted for. It may, I believe, be shown that certain cardinal traits of animals and plants at large are still unexplained; and that a further factor must be recognized. To show this, however, will require another paper.