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полная версияThe Foundations of the Origin of Species

Чарльз Дарвин
The Foundations of the Origin of Species

Variation in the mental attributes of wild animals

With respect to the variation284 of the mental powers of animals in a wild state, we know that there is a considerable difference in the disposition of different individuals of the same species, as is recognised by all those who have had the charge of animals in a menagerie. With respect to the wildness of animals, that is fear directed particularly against man, which appears to be as true an instinct as the dread of a young mouse of a cat, we have excellent evidence that it is slowly acquired and becomes hereditary. It is also certain that, in a natural state, individuals of the same species lose or do not practice their migratory instincts – as woodcocks in Madeira. With respect to any variation in the more complicated instincts, it is obviously most difficult to detect, even more so than in the case of corporeal structure, of which it has been admitted the variation is exceedingly small, and perhaps scarcely any in the majority of species at any one period. Yet, to take one excellent case of instinct, namely the nests of birds, those who have paid most attention to the subject maintain that not only certain individuals «? species» seem to be able to build very imperfectly, but that a difference in skill may not unfrequently be detected between individuals285. Certain birds, moreover, adapt their nests to circumstances; the water-ouzel makes no vault when she builds under cover of a rock – the sparrow builds very differently when its nest is in a tree or in a hole, and the golden-crested wren sometimes suspends its nest below and sometimes places it on the branches of trees.

Principles of Selection applicable to instincts

As the instincts of a species are fully as important to its preservation and multiplication as its corporeal structure, it is evident that if there be the slightest congenital differences in the instincts and habits, or if certain individuals during their lives are induced or compelled to vary their habits, and if such differences are in the smallest degree more favourable, under slightly modified external conditions, to their preservation, such individuals must in the long run have a better chance of being preserved and of multiplying286. If this be admitted, a series of small changes may, as in the case of corporeal structure, work great changes in the mental powers, habits and instincts of any species.

Difficulties in the acquirement of complex instincts by Selection

Every one will at first be inclined to explain (as I did for a long time) that many of the more complicated and wonderful instincts could not be acquired in the manner here supposed287. The Second Part of this work is devoted to the general consideration of how far the general economy of nature justifies or opposes the belief that related species and genera are descended from common stocks; but we may here consider whether the instincts of animals offer such a primâ facie case of impossibility of gradual acquirement, as to justify the rejection of any such theory, however strongly it may be supported by other facts. I beg to repeat that I wish here to consider not the probability but the possibility of complicated instincts having been acquired by the slow and long-continued selection of very slight (either congenital or produced by habit) modifications of foregoing simpler instincts; each modification being as useful and necessary, to the species practising it, as the most complicated kind.

First, to take the case of birds’-nests; of existing species (almost infinitely few in comparison with the multitude which must have existed, since the period of the new Red Sandstone of N. America, of whose habits we must always remain ignorant) a tolerably perfect series could be made from eggs laid on the bare ground, to others with a few sticks just laid round them, to a simple nest like the wood-pigeons, to others more and more complicated: now if, as is asserted, there occasionally exist slight differences in the building powers of an individual, and if, which is at least probable, that such differences would tend to be inherited, then we can see that it is at least possible that the nidificatory instincts may have been acquired by the gradual selection, during thousands and thousands of generations, of the eggs and young of those individuals, whose nests were in some degree better adapted to the preservation of their young, under the then existing conditions. One of the most surprising instincts on record is that of the Australian bush-turkey, whose eggs are hatched by the heat generated from a huge pile of fermenting materials, which it heaps together; but here the habits of an allied species show how this instinct might possibly have been acquired. This second species inhabits a tropical district, where the heat of the sun is sufficient to hatch its eggs; this bird, burying its eggs, apparently for concealment, under a lesser heap of rubbish, but of a dry nature, so as not to ferment. Now suppose this bird to range slowly into a climate which was cooler, and where leaves were more abundant, in that case, those individuals, which chanced to have their collecting instinct strongest developed, would make a somewhat larger pile, and the eggs, aided during some colder season, under the slightly cooler climate by the heat of incipient fermentation, would in the long run be more freely hatched and would probably produce young ones with the same more highly developed collecting tendencies; of these again, those with the best developed powers would again tend to rear most young. Thus this strange instinct might possibly be acquired, every individual bird being as ignorant of the laws of fermentation, and the consequent development of heat, as we know they must be.

Secondly, to take the case of animals feigning death (as it is commonly expressed) to escape danger. In the case of insects, a perfect series can be shown, from some insects, which momentarily stand still, to others which for a second slightly contract their legs, to others which will remain immovably drawn together for a quarter of an hour, and may be torn asunder or roasted at a slow fire, without evincing the smallest sign of sensation. No one will doubt that the length of time, during which each remains immovable, is well adapted to «favour the insect’s» escape «from» the dangers to which it is most exposed, and few will deny the possibility of the change from one degree to another, by the means and at the rate already explained. Thinking it, however, wonderful (though not impossible) that the attitude of death should have been acquired by methods which imply no imitation, I compared several species, when feigning, as is said, death, with others of the same species really dead, and their attitudes were in no one case the same.

Thirdly, in considering many instincts it is useful to endeavour to separate the faculty288 by which they perform it, and the mental power which urges to the performance, which is more properly called an instinct. We have an instinct to eat, we have jaws &c. to give us the faculty to do so. These faculties are often unknown to us: bats, with their eyes destroyed, can avoid strings suspended across a room, we know not at present by what faculty they do this. Thus also, with migratory birds, it is a wonderful instinct which urges them at certain times of the year to direct their course in certain directions, but it is a faculty by which they know the time and find their way. With respect to time289, man without seeing the sun can judge to a certain extent of the hour, as must those cattle which come down from the inland mountains to feed on sea-weed left bare at the changing hour of low-water290. A hawk (D’Orbigny) seems certainly to have acquired a knowledge of a period of every 21 days. In the cases already given of the sheep which travelled to their birth-place to cast their lambs, and the sheep in Spain which know their time of march291, we may conjecture that the tendency to move is associated, we may then call it instinctively, with some corporeal sensations. With respect to direction we can easily conceive how a tendency to travel in a certain course may possibly have been acquired, although we must remain ignorant how birds are able to preserve any direction whatever in a dark night over the wide ocean. I may observe that the power of some savage races of mankind to find their way, although perhaps wholly different from the faculty of birds, is nearly as unintelligible to us. Bellinghausen, a skilful navigator, describes with the utmost wonder the manner in which some Esquimaux guided him to a certain point, by a course never straight, through newly formed hummocks of ice, on a thick foggy day, when he with a compass found it impossible, from having no landmarks, and from their course being so extremely crooked, to preserve any sort of uniform direction: so it is with Australian savages in thick forests. In North and South America many birds slowly travel northward and southward, urged on by the food they find, as the seasons change; let them continue to do this, till, as in the case of the sheep in Spain, it has become an urgent instinctive desire, and they will gradually accelerate their journey. They would cross narrow rivers, and if these were converted by subsidence into narrow estuaries, and gradually during centuries to arms of the sea, still we may suppose their restless desire of travelling onwards would impel them to cross such an arm, even if it had become of great width beyond their span of vision. How they are able to preserve a course in any direction, I have said, is a faculty unknown to us. To give another illustration of the means by which I conceive it possible that the direction of migrations have been determined. Elk and reindeer in N. America annually cross, as if they could marvellously smell or see at the distance of a hundred miles, a wide tract of absolute desert, to arrive at certain islands where there is a scanty supply of food; the changes of temperature, which geology proclaims, render it probable that this desert tract formerly supported some vegetation, and thus these quadrupeds might have been annually led on, till they reached the more fertile spots, and so acquired, like the sheep of Spain, their migratory powers.

 

Fourthly, with respect to the combs of the hive-bee292; here again we must look to some faculty or means by which they make their hexagonal cells, without indeed we view these instincts as mere machines. At present such a faculty is quite unknown: Mr Waterhouse supposes that several bees are led by their instinct to excavate a mass of wax to a certain thinness, and that the result of this is that hexagons necessarily remain. Whether this or some other theory be true, some such means they must possess. They abound, however, with true instincts, which are the most wonderful that are known. If we examine the little that is known concerning the habits of other species of bees, we find much simpler instincts: the humble bee merely fills rude balls of wax with honey and aggregates them together with little order in a rough nest of grass. If we knew the instinct of all the bees, which ever had existed, it is not improbable that we should have instincts of every degree of complexity, from actions as simple as a bird making a nest, and rearing her young, to the wonderful architecture and government of the hive-bee; at least such is possible, which is all that I am here considering.

Finally, I will briefly consider under the same point of view one other class of instincts, which have often been advanced as truly wonderful, namely parents bringing food to their young which they themselves neither like nor partake of293; – for instance, the common sparrow, a granivorous bird, feeding its young with caterpillars. We might of course look into the case still earlier, and seek how an instinct in the parent, of feeding its young at all, was first derived; but it is useless to waste time in conjectures on a series of gradations from the young feeding themselves and being slightly and occasionally assisted in their search, to their entire food being brought to them. With respect to the parent bringing a different kind of food from its own kind, we may suppose either that the remote stock, whence the sparrow and other congenerous birds have descended, was insectivorous, and that its own habits and structure have been changed, whilst its ancient instincts with respect to its young have remained unchanged; or we may suppose that the parents have been induced to vary slightly the food of their young, by a slight scarcity of the proper kind (or by the instincts of some individuals not being so truly developed), and in this case those young which were most capable of surviving were necessarily most often preserved, and would themselves in time become parents, and would be similarly compelled to alter their food for their young. In the case of those animals, the young of which feed themselves, changes in their instincts for food, and in their structure, might be selected from slight variations, just as in mature animals. Again, where the food of the young depends on where the mother places her eggs, as in the case of the caterpillars of the cabbage-butterfly, we may suppose that the parent stock of the species deposited her eggs sometimes on one kind and sometimes on another of congenerous plants (as some species now do), and if the cabbage suited the caterpillars better than any other plant, the caterpillars of those butterflies, which had chosen the cabbage, would be most plentifully reared, and would produce butterflies more apt to lay their eggs on the cabbage than on the other congenerous plants.

However vague and unphilosophical these conjectures may appear, they serve, I think, to show that one’s first impulse utterly to reject any theory whatever, implying a gradual acquirement of these instincts, which for ages have excited man’s admiration, may at least be delayed. Once grant that dispositions, tastes, actions or habits can be slightly modified, either by slight congenital differences (we must suppose in the brain) or by the force of external circumstances, and that such slight modifications can be rendered inheritable, – a proposition which no one can reject, – and it will be difficult to put any limit to the complexity and wonder of the tastes and habits which may possibly be thus acquired.

Difficulties in the acquirement by Selection of complex corporeal structures

After the past discussion it will perhaps be convenient here to consider whether any particular corporeal organs, or the entire structure of any animals, are so wonderful as to justify the rejection primâ facie of our theory294. In the case of the eye, as with the more complicated instincts, no doubt one’s first impulse is to utterly reject every such theory. But if the eye from its most complicated form can be shown to graduate into an exceedingly simple state, – if selection can produce the smallest change, and if such a series exists, then it is clear (for in this work we have nothing to do with the first origin of organs in their simplest forms295) that it may possibly have been acquired by gradual selection of slight, but in each case, useful deviations296. Every naturalist, when he meets with any new and singular organ, always expects to find, and looks for, other and simpler modifications of it in other beings. In the case of the eye, we have a multitude of different forms, more or less simple, not graduating into each other, but separated by sudden gaps or intervals; but we must recollect how incomparably greater would the multitude of visual structures be if we had the eyes of every fossil which ever existed. We shall discuss the probable vast proportion of the extinct to the recent in the succeeding Part. Notwithstanding the large series of existing forms, it is most difficult even to conjecture by what intermediate stages very many simple organs could possibly have graduated into complex ones: but it should be here borne in mind, that a part having originally a wholly different function, may on the theory of gradual selection be slowly worked into quite another use; the gradations of forms, from which naturalists believe in the hypothetical metamorphosis of part of the ear into the swimming bladder in fishes297, and in insects of legs into jaws, show the manner in which this is possible. As under domestication, modifications of structure take place, without any continued selection, which man finds very useful, or valuable for curiosity (as the hooked calyx of the teazle, or the ruff round some pigeons’ necks), so in a state of nature some small modifications, apparently beautifully adapted to certain ends, may perhaps be produced from the accidents of the reproductive system, and be at once propagated without long-continued selection of small deviations towards that structure298. In conjecturing by what stages any complicated organ in a species may have arrived at its present state, although we may look to the analogous organs in other existing species, we should do this merely to aid and guide our imaginations; for to know the real stages we must look only through one line of species, to one ancient stock, from which the species in question has descended. In considering the eye of a quadruped, for instance, though we may look at the eye of a molluscous animal or of an insect, as a proof how simple an organ will serve some of the ends of vision; and at the eye of a fish as a nearer guide of the manner of simplification; we must remember that it is a mere chance (assuming for a moment the truth of our theory) if any existing organic being has preserved any one organ, in exactly the same condition, as it existed in the ancient species at remote geological periods.

 

The nature or condition of certain structures has been thought by some naturalists to be of no use to the possessor299, but to have been formed wholly for the good of other species; thus certain fruit and seeds have been thought to have been made nutritious for certain animals – numbers of insects, especially in their larval state, to exist for the same end – certain fish to be bright coloured to aid certain birds of prey in catching them, &c. Now could this be proved (which I am far from admitting) the theory of natural selection would be quite overthrown; for it is evident that selection depending on the advantage over others of one individual with some slight deviation would never produce a structure or quality profitable only to another species. No doubt one being takes advantage of qualities in another, and may even cause its extermination; but this is far from proving that this quality was produced for such an end. It may be advantageous to a plant to have its seeds attractive to animals, if one out of a hundred or a thousand escapes being digested, and thus aids dissemination: the bright colours of a fish may be of some advantage to it, or more probably may result from exposure to certain conditions in favourable haunts for food, notwithstanding it becomes subject to be caught more easily by certain birds.

If instead of looking, as above, at certain individual organs, in order to speculate on the stages by which their parts have been matured and selected, we consider an individual animal, we meet with the same or greater difficulty, but which, I believe, as in the case of single organs, rests entirely on our ignorance. It may be asked by what intermediate forms could, for instance, a bat possibly have passed; but the same question might have been asked with respect to the seal, if we had not been familiar with the otter and other semi-aquatic carnivorous quadrupeds. But in the case of the bat, who can say what might have been the habits of some parent form with less developed wings, when we now have insectivorous opossums and herbivorous squirrels fitted for merely gliding through the air300. One species of bat is at present partly aquatic in its habits301. Woodpeckers and tree-frogs are especially adapted, as their names express, for climbing trees; yet we have species of both inhabiting the open plains of La Plata, where a tree does not exist302. I might argue from this circumstance that a structure eminently fitted for climbing trees might descend from forms inhabiting a country where a tree did not exist. Notwithstanding these and a multitude of other well-known facts, it has been maintained by several authors that one species, for instance of the carnivorous order, could not pass into another, for instance into an otter, because in its transitional state its habits would not be adapted to any proper conditions of life; but the jaguar303 is a thoroughly terrestrial quadruped in its structure, yet it takes freely to the water and catches many fish; will it be said that it is impossible that the conditions of its country might become such that the jaguar should be driven to feed more on fish than they now do; and in that case is it impossible, is it not probable, that any the slightest deviation in its instincts, its form of body, in the width of its feet, and in the extension of the skin (which already unites the base of its toes) would give such individuals a better chance of surviving and propagating young with similar, barely perceptible (though thoroughly exercised), deviations304? Who will say what could thus be effected in the course of ten thousand generations? Who can answer the same question with respect to instincts? If no one can, the possibility (for we are not in this chapter considering the probability) of simple organs or organic beings being modified by natural selection and the effects of external agencies into complicated ones ought not to be absolutely rejected.

284A short discussion of a similar kind occurs in the Origin, Ed. i. p. 211, vi. p. 324.
285This sentence agrees with the MS., but is clearly in need of correction.
286This corresponds to Origin, Ed. i. p. 212, vi. p. 325.
287This discussion is interesting in differing from the corresponding section of the Origin, Ed. i. p. 216, vi. p. 330, to the end of the chapter. In the present Essay the subjects dealt with are nest-making instincts, including the egg-hatching habit of the Australian bush-turkey. The power of “shamming death.” “Faculty” in relation to instinct. The instinct of lapse of time, and of direction. Bees’ cells very briefly given. Birds feeding their young on food differing from their own natural food. In the Origin, Ed. i., the cases discussed are the instinct of laying eggs in other birds’ nests; the slave-making instinct in ants; the construction of the bee’s comb, very fully discussed.
288The distinction between faculty and instinct corresponds in some degree to that between perception of a stimulus and a specific reaction. I imagine that the author would have said that the sensitiveness to light possessed by a plant is faculty, while instinct decides whether the plant curves to or from the source of illumination.
289«Note in the original in an unknown handwriting.» At the time when corn was pitched in the market instead of sold by sample, the geese in the town fields of Newcastle «Staffordshire?» used to know market day and come in to pick up the corn spilt.
290«Note in original.» Macculloch and others.
291I can find no reference to the transandantes sheep in Darwin’s published work. He was possibly led to doubt the accuracy of the statement on which he relied. For the case of the sheep returning to their birth-place see .
292Origin, Ed. i. p. 224, vi. p. 342.
293This is an expansion of an obscure passage in the Essay of 1842, p. .
294The difficulties discussed in the Origin, Ed. i. p. 171, vi. p. 207, are the rarity of transitional varieties, the origin of the tail of the giraffe; the otter-like polecat (Mustela vison); the flying habit of the bat; the penguin and the logger-headed duck; flying fish; the whale-like habit of the bear; the woodpecker; diving petrels; the eye; the swimming bladder; Cirripedes; neuter insects; electric organs. Of these, the polecat, the bat, the woodpecker, the eye, the swimming bladder are discussed in the present Essay, and in addition some botanical problems.
295In the Origin, Ed. vi. p. 275, the author replies to Mivart’s criticisms (Genesis of Species, 1871), referring especially to that writer’s objection “that natural selection is incompetent to account for the incipient stages of useful structures.”
296«The following sentence seems to have been intended for insertion here» “and that each eye throughout the animal kingdom is not only most useful, but perfect for its possessor.”
297Origin, Ed. i. p. 190, vi. p. 230.
298This is one of the most definite statements in the present Essay of the possible importance of sports or what would now be called mutations. As is well known the author afterwards doubted whether species could arise in this way. See Origin, Ed. v. p. 103, vi. p. 110, also Life and Letters, vol. iii. p. 107.
299See Origin, Ed. i. p. 210, vi. p. 322, where the question is discussed for the case of instincts with a proviso that the same argument applies to structure. It is briefly stated in its general bearing in Origin, Ed. i. p. 87, vi. p. 106.
300«Note in original.» No one will dispute that the gliding is most useful, probably necessary for the species in question.
301«Note in original.» Is this the Galeopithecus? I forget. «Galeopithecus “or the flying Lemur” is mentioned in the corresponding discussion in the Origin, Ed. i. p. 181, vi. p. 217, as formerly placed among the bats. I do not know why it is described as partly aquatic in its habits.»
302In the Origin, Ed. vi. p. 221, the author modified the statement that it never climbs trees; he also inserted a sentence quoting Mr Hudson to the effect that in other districts this woodpecker climbs trees and bores holes. See Mr Darwin’s paper, Zoolog. Soc. Proc., 1870, and Life and Letters, iii. p. 153.
303Note by the late Alfred Newton. Richardson in Fauna Boreali-Americana, i. p. 49.
304«Note in original.» See Richardson a far better case of a polecat animal «Mustela vison», which half-year is aquatic. «Mentioned in Origin, Ed. i. p. 179, vi. p. 216.»
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