Essays: Scientific, Political, & Speculative; Vol. 1 of 3 / Library Edition (1891), Containing Seven Essays not before Republished, and Various other Additions.

Now, this criticism would have been very much to the point did the word Evolution truly express the process it names. If this process, as scientifically defined, really involved that conception which the word evolution was originally designed to convey, the implications would be those Mr. Martineau alleges. But, unfortunately for him, the word, having been in possession of the field before the process was understood, has been adopted merely because displacing it by another word seemed impracticable. And this adoption of it has been joined with a caution against misunderstandings arising from its unfitness. Here is a part of the caution:—"Evolution has other meanings, some of which are incongruous with, and some even directly opposed to, the meaning here given to it.... The antithetical word, Involution, would much more truly express the nature of the process; and would, indeed, describe better the secondary characters of the process which we shall have to deal with presently."[38] So that the meanings which the word involves, and which Mr. Martineau regards as fatal to the hypothesis, are already repudiated as not belonging to the hypothesis.

And now, having dealt with the essential objections raised by Mr. Martineau to the Hypothesis of Evolution as it is presented under that purely scientific form which generalizes the process of things, firstly as observed and secondly as inferred from certain ultimate principles, let me go on to examine that form of the Hypothesis which he propounds—Evolution as determined by Mind and Will—Evolution as pre-arranged by a Divine Actor. For Mr. Martineau apparently abandons the primitive theory of creation by "fiat of Almighty Will", and also the theory of creation by manufacture—by "a contriving and adapting power," and seems to believe in evolution: requiring only that "an originating Mind" shall be taken as its antecedent. Let us ask, first, in what relation Mr. Martineau conceives the "originating Mind" to stand to the evolving Universe. From some passages it is inferable that he considers the "presence of mind" to be everywhere needful. He says:—

This seems to be an unmistakable assertion that, wherever Evolution is going on, Mind is then and there behind it. At the close of the argument, however, a quite different conception is implied. Mr. Martineau says:—

It would take too much space to deal fully with the various questions which this last passage raises. There is the question—Whence come these "Forces," spoken of as separate from the "Will of God"—did they pre-exist? Then what becomes of the Divine Power? Do they exist by the Divine Will? Then what kind of nature is that by which they act apart from the Divine Will? Again, there is the question—How do these deputy-forces co-operate in each particular phenomenon, if the presiding Will is not there present to control them? Either an organ which develops into fitness for its function, develops by the co-operation of these forces under the direction of Mind then present, or it so develops in the absence of Mind. If it develops in the absence of Mind, the hypothesis is given up; and if the "originating Mind" is required to be then and there present, we must suppose a particular providence to be present in each particular organ of each particular creature throughout the universe. Once more there is the question—If "His thought is related to them [these Forces] as, in Man, the mental force is related to all below it," how can "His thought" be regarded as the cause of Evolution? In man the mental force is related to the forces below it neither as a creator of them nor as a regulator of them, save in a very limited way: the greater part of the forces present in man, both structural and functional, defy the mental force absolutely. Nay, more, it needs but to injure a nerve to see that the power of the mental force over the physical forces is dependent on conditions which are themselves physical; and one who takes morphia in mistake for magnesia, discovers that the power of the physical forces over the mental is unconditioned by any thing mental.

Not dwelling on these questions, however, I will merely draw attention to the entire incongruity of this conception with the previous conception which I have quoted. Assuming that, when the choice is pressed on him, Mr. Martineau will choose the first, which alone has any thing like defensibility, let us go on to ask how far Evolution is made more comprehensible by postulating Mind, universally immanent, as its cause.

In metaphysical controversy, many of the propositions propounded and accepted as quite believable, are absolutely inconceivable. There is a perpetual confusing of actual ideas with what are nothing but pseud-ideas. No distinction is made between propositions that contain real thoughts, and propositions that are only the forms of thoughts. A thinkable proposition is one of which the two terms can be brought together in consciousness under the relation said to exist between them. But very often, when the subject of a proposition has been thought of as something known, and when the predicate has been thought of as something known, and when the relation alleged between them has been thought of as a known relation, it is supposed that the proposition itself has been thought. The thinking separately of the elements of a proposition is mistaken for the thinking of them in the combination which the proposition affirms. And hence it continually happens that propositions which cannot be rendered into thought at all, are supposed to be not only thought but believed. The proposition that Evolution is caused by Mind is one of this nature. The two terms are separately intelligible; but they can be regarded in the relation of effect and cause only so long as no attempt is made to put them together in this relation.

The only thing which any one knows as Mind is the series of his own states of consciousness; and if he thinks of any mind other than his own, he can think of it only in terms derived from his own. If I am asked to frame a notion of Mind divested of all those structural traits under which alone I am conscious of mind in myself, I cannot do it. I know nothing of thought save as carried on in ideas originally traceable to the effects wrought by objects and forces on me. A mental act is an unintelligible phrase if I am not to regard it as an act in which states of consciousness are severally known as like other states in the series that has gone by, and in which the relations between them are severally known as like past relations in the series. If, then, I have to conceive Evolution as caused by an "originating Mind," I must conceive this Mind as having attributes akin to those of the only mind I know, and without which I cannot conceive Mind at all.

I will not dwell on the many incongruities hence resulting, by asking how the "originating Mind" is to be thought of as having states produced by things objective to it; as discriminating among these states, and classing them as like and unlike; and as preferring one objective result to another. I will simply ask—What happens if we ascribe to the "originating Mind" the character absolutely essential to the conception of Mind, that it consists of a series of states of consciousness? Put a series of states of consciousness as cause, and the evolving Universe as effect, and then endeavor to see the last as flowing from the first. I find it possible to imagine in some dim way a series of states of consciousness serving as antecedent to any one of the movements I see going on; for my own states of consciousness are often indirectly the antecedents to such movements. But how if I attempt to think of such a series as antecedent to all actions throughout the Universe—to the motions of the multitudinous stars through space, to the revolutions of all their planets round them, to the gyrations of all these planets on their axes, to the infinitely-multiplied physical processes going on in each of these suns and planets? I cannot think of a single series of states of consciousness as causing even the relatively small group of actions going on over the Earth's surface. I cannot think of it even as antecedent to all the various winds and the dissolving clouds they bear, to the currents of all the rivers, and the grinding actions of all the glaciers; still less can I think of it as antecedent to the infinity of processes simultaneously going on in all the plants that cover the globe, from scattered polar lichens to crowded tropical palms, and in all the millions of quadrupeds that roam among them, and the millions of millions of insects that buzz about them. Even to a single small set of these multitudinous terrestrial changes, I cannot conceive as antecedent a single series of states of consciousness—cannot, for instance, think of it as causing the hundred thousand breakers that are at this instant curling over on the shores of England. How, then, is it possible for me to conceive an "originating Mind," which I must represent to myself as a single series of states of consciousness, working the infinitely-multiplied sets of changes simultaneously going on in worlds too numerous to count, dispersed throughout a space that baffles imagination?

If, to account for this infinitude of physical changes everywhere going on, "Mind must be conceived as there" "under the guise of simple Dynamics," then the reply is that, to be so conceived, Mind must be divested of all attributes by which it is distinguished; and that, when thus divested of its distinguishing attributes, the conception disappears—the word Mind stands for a blank. If Mr. Martineau takes refuge in the entirely different and, as it seems to me, incongruous hypothesis of something like a plurality of minds—if he accepts, as he seems to do, the doctrine that you cannot explain Evolution "unless among your primordial elements you scatter already the germs of Mind as well as the inferior elements"—if the insuperable difficulties I have just pointed out are to be met by assuming a local series of states of consciousness for each phenomenon, then we are obviously carried back to something like the alleged fetichistic notion, with the difference only, that the assumed spiritual agencies are indefinitely multiplied.

Clearly, therefore, the proposition that an "originating Mind" is the cause of Evolution, is a proposition that can be entertained so long only as no attempt is made to unite in thought its two terms in the alleged relation. That it should be accepted as a matter of faith, may be a defensible position, provided good cause is shown why it should be so accepted; but that it should be accepted as a matter of understanding—as a statement making the order of the universe comprehensible—is a quite indefensible position.

Here let me guard myself against a misinterpretation very likely to be put upon the foregoing arguments; especially by those who have read the Essay to which they reply. The statements of that Essay carry the implication that all who adhere to the hypothesis it combats, imagine they have solved the mystery of things when they have shown the processes of Evolution to be naturally caused. Mr. Martineau tacitly represents them as believing that, when every thing has been interpreted in terms of Matter and Motion, nothing remains to be explained. This, however, is by no means the fact. The Doctrine of Evolution, under its purely scientific form, does not involve Materialism, though its opponents persistently represent it as doing so. Indeed, among adherents of it who are friends of mine, there are those who speak of the Materialism of Buechner and his school, with a contempt certainly not less than that felt by Mr. Martineau. To show how anti-materialistic my own view is, I may, perhaps, without impropriety, quote some out of many passages which I have written on the question elsewhere:

And again:

It is thus, I think, manifest that the difference between Mr. Martineau's view and the view he opposes is by no means so wide as he makes it appear; and further, it seems to me that such difference as exists is rather the reverse of that indicated by his exposition. Briefly expressed, the difference is that, where he thinks there is no mystery, the doctrine he combats recognizes a mystery. Speaking for myself only, I may say that, agreeing entirely with Mr. Martineau in repudiating the materialistic interpretation as utterly futile, I differ from him simply in this, that while he says he has found another interpretation, I confess that I cannot find any interpretation; while he holds that he can understand the Power which is manifested in things, I feel obliged to admit, after many failures, that I cannot understand it. So that, in presence of the transcendent problem which the universe presents, Mr. Martineau regards the human intellect as capable, and I as incapable. This contrast does not appear to me of the kind which his Essay tacitly asserts. If there is such a thing as the "pride of Science," it is obviously exceeded by the pride of Theology. I fail to perceive humility in the belief that the human mind is able to comprehend that which is behind appearances; and I do not see how piety is especially exemplified in the assertion that the Universe contains no mode of existence higher in Nature than that which is present to us in consciousness. On the contrary, I think it quite a defensible proposition that humility is better shown by a confession of incompetence to grasp in thought the Cause of all things; and that the religious sentiment may find its highest sphere in the belief that the Ultimate Power is no more representable in terms of human consciousness than human consciousness is representable in terms of a plant's functions.

THE FACTORS OF ORGANIC EVOLUTION.

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 re-moulding 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 armour 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, &c., &c. Still more obviously impossible is it to explain as due to use or disuse the colours 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 how 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. Having been myself one of these few, I look back with surprise at the way in which the facts which were congruous with the espoused view monopolized consciousness and kept out the facts which 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 favourable variations might have produced it; and the like holds of the no less remarkable appliance of the Venus's Fly-trap, 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 colours of each part, Mr. Darwin has shown us how, by the establishment of favourable 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 afterwards 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 vertebræ; 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 vertebræ, 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 cannot 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 favourable 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 colours 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 favourable 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 favourable 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 favourable variations, such changes of structure as adapt an organism to some useful action in which many different parts co-operate. 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, &c. 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 is 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."[41]

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 1864 (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 favourable 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 downwards and forwards. 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 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.

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.