Origin of Species by Charles Darwin
Chapter IV. Natural Selection; or the Survival of the Fittest
How will the struggle for existence, briefly discussed in the last chapter, act in regard to variation? Can the principle of selection, which we have seen is so potent in the hands of man, apply under nature? I think we shall see that it can act most efficiently. Let the endless number of slight variations and individual differences occurring in our domestic productions, and, in a lesser degree, in those under nature, be borne in mind; as well as the strength of the hereditary tendency. Under domestication, it may truly be said that the whole organisation becomes in some degree plastic. But the variability, which we almost universally meet with in our domestic productions is not directly produced, as Hooker and Asa Gray have well remarked, by man; he can neither originate varieties nor prevent their occurrence; he can only preserve and accumulate such as do occur. Unintentionally he exposes organic beings to new and changing conditions of life, and variability ensues; but similar changes of conditions might and do occur under nature. Let it also be borne in mind how infinitely complex and close-fitting are the mutual relations of all organic beings to each other and to their physical conditions of life; and consequently what infinitely varied diversities of structure might be of use to each being under changing conditions of life. Can it then be thought improbable, seeing that variations useful to man have undoubtedly occurred, that other variations useful in some way to each being in the great and complex battle of life, should occur in the course of many successive generations? If such do occur, can we doubt (remembering that many more individuals are born than can possibly survive) that individuals having any advantage, however slight, over others, would have the best chance of surviving and procreating their kind? On the other hand, we may feel sure that any variation in the least degree injurious would be rigidly destroyed. This preservation of favourable individual differences and variations, and the destruction of those which are injurious, I have called Natural Selection, or the Survival of the Fittest. Variations neither useful nor injurious would not be affected by natural selection, and would be left either a fluctuating element, as perhaps we see in certain polymorphic species, or would ultimately become fixed, owing to the nature of the organism and the nature of the conditions.
Several writers have misapprehended or objected to the term Natural Selection. Some have even imagined that natural selection induces variability, whereas it implies only the preservation of such variations as arise and are beneficial to the being under its conditions of life. No one objects to agriculturists speaking of the potent effects of man's selection; and in this case the individual differences given by nature, which man for some object selects, must of necessity first occur. Others have objected that the term selection implies conscious choice in the animals which become modified; and it has even been urged that, as plants have no volition, natural selection is not applicable to them! In the literal sense of the word, no doubt, natural selection is a false term; but who ever objected to chemists speaking of the elective affinities of the various elements?--and yet an acid cannot strictly be said to elect the base with which it in preference combines. It has been said that I speak of natural selection as an active power or Deity; but who objects to an author speaking of the attraction of gravity as ruling the movements of the planets? Every one knows what is meant and is implied by such metaphorical expressions; and they are almost necessary for brevity. So again it is difficult to avoid personifying the word Nature; but I mean by nature, only the aggregate action and product of many natural laws, and by laws the sequence of events as ascertained by us. With a little familiarity such superficial objections will be forgotten.
We shall best understand the probable course of natural selection by taking the case of a country undergoing some slight physical change, for instance, of climate. The proportional numbers of its inhabitants will almost immediately undergo a change, and some species will probably become extinct. We may conclude, from what we have seen of the intimate and complex manner in which the inhabitants of each country are bound together, that any change in the numerical proportions of the inhabitants, independently of the change of climate itself, would seriously affect the others. If the country were open on its borders, new forms would certainly immigrate, and this would likewise seriously disturb the relations of some of the former inhabitants. Let it be remembered how powerful the influence of a single introduced tree or mammal has been shown to be. But in the case of an island, or of a country partly surrounded by barriers, into which new and better adapted forms could not freely enter, we should then have places in the economy of nature which would assuredly be better filled up if some of the original inhabitants were in some manner modified; for, had the area been open to immigration, these same places would have been seized on by intruders. In such cases, slight modifications, which in any way favoured the individuals of any species, by better adapting them to their altered conditions, would tend to be preserved; and natural selection would have free scope for the work of improvement.
We have good reason to believe, as shown in the first chapter, that changes in the conditions of life give a tendency to increased variability; and in the foregoing cases the conditions the changed, and this would manifestly be favourable to natural selection, by affording a better chance of the occurrence of profitable variations. Unless such occur, natural selection can do nothing. Under the term of "variations," it must never be forgotten that mere individual differences are included. As man can produce a great result with his domestic animals and plants by adding up in any given direction individual differences, so could natural selection, but far more easily from having incomparably longer time for action. Nor do I believe that any great physical change, as of climate, or any unusual degree of isolation, to check immigration, is necessary in order that new and unoccupied places should be left for natural selection to fill up by improving some of the varying inhabitants. For as all the inhabitants of each country are struggling together with nicely balanced forces, extremely slight modifications in the structure or habits of one species would often give it an advantage over others; and still further modifications of the same kind would often still further increase the advantage, as long as the species continued under the same conditions of life and profited by similar means of subsistence and defence. No country can be named in which all the native inhabitants are now so perfectly adapted to each other and to the physical conditions under which they live, that none of them could be still better adapted or improved; for in all countries, the natives have been so far conquered by naturalised productions that they have allowed some foreigners to take firm possession of the land. And as foreigners have thus in every country beaten some of the natives, we may safely conclude that the natives might have been modified with advantage, so as to have better resisted the intruders.
As man can produce, and certainly has produced, a great result by his methodical and unconscious means of selection, what may not natural selection effect? Man can act only on external and visible characters: Nature, if I may be allowed to personify the natural preservation or survival of the fittest, cares nothing for appearances, except in so far as they are useful to any being. She can act on every internal organ, on every shade of constitutional difference, on the whole machinery of life. Man selects only for his own good; Nature only for that of the being which she tends. Every selected character is fully exercised by her, as is implied by the fact of their selection. Man keeps the natives of many climates in the same country. He seldom exercises each selected character in some peculiar and fitting manner; he feeds a long and a short-beaked pigeon on the same food; he does not exercise a long-backed or long-legged quadruped in any peculiar manner; he exposes sheep with long and short wool to the same climate; does not allow the most vigorous males to struggle for the females; he does not rigidly destroy all inferior animals, but protects during each varying season, as far as lies in his power, all his productions. He often begins his selection by some half-monstrous form, or at least by some modification prominent enough to catch the eye or to be plainly useful to him. Under nature, the slightest differences of structure or constitution may well turn the nicely-balanced scale in the struggle for life, and so be preserved. How fleeting are the wishes and efforts of man! How short his time, and consequently how poor will be his results, compared with those accumulated by Nature during whole geological periods! Can we wonder, then, that Nature's productions should be far "truer" in character than man's productions; that they should be infinitely better adapted to the most complex conditions of life, and should plainly bear the stamp of far higher workmanship?
It may metaphorically be said that natural selection is daily and hourly scrutinising, throughout the world, the slightest variations; rejecting those that are bad, preserving and adding up all that are good; silently and insensibly working, whenever and wherever opportunity offers, at the improvement of each organic being in relation to its organic and inorganic conditions of life. We see nothing of these slow changes in progress, until the hand of time has marked the long lapse of ages, and then so imperfect is our view into long-past geological ages that we see only that the forms of life are now different from what they formerly were.
In order that any great amount of modification should be effected in a species, a variety, when once formed must again, perhaps after a long interval of time, vary or present individual differences of the same favourable nature as before; and these must again be preserved, and so onward, step by step. Seeing that individual differences of the same kind perpetually recur, this can hardly be considered as an unwarrantable assumption. But whether it is true, we can judge only by seeing how far the hypothesis accords with and explains the general phenomena of nature. On the other hand, the ordinary belief that the amount of possible variation is a strictly limited quantity, is likewise a simple assumption.
Although natural selection can act only through and for the good of each being, yet characters and structures, which we are apt to consider as of very trifling importance, may thus be acted on. When we see leaf-eating insects green, and bark-feeders mottled-grey; the alpine ptarmigan white in winter, the red-grouse the colour of heather, we must believe that these tints are of service to these birds and insects in preserving them from danger. Grouse, if not destroyed at some period of their lives, would increase in countless numbers; they are known to suffer largely from birds of prey; and hawks are guided by eyesight to their prey,--so much so that on parts of the continent persons are warned not to keep white pigeons, as being the most liable to destruction. Hence natural selection might be effective in giving the proper colour to each kind of grouse, and in keeping that colour, when once acquired, true and constant. Nor ought we to think that the occasional destruction of an animal of any particular colour would produce little effect; we should remember how essential it is in a flock of white sheep to destroy a lamb with the faintest trace of black. We have seen how the colour of hogs, which feed on the "paint-root" in Virginia, determines whether they shall live or die. In plants, the down on the fruit and the colour of the flesh are considered by botanists as characters of the most trifling importance; yet we hear from an excellent horticulturist, Downing, that in the United States smooth-skinned fruits suffer far more from a beetle, a Curculio, than those with down; that purple plums suffer far more from a certain disease than yellow plums; whereas another disease attacks yellow-fleshed peaches far more than those with other coloured flesh. If, with all the aids of art, these slight differences make a great difference in cultivating the several varieties, assuredly, in a state of nature, where the trees would have to struggle with other trees and with a host of enemies, such differences would effectually settle which variety, whether a smooth or downy, a yellow or a purple-fleshed fruit, should succeed.
In looking at many small points of difference between species, which, as far as our ignorance permits us to judge, seem quite unimportant, we must not forget that climate, food, etc., have no doubt produced some direct effect. It is also necessary to bear in mind that, owing to the law of correlation, when one part varies and the variations are accumulated through natural selection, other modifications, often of the most unexpected nature, will ensue.
As we see that those variations which, under domestication, appear at any particular period of life, tend to reappear in the offspring at the same period; for instance, in the shape, size and flavour of the seeds of the many varieties of our culinary and agricultural plants; in the caterpillar and cocoon stages of the varieties of the silkworm; in the eggs of poultry, and in the colour of the down of their chickens; in the horns of our sheep and cattle when nearly adult; so in a state of nature natural selection will be enabled to act on and modify organic beings at any age, by the accumulation of variations profitable at that age, and by their inheritance at a corresponding age. If it profit a plant to have its seeds more and more widely disseminated by the wind, I can see no greater difficulty in this being effected through natural selection, than in the cotton-planter increasing and improving by selection the down in the pods on his cotton-trees. Natural selection may modify and adapt the larva of an insect to a score of contingencies, wholly different from those which concern the mature insect; and these modifications may affect, through correlation, the structure of the adult. So, conversely, modifications in the adult may affect the structure of the larva; but in all cases natural selection will ensure that they shall not be injurious: for if they were so, the species would become extinct.
Natural selection will modify the structure of the young in relation to the parent and of the parent in relation to the young. In social animals it will adapt the structure of each individual for the benefit of the whole community; if the community profits by the selected change. What natural selection cannot do, is to modify the structure of one species, without giving it any advantage, for the good of another species; and though statements to this effect may be found in works of natural history, I cannot find one case which will bear investigation. A structure used only once in an animal's life, if of high importance to it, might be modified to any extent by natural selection; for instance, the great jaws possessed by certain insects, used exclusively for opening the cocoon--or the hard tip to the beak of unhatched birds, used for breaking the eggs. It has been asserted, that of the best short-beaked tumbler-pigeons a greater number perish in the egg than are able to get out of it; so that fanciers assist in the act of hatching. Now, if nature had to make the beak of a full-grown pigeon very short for the bird's own advantage, the process of modification would be very slow, and there would be simultaneously the most rigorous selection of all the young birds within the egg, which had the most powerful and hardest beaks, for all with weak beaks would inevitably perish: or, more delicate and more easily broken shells might be selected, the thickness of the shell being known to vary like every other structure.
It may be well here to remark that with all beings there must be much fortuitous destruction, which can have little or no influence on the course of natural selection. For instance, a vast number of eggs or seeds are annually devoured, and these could be modified through natural selection only if they varied in some manner which protected them from their enemies. Yet many of these eggs or seeds would perhaps, if not destroyed, have yielded individuals better adapted to their conditions of life than any of those which happened to survive. So again a vast number of mature animals and plants, whether or not they be the best adapted to their conditions, must be annually destroyed by accidental causes, which would not be in the least degree mitigated by certain changes of structure or constitution which would in other ways be beneficial to the species. But let the destruction of the adults be ever so heavy, if the number which can exist in any district be not wholly kept down by such causes--or again let the destruction of eggs or seeds be so great that only a hundredth or a thousandth part are developed--yet of those which do survive, the best adapted individuals, supposing that there is any variability in a favourable direction, will tend to propagate their kind in larger numbers than the less well adapted. If the numbers be wholly kept down by the causes just indicated, as will often have been the case, natural selection will be powerless in certain beneficial directions; but this is no valid objection to its efficiency at other times and in other ways; for we are far from having any reason to suppose that many species ever undergo modification and improvement at the same time in the same area.