# The Principles of Biology Vol. I/Chapter II.5

CHAPTER V.
ADAPTATION.
§ 67. In plants waste and repair being scarcely appreciable, there are not likely to arise appreciable changes in the proportions of already-formed parts. The only divergences from the average structures of a species, which we may expect particular conditions to produce, are those producible by the action of these conditions on parts in course of formation; and such divergences we do find. We know that a tree which, standing alone in an exposed position, has a short and thick stem, has a tall and slender stem when it grows in a wood; and that also its branches then take a different inclination. We know that potato-sprouts which, on reaching the light, develop into foliage, will, in the absence of light, grow to a length of several feet without foliage. And every in-door plant furnishes proof that shoots and leaves, by habitually turning themselves to the light, exhibit a certain adaptation—an adaptation due, as we must suppose; to the special effects of the special conditions on the still growing parts. In animals, however, besides analogous structural changes wrought during the period of growth, by subjection to circumstances unlike the ordinary circumstances, there are structural changes similarly wrought after maturity has been reached. Organs that have arrived at their full sizes possess a certain modifiability; so that while the ​organism as a whole retains pretty nearly the same bulk, the proportions of its parts may be considerably varied. Their variations, here treated of under the title Adaptation, depend on specialities of individual action. In the last chapter we saw that the actions of organisms entail re-actions on them; and that specialities of action entail specialities of re-action. Here it remains to be pointed out that these special actions and re-actions do not end with temporary changes, but work permanent changes.
If, in an adult animal, the waste and repair in all parts were exactly balanced—if each organ daily gained by nutrition exactly as much as it lost daily by the discharge of its function—if excess of function were followed only by such excess of nutrition as balanced the extra waste; it is clear that there would occur no change in the relative sizes of organs. But there is no such exact balance. If the excess of function, and consequent excess of waste, is moderate, it is not simply compensated by repair but more than compensated—there is a certain increase of bulk. This is true to some degree of the organism as a whole, when the organism is framed for activity. A considerable waste giving considerable power of assimilation, is more favourable to accumulation of tissue than is quiescence with its comparatively feeble assimilation: whence results a certain adaptation of the whole organism to its requirements. But it is more especially true of the parts of an organism in relation to one another. The illustrations fall into several groups. The growth of muscles exercised to an unusual degree is a matter of common observation. In the often-cited blacksmith's arm, the dancer's legs and the jockey's crural adductors, we have marked examples of a modifiability which almost every one has to some extent experienced. It is needless to multiply proofs. The occurrence of changes in the structure of the skin, where the skin is exposed to unusual stress of function, is also familiar. That thickening of the epidermis on a labourer's palm results from continual pressure and ​friction, is certain. Those who have not before exerted their hands, find that such an exercise as rowing soon begins to produce a like thickening. This relation of cause and effect is still better shown by the marked indurations at the ends of a violinist's fingers. Even in mucous membrane, which ordinarily is not subject to mechanical forces of any intensity, similar modifications are possible: witness the callosity of the gums which arises in those who have lost their teeth, and have to masticate without teeth. The vascular system furnishes good instances of the increased growth that follows increased function. When, because of some permanent obstruction to the circulation, the heart has to exert a greater contractile force on the mass of blood which it propels at each pulsation, and when there results the laboured action known as palpitation, there usually occurs dilatation, or hypertrophy, or a mixture of the two: the dilatation, which is a yielding of the heart's structure under the increased strain, implying a failure to meet the emergency; but the hypertrophy, which consists in a thickening of the heart's muscular walls, being an adaptation of it to the additional effort required. Again, when an aneurism in some considerable artery has been obliterated, either artifically or by a natural inflammatory process; and when this artery has consequently ceased to be a channel for the blood; some of the adjacent arteries which anastomose with it become enlarged, so as to carry the needful quantity of blood to the parts supplied. Though we have no direct proof of analogous modifications in nervous structures, yet indirect proof is given by the greater efficiency that follows greater activity. This is manifested alike in the senses and the intellect. The palate may be cultivated into extreme sensitiveness, as in professional tea-tasters. An orchestral conductor gains, by continual practice, an unusually great ability to discriminate differences of sound. In the finger-reading of the blind we have evidence that the sense of touch may be brought by exercise to a far higher capability than is ​ordinary. The increase of power which habitual exertion gives to mental faculties needs no illustration: every person of education has personal experience of it. Even from the osseous structures evidence may be drawn. The bones of men accustomed to great muscular action are more massive, and have more strongly marked processes for the attachment of muscles, than the bones of men who lead sedentary lives; and a like contrast holds between the bones of wild and tame animals of the same species. Adaptations of another order, in which there is a qualitative rather than a quantitative modification, arise after certain accidents to which the skeleton is liable. When the hip-joint has been dislocated, and long delay has made it impossible to restore the parts to their proper places, the head of the thigh-bone, imbedded in the surrounding muscles, becomes fixed in its new position by attachments of fibrous tissue, which afford support enough to permit a halting walk. But the most remarkable modification of this order occurs in united ends of fractured bones. "False joints" are often formed—joints which rudely simulate the hinge structure or the ball-and-socket structure, according as the muscles tend to produce a motion of flexion and extension or a motion of rotation. In the one case, according to Rokitansky, the two ends of the broken bone become smooth and covered with periosteum and fibrous tissue, and are attached by ligaments that allow a certain backward and forward motion; and in the other case the ends, similarly clothed with the appropriate membranes, become the one convex and the other concave, are inclosed in a capsule, and are even occasionally supplied with synovial fluid!
The general truth that extra function is followed by extra growth, must be supplemented by the equally general truth, ​that beyond a limit, usually soon reached, very little, if any, further modification can be produced. The experiences which we colligate into the one induction thrust the other upon us. After a time no training makes the pugilist or the athlete any stronger. The adult gymnast at last acquires the power to perform certain difficult feats; but certain more difficult feats no additional practice enables him to perform. Years of discipline give the singer a particular loudness and range of voice, beyond which further discipline does not give greater loudness or wider range: on the contrary, increased vocal exercise, causing a waste in excess of repair, is often followed by decrease of power. In the exaltation of the perceptions we see similar limits. The culture which raises the susceptibility of the ear to the intervals and harmonies of notes, will not turn a bad ear into a good one. Lifelong effort fails to make this artist a correct draftsman or that a fine colourist: each does better than he did at first, but each falls short of the power attained by some other artists. Nor is this truth less clearly illustrated among the more complex mental powers. A man may have a mathematical faculty, a poetical faculty, or an oratorical faculty, which special education improves to a certain extent. But unless he is unusually endowed in one of those directions, no amount of education will make him a first-rate mathematician, a first-rate poet, or a first-rate orator. Thus the general fact appears to be that while in each individual certain changes in the proportions of parts may be caused by variations of functions, the congenital structure of each individual puts a limit to the modifiability of every part. Nor is this true of individuals only: it holds, in a sense, of species. Leaving open the question whether, in indefinite times, indefinite modifications may not be produced by inheritance of functionally wrought adaptations; experience proves that within assigned times, the changes wrought in races of organisms by changes of conditions fall within narrow limits. Though by discipline, aided by ​selective breeding, one variety of horse has had its locomotive power increased considerably beyond the locomotive powers of other varieties; yet further increase takes place, if at all, at an inappreciable rate. The different kinds of dogs, too, in which different forms and capacities have been established, do not now show aptitudes for diverging in the same directions at considerable rates. In domestic animals generally, certain accessions of intelligence have been produced by culture; but accessions beyond these are inconspicuous. It seems that in each species of organism there is a margin for functional oscillations on all sides of a mean state, and a consequent margin for structural variations; that it is possible rapidly to push functional and structural changes towards the extreme of this margin in any direction, both in an individual and in a race; but that to push these changes further in any direction, and so to alter the organism as to bring its mean state up to the extreme of the margin in that direction, is a comparatively slow process.
We also have to note that the limited increase of size produced in any organ by a limited increase of its function, is not maintained unless the increase of function is permanent. A mature man or other animal, led by circumstances into exerting particular members in unusual degrees, and acquiring extra sizes in these members, begins to lose such extra sizes on ceasing to exert the members; and eventually lapses more or less nearly into the original state. Legs strengthened by a pedestrian tour, become relatively weak again after a prolonged return to sedentary life. The acquired ability to perform feats of skill disappears in course of time, if the performance of them be given up. For comparative failure in executing a piece of music, in playing a game at chess, or in anything requiring special culture, the being out of practice ​is a reason which every one recognizes as valid. It is observable, too, that the rapidity and completeness with which an artificial power is lost, is proportionate to the shortness of the cultivation which evoked it. One who has for many years persevered in habits which exercise special muscles or special faculties of mind, retains the extra capacity produced, to a very considerable degree, even after a long period of desistance; but one who has persevered in such habits for but a short time has, at the end of a like period, scarcely any of the facility he had gained. Here too, as before, successions of organisms present an analogous fact. A species in which domestication continued through many generations, has organized certain peculiarities; and which afterwards, escaping domestic discipline, returns to something like its original habits; soon loses, in great measure, such peculiarities. Though it is not true, as alleged, that it resumes completely the structure it had before domestication, yet it approximates to that structure. The Dingo, or wild dog of Australia, is one of the instances given of this; and the wild horse of South America is another. Mankind, too, supplies us with instances. In the Australian bush and in the backwoods of America, the Anglo-Saxon race, in which civilization has developed the higher feelings to a considerable degree, rapidly lapses into comparative barbarism: adopting the moral code, and sometimes the habits, of savages.
§ 68. It is important to reach, if possible, some rationale of these general truths—especially of the last two. A right understanding of these laws of organic modification underlies a right understanding of the great question of species. While, as before hinted ( § 40), the action of structure on function is one of the factors in that process of differentiation by which unlike forms of plants and animals are produced, the reaction of function on structure is another factor. Hence, it is well worth while inquiring how far these inductions are deductively interpretable.
​The first of them is the most difficult to deal with. Why an organ exerted somewhat beyond its wont should presently grow, and thus meet increase of demand by increase of supply, is not obvious. We know, indeed, (First Principles, §§ 85, 173,) that of necessity, the rhythmical changes produced by antagonistic organic actions cannot any of them be carried to an excess in one direction, without there being produced an equivalent excess in the opposite direction. It is a corollary from the persistence of force, that any deviation effected by a disturbing cause, acting on some member of a moving equilibrium, must (unless it altogether destroys the moving equilibrium) be eventually followed by a compensating deviation. Hence, that excess of repair should succeed excess of waste, is to be expected. But how happens the mean state of the organ to be changed? If daily extra waste naturally brings about daily extra repair only to an equivalent extent, the mean state of the organ should remain constant. How then comes the organ to augment in size and power?
Such answer to this question as we may hope to find, must be looked for in the effects wrought on the organism as a whole by increased function in one of its parts. For since the discharge of its function by any part is possible only on condition that those various other functions on which its own is immediately dependent are also discharged, it follows that excess in its function presupposes some excess in their functions. Additional work given to a muscle implies additional work given to the branch arteries which bring it blood, and additional work, smaller in proportion, to the arteries from which these branch arteries come. Similarly, the smaller and larger veins which take away the blood, as well as those structures which deal with effete products, must have more to do. And yet further, on the nervous centres which excite the muscle a certain extra duty must fall. But excess of waste will entail excess of repair, in these parts as well as in the muscle. The several appliances by which the nutrition ​and excitation of an organ are carried on, must also be influenced by this rhythm of action and reaction; and therefore, after losing more than usual by the destructive process they must gain more than usual by the constructive process. But temporarily-increased efficiency in these appliances by which blood and nervous force are brought to an organ, will cause extra assimilation in the organ, beyond that required to balance its extra expenditure. Regarding the functions as constituting a moving equilibrium, we may say that divergence of any function in the direction of increase, causes the functions with which it is bound up to diverge in the same direction; that these, again, cause the functions which they are bound up with, also to diverge in the same direction; and that these divergences of the connected functions allow the specially-affected function to be carried further in this direction than it could otherwise be—further than the perturbing force could carry it if it had a fixed basis.
It must be admitted that this is but a vague explanation. Among actions so involved as these, we can scarcely expect to do more than dimly discern a harmony with first principles. That the facts are to be interpreted in some such way, may, however, be inferred from the circumstance that an extra supply of blood continues for some time to be sent to an organ that has been unusually exercised; and that when unusual exercise is long continued a permanent increase of vascularity results.
§ 69. Answers to the questions—Why do these adaptive modifications in an individual animal soon reach a limit? and why, in the descendants of such animal, similarly conditioned, is this limit very slowly extended?—are to be found in the same direction as was the answer to the last question. And here the connexion of cause and consequence is more manifest.
Since the function of any organ is dependent on the functions of the organs which supply it with materials and ​stimuli; and since the functions of these subsidiary organs are dependent on the functions of organs which supply them with materials and stimuli; it follows that before any great extra power of discharging its function can be gained by a specially-exercised organ, a considerable extra power must be gained by a series of immediately-subservient organs, and some extra power by a secondary series of remotely-subservient organs. Thus there are required numerous and wide-spread modifications. Before the artery which feeds a hard-worked muscle can permanently furnish a large additional quantity of blood, it must increase in diameter; and that its increase of diameter may be of use, the main artery from which it diverges must also be so far modified as to bring this additional quantity of blood to the branch artery. Similarly with the veins; similarly with the structures which remove waste-products; similarly with the nerves. And when we ask what these subsidiary changes imply, we are forced to conclude that there must be an analogous group of more numerous changes ramifying throughout the system. The growth of the arteries primarily and secondarily implicated, cannot go to any extent without growth in the minor blood-vessels on which their nutrition depends; while their greater contractile power involves enlargement of the nerves which excite them, and some modification of that part of the spinal cord whence these nerves proceed. Thus, without tracing the like remote alterations implied by extra growth of the veins, lymphatics, glandular organs, and other agencies, it is manifest that a large amount of rebuilding must be done throughout the organism, before any organ of importance can be permanently increased in size and power to a great extent. Hence, though such extra growth in any part as does not necessitate considerable changes throughout the rest of the organism, may rapidly take place; a further growth in this part, requiring a re-modelling of numerous parts remotely and slightly affected, must take place but slowly.
​We have before found our conceptions of vital processes made clearer by studying analogous social processes. In societies there is a mutual dependence of functions, essentially like that which exists in organisms; and there is also an essentially like reaction of functions on structures. From the laws of adaptive modification in societies, we may therefore hope to get a clue to the laws of adaptive modification in organisms. Let us suppose, then, that a society has arrived at a state of equilibrium analogous to that of a mature animal—a state not like our own, in which growth and structural development are rapidly going on, but a state of settled balance among the functional powers of the various classes and industrial bodies, and a consequent fixity in the relative sizes of such classes and bodies. Further, let us suppose that in a society thus balanced there occurs something which throws an unusual demand on one industry—say an unusual demand for ships (which we will assume to be built of iron) in consequence of a competing mercantile nation having been prostrated by famine or pestilence. The immediate result of this additional demand for iron ships is the employment of more workmen, and the purchase of more iron, by the ship-builders; and when, presently, the demand continuing, the ship-builders find their premises and machinery insufficient, they enlarge them. If the extra requirement persists, the high interest and high wages bring such extra capital and labour into the business as are needed for new ship-building establishments. But such extra capital and labour do not come quickly; since, in a balanced community, not increasing in population and wealth, labour and capital have to be drawn from other industries, where they are already yielding the ordinary returns. Let us now go a step further. Suppose that this iron-ship-building industry, having enlarged as much as the available capital and labour permit, is still unequal to the demand; what limits its immediate further growth? The lack of iron. By the hypothesis, the iron-producing industry, like all the ​other industries throughout the community, yields only as much iron as is habitually required for all the purposes to which iron is applied: ship-building being only one. If, then, extra iron is required for ship-building, the first effect is to withdraw part of the iron habitually consumed for other purposes, and to raise the price of iron. Presently, the iron-makers feel this change and their stocks dwindle. As, however, the quantity of iron required for ship-building forms but a small part of the total quantity required for all purposes, the extra demand on the iron-makers can be nothing like so great in proportion as is the extra demand on the ship-builders. Whence it follows that there will be much less tendency to an immediate enlargement of the iron-producing industry; since the extra quantity will for some time be obtained by working extra hours. Nevertheless if, as fast as more iron can be thus supplied, the ship-building industry goes on growing—if, consequently, the iron-makers experience a permanently-increased demand, and out of their greater profits get higher interest on capital, as well as pay higher wages; there will eventually be an abstraction of capital and labour from other industries to enlarge the iron-producing industry: new blast-furnaces, new rolling-mills, new cottages for workmen, will be erected. But obviously, the inertia of capital and labour to be overcome before the iron-producing industry can grow by a decrease of certain other industries, will prevent its growth from taking place until long after the increased ship-building industry has demanded it; and meanwhile, the growth of the ship-building industry must be limited by the deficiency of iron. A remoter restraint of the same nature meets us if we go a step further—a restraint which can be overcome only in a still longer time. For the manufacture of iron depends on the supply of coal. The production of coal being previously in equilibrium with the consumption; and the consumption of coal for the manufacture of iron being but a small part of the total consumption; it follows that a ​considerable extension of the iron manufacture, when it at length takes place, will cause but a comparatively small additional demand on the coal-owners and coal-miners—a demand which will not, for a long period, suffice to cause enlargement of the coal-trade, by drawing capital and labour from other investments and occupations. And until the permanent extra demand for coal has become great enough to draw from other investments and occupations sufficient capital and labour to sink new mines, the increasing production of iron must be restricted by the scarcity of coal, and the multiplication of ship-yards and ship-builders must be checked by the want of iron. Thus, in a community which has reached a state of moving equilibrium, though any one industry directly affected by an additional demand may rapidly undergo a small extra growth, yet a growth beyond this, requiring as it does the building-up of subservient industries, less directly and strongly affected, as well as the partial unbuilding of other industries, can take place only with comparative slowness. And a still further growth, requiring structural modifications of industries still more distantly affected, must take place still more slowly.
On returning from this analogy, we see more clearly the truth that any considerable member of an animal organism, cannot be greatly enlarged without some general reorganization. Besides a building up of the primary, secondary, and tertiary groups of the subservient parts, there must be an unbuilding of sundry non-subservient parts; or, at any rate, there must be permanently established a lower nutrition of such non-subservient parts. For it must be remembered that in a mature animal, or one which has reached a balance between assimilation and expenditure, there cannot (supposing general conditions to remain constant) be an increase in the nutrition of some organs without a decrease in the nutrition of others; and an organic establishment of the increase implies an organic establishment of the decrease—implies more or less change in the processes and structures ​throughout the entire system. And here, indeed, is disclosed one reason why growing animals undergo adaptations so much more readily than adult ones. For while there is surplus nutrition, it is possible for specially-exercised parts to be specially enlarged without any positive deduction from other parts. There is required only that negative deduction implied in the diminished growth of other parts.
§ 70. Pursuing the argument further, we reach an explanation of the third general truth; namely that organisms, and species of organisms, which, under new conditions, have undergone adaptive modifications, soon return to something like their original structures when restored to their original conditions. Seeing, as we have done, how excess of action and excess of nutrition in any part of an organism, must affect action and nutrition in subservient parts, and these again in other parts, until the re-action has divided and subdivided itself throughout the organism, affecting in decreasing degrees the more and more numerous parts more and more remotely implicated; we see that the consequent changes in the parts remotely implicated, constituting the great mass of the organism, must be extremely slow. Hence, if the need for the adaptive modification ceases before the great mass of the organism has been much altered in its structure by these ramified but minute reactions, we shall have a condition in which the specially-modified part is not in equilibrium with the rest. All the remotely-affected organs, as yet but little changed, will, in the absence of the perturbing cause, resume very nearly their previous actions. The parts that depend on them will consequently by and by do the same. Until at length, by a reversal of the adaptive process, the organ at first affected will be brought back almost to its original state. Reconsidering the above-drawn analogy between an organism and a society, will enable us better to recognize this necessity. If, in the case supposed, the extra demand for iron ships, after causing ​the erection of some additional ship-yards and the drawing of iron from other manufactures, were to cease; the old dimensions of the ship-building trade would be quickly returned to: discharged workmen would seek fresh occupations, and the new yards would be devoted to other uses. But if the increased need for ships lasted long enough, and became great enough, to cause a flow of capital and labour from other industries into the iron-manufacture, a falling off in the demand for ships, would much less rapidly entail a dwindling of the ship-building industry. For iron being now produced in greater quantity, a diminished consumption of it for ships would cause a fall in its price, and a consequent fall in the cost of ships: thus enabling the ship-builders to meet the competition which we may suppose led to a decrease in the orders they received. And since, when new blast-furnaces and rolling-mills, &c., had been built with capital drawn from other industries, its transference back into other industries would involve great loss; the owners, rather than transfer it, would accept unusually low interest, and an excess of iron would continue to be produced; resulting in an undue cheapness of ships, and a maintenance of the ship-building industry at a size beyond the need. Eventually, however, if the number of ships required still diminished, the production of iron in excess would become very unremunerative: some of the blast-furnaces would be blown out; and as much of the capital and labour as remained available would be re-distributed among other occupations. Without repeating the steps of the argument, it will be clear that were the enlargement of the ship-building industry great enough, and did it last long enough to cause an increase in the number of coal-mines, the ship-building industry would be still better able to maintain itself under adverse circumstances; but that it would, though at a more distant period, end by sinking down to the needful dimensions. Thus our conclusions are:—First, that if the extra growth caused by extra activity in a particular industry has lasted long enough ​only to remodel the proximately-affected industries; it will dwindle away again after a moderate period, if the need for it disappears. Second, that a long period must be required before the re-actions produced by an enlarged industry can cause a re-construction of the whole society, and before the countless re-distributions of capital and labour can again reach a state of equilibrium. And third, that only when such a new state of equilibrium is eventually reached, can the adaptive modification become a permanent one. How, in animal organisms the like argument holds, need not be pointed out. The reader will readily follow the parallel.
That organic types should be comparatively stable, might be anticipated on the hypothesis of Evolution. The structure of any organism being a product of the almost infinite series of actions and reactions to which ancestral organisms have been exposed; any unusual actions and reactions brought to bear on an individual, can have but an infinitesimal effect in permanently changing the structure of the organism as a whole. The new set of forces, compounded with all the antecedent sets of forces, can but inappreciably modify that moving equilibrium of functions which all these antecedent sets of forces have established. Though there may result a considerable perturbation of certain functions—a considerable divergence from their ordinary rhythms—yet the general centre of equilibrium cannot be sensibly changed. On the removal of the perturbing cause the previous balance will be quickly restored: the effect of the new forces being almost obliterated by the enormous aggregate of forces which the previous balance expresses.
§ 71. As thus understood, the phenomena of adaptation fall into harmony with first principles. The inference that organic types are fixed, because the deviations from them which can be produced within assignable periods are relatively small, and because, when a force producing deviation ceases, there is a return to something like the original state; proves to ​be an invalid inference. Without assuming fixity of species, we find good reasons for anticipating that kind and degree of stability which is observed. We find grounds for concluding, a priori, that an adaptive change of structure will soon reach a point beyond which further adaptation will be slow; for concluding that when the modifying cause has been but a short time in action, the modification generated will be evanescent; for concluding that a modifying cause acting even for many generations, will do but little towards permanently altering the organic equilibrium of a race; and for concluding that on the cessations of such cause, its effects will become unapparent in the course of a few generations.
* ↑ In the account of James Mitchell, a boy born blind and deaf, given by James Wardrop, F.R.S. (Edin. 1813), it is said that he acquired a "preternatural acuteness of touch and smell." The deaf Dr. Kitto described himself as having an extremely strong visual memory: he retained "a clear impression or image of everything at which he ever looked."
* ↑ Here, as in sundry places throughout this chapter, the necessities of the argument have obliged me to forestall myself, by assuming the conclusion reached in a subsequent chapter, that modifications of structure produced by modifications of function are transmitted to offspring.