Chapter XIX: THE GERM-PLASM THEORY (continued) 392 (3)

Among pigeons, as well as among other domesticated animals, some changes have been brought about, which are not only of no use to their possessors, but would be actually disadvantageous if they were living under natural conditions. Some of the very short-billed breeds of pigeons have the bill so short and soft that the young can no longer use it to scratch and break the egg-shell, and would perish miserably if human aid were not at hand. The Yorkshire pig has become such a colossus of fat on weak, short legs, that if it were dependent on its own resources, it could not secure its food, much less escape from a beast of prey; and among horses the heavy cart-horse and the racer are alike unfit to cope with the dangers of a wild life, or the vicissitudes of weather.

Breeding has done much to bring about variations useful to man. Thus we have breeds of cattle which excel in flesh, or in milk, or as draught animals, and sheep which excel in flesh or in wool, and to what a height the perfecting of a useful quality can be brought is shown, in regard to fineness of wool, by that finest breed of sheep, the merino, which instead of the 5,500 hairs borne by the old German sheep on a square inch, possesses 48,000.

Not infrequently it is a particular stage of a species that has been bred by man, and the other stages have remained more or less unaltered. Thus it is with one of the few domesticated insects, the silk-moth. Only the cocoon is of use to man, and according to the cocoon different breeds are distinguished, differing in fineness, colour, &c.; but no breeds can be distinguished in reference to the larvæ, or the perfect insects. Among gooseberries there are about a hundred varieties distinguished according to the form, colour, size, thickness of skin, hairiness, &c., of the fruits, but the little, inconspicuous, green blossoms, of which the breeders take no account, are alike in them all. In the pansies (_Viola tricolor_), on the other hand, it is only by the flowers that the varieties are distinguished, while the seeds have remained alike in all.

It may be asked how it could have occurred to any one, when pigeons, for instance, first began to be domesticated, to wish to produce fantails or pouters, since he could have no mental picture of them in advance. Darwin replies to this objection, that it was not always conscious and methodical artificial selection, such as is now practised, that brought about the origin of breeds, but that they have very often resulted, and at first perhaps always, from unconscious selection. When savages tamed a dog, they used the 'best' of their dogs for breeding, that is, they chose those which had in the highest degree the qualities they valued, watchfulness, for instance, or if the dog were intended for the chase, keen scent and swiftness. In this way the body of the animal would be changed in a definite direction, especially if rivalry helped, and if it was the ambition of each to possess a dog as good as, or better than those of his tribal companions. That perfectly definite changes in bodily form can thus be brought about unconsciously is well illustrated by the case of a racehorse. This has arisen within the last two hundred years simply because the fleetest of the products of crossing between the Arab and the English horse were always chosen for breeding. It could not have been predicted that horses with thin neck, small head, long rump, and slender legs would necessarily be the swiftest runners; but this is the form which has resulted from the selection,--a very ugly, but very swift horse. This unconscious selection must undoubtedly have played a large part in the early stages of the evolution of the breeds of our domestic animals.

But even in the fully conscious and methodical selective breeding of particular characters, the breeder rarely alters only the one his attention is fixed on; generally quite a number of other characters alter apart from his intention as an inevitable accompaniment of the desired variation on which attention was riveted. There are breeds of rabbits whose ears hang limply down instead of standing erect, and in these so-called lop-eared rabbits the ear-muscles are partly degenerated, and as a consequence of this lack of muscular strain the skull has assumed another form. Thus the variation of one part may influence the development of a second and a third organ, and may even not stop there, for very often the influence has penetrated much deeper and affected quite remote parts of the body.

If any one were to succeed in adding a heavy pair of horns to a breed of hornless sheep, there would run parallel with the course of this variation, which was directly aimed at, a long series of secondary changes which would affect at least the whole of the anterior half of the body; the skull would become thicker and stronger to support the weight of the heavy horns; the neck-tendon (_ligamentum nuchæ_) would have to become thicker to hold up the heavy head, and so also with the muscles of the neck; the spinous processes of the cervical and dorsal vertebrae would become longer and stronger, and the forelegs, too, would need to adapt themselves to the heavier burden. Every organism thus resembles, as it were, a mosaic, out of which no individual group of pieces can be taken and replaced by another without in some measure disturbing the correlation and harmony of the whole: in order to restore this, the pieces all round about the changed part must be moved or replaced by others.

According to Darwin, it is to this correlation of parts that we must refer the variation of other parts besides the one intentionally altered in the course of breeding. It must be admitted that the mutual dependence of the parts plays a very important rôle in the economy and development of the animal body, as we shall see later, and these connexions still remain very mysterious to us. Especially is this the case with the connexion between the reproductive organs and the so-called secondary sexual characters. Removal of the reproductive organs or gonads induces, in Man, for instance, if it be effected in youth, the persistence of the childish voice and the non-development of the beard; in the stag the antlers do not appear, and in the cock the comb does not develop perfectly, &c., but we are not yet able to understand clearly why this should be so.

LECTURE III

THE DARWINIAN THEORY (_continued_)

Natural selection--Variation--Struggle for existence--Geometric ratio of rate of increase--Normal number and ratio of elimination in a species--Accidental causes of extinction--Dependence of the strength of a species on enemies--Struggle for existence between individuals of the same species--Natural selection affects all organs and stages--Summary.

IN artificial selection, through which, with or without conscious intention, our domesticated animals and cultivated plants have arisen, there must obviously be three kinds of co-operative factors: first, the _variability_ of the species; second, the capacity of the organism for _transmitting_ its particular characters to its progeny; and third, the _breeder_ who selects particular qualities for breeding. No one of the factors can be dispensed with; the breeder could effect nothing, were there not presented to him the variations of parts in the particular direction in which he wishes them to vary; an indefinite variation, that is, a variation not guided by selection, would never lead to the formation of new breeds; the species would probably become in time a motley mixture of all sorts of variations, but a breed with definite characters, transmissible in their purity to its descendants, could never be formed. Finally, every process of selective breeding would be futile, if the variations which appeared could not be transmitted.

Darwin assumes that processes of transformation quite similar to those which take place under the guidance of Man occur also in nature, and that it is mainly these which have brought about and guided the transformations of species which have taken place in the course of the earth's history. This process he calls _natural selection_.

It will readily be admitted that two out of the three factors necessary to a process of selective breeding are present also in the natural conditions of the life of species. Variability in some degree or other is absent from no species of animal or plant, though it may be greater in one than in another, and it cannot be doubted that the inborn differences which distinguish one individual from another are capable of transmission. It is only to untrained observers that all the individuals of a species appear alike; for instance, all garden whites, or all the individuals of the small tortoiseshell butterfly (_Vanessa urticæ_), or all the chaffinches. If the individuals are carefully compared it will be recognized that, even in these relatively constant species, no individual exactly resembles another; that even among butterflies twenty black scales may go to form a particular spot on the wings in one individual and thirty or twenty-five in others; that the length of the body, the legs, the antennæ, the proboscis exhibit minute differences; and it is probable that the same combination of quite similar parts never occurs twice. In many animals this cannot, of course, be proved, because our power of diagnosis is not fine enough to be able to estimate the differences directly, and because a comparison of measurements of all the parts in detail is not practicable. So we may here confine ourselves to the differences in the human race, which we can recognize with ease and certainty. Even as regards the face alone, all men differ from one another, and, numerous and complete as likenesses may be, it is impossible to find two human beings in which even the characters of the face are exactly similar. Even so-called 'identical twins' can always be distinguished if they are directly compared either in person or in a photograph, and if the rest of the body be also taken into consideration we find numerous small, sometimes even measurable differences.

The same is true of animals, and it is only our lack of practice that is at fault if we frequently fail to detect their individual differences. The Bohemian shepherds are said to know personally, and be able to distinguish from all the rest, every sheep in their herds of many thousands. Thus the factors of variability and transmissibility must be granted, and it remains only to ask: Who plays the part of selecting breeder in wild nature? The answer to this question forms the kernel to the whole Darwinian theory, which ascribes this rôle to the conditions of life, to definite relations of individuals to the external influences which they meet with during the course of their lives, and which together make up their 'struggle for existence.'

To make this idea clear I must to some extent diverge.

It is a generally observed fact that, in every species of animals or of plants, more germs and more individuals are produced than grow to maturity, or become capable of reproduction. Numerous young individuals perish at an early stage, often because of unfavourable circumstances--cold, drought, damp, or through hunger, or at the hands of their enemies. When we ask which of the progeny perish early, and which survive to carry on the species, we are at first sight inclined to suppose that this is entirely a matter of chance; but this is just what Darwin disputed. It is not chance alone, it is, above all, the differences between individuals, which enable them to withstand adverse circumstances better or worse, and thus decide, according to his view, which shall perish and which shall survive. If this be so, then we have a veritable process of selection, and one which secures that the 'best,' that is, the most capable of resistance, survive to breed, being thus, so to speak, 'selected.'

It may be asked, however, why so many individuals must perish in youth, and whether it could not have been arranged that all, or at least most, should survive till they had reproduced. But this is an impossibility, unrealizable for this among other reasons, that organisms multiply in geometrical progression, and that their progeny would very soon exceed the limits of computability. This does not occur, for there is a limit set which they can in no case overstep,--which, indeed, as we shall see, they never reach--I mean the limits of space and food-supply. Every species, by the natural requirements of its life, is restricted to a particular habitat, to land or to water, but most are still more strictly limited to a definite area of the earth's surface, which alone affords the climate suited to them, or where alone the still more specialized conditions of their existence can be realized. Thus, for instance, the occurrence of a particular species of plant determines that of the animal which is dependent on it for its food-supply. If they could multiply unchecked, that is, without the loss of many of their progeny, every species would fill up its area of occurrence and exhaust the whole of its food-supply, and thus bring about its own extermination. This seems to be prevented in some way, for as a matter of fact it does not happen.

It may, perhaps, be imagined that this might be prevented by a regulation of the productivity of the species, and that those which have not a large area of distribution, or can only count on a relatively limited food-supply, have also a low rate of multiplication, but this is not the case; even the lowest rate of multiplication would very soon suffice to make any species fill up its whole available space and completely exhaust its food-supply. Darwin takes as an example the elephant, which only begins to breed at thirty years of age, and continues to do so till about ninety, but so slowly that in these sixty years only three pairs of young are produced. Nevertheless, in 500 years an elephant pair would be represented by fifteen millions of descendants, if all the young survived till they were capable of reproduction. A species of bird with a duration of life of five years, during which it breeds four times, producing and rearing four young each time, would in the course of fifteen years have 2,000 millions of descendants.

Thus, although the fertility of each species is, as a matter of fact, precisely regulated, a low rate of multiplication is not in itself sufficient to prevent the excessive increase of any species, nor is the quantity of the relevant food-supply. Whether this be very large or very small, we see that in reality it is never entirely used up, that, as a matter of fact, a much greater quantity is always left over than has been consumed. If increase depended only on food-supply, there would, for instance, be food enough in their tropical home for many thousand times more elephants than actually occur; and among ourselves the cockchafers might appear in much greater numbers than they do even in the worst cockchafer year, for all the leaves of all the trees are never eaten up; a great many leaves and a great many trees are left untouched even in the years when the voracious insects are the most numerous. Nor do the rose-aphides, notwithstanding their enormously rapid multiplication, ever destroy all the young shoots of a rose-bush, or all the rose-bushes of a garden, or of the whole area in which roses grow.

At the same time it must be noted, that the number of individuals in a species undoubtedly does bear some relation to the amount of the food-supply available; for instance, it is very low among the large carnivores, the lion, the eagle, and the like. In our Alps the eagles have become rarer with the decrease of game, and where one eagle pair make their eyrie they rule alone over a hunting territory of more than sixty miles, a preserve on which no others of the same species are allowed to intrude. If there were several pairs of eagles in such a preserve, they would soon have so decimated the food-supply that they would starve. On the other hand, numerous herbivores, e.g. chamois and marmots, live within the bounds of the pair of eagles' hunting grounds, since the food they require is present in enormously greater quantity.

While it is true that the number of individuals of a given species which live in a particular area is not exactly the same year in year out, being subject to small, and sometimes, as in the case of the aphides and cockchafers, to very great fluctuations, nevertheless we may assume that the _average number_ remains the same, that in the course of a century, or, let us say, of a thousand years, the number of mature individuals inhabiting the particular area remains the same. This, of course, only holds true on the supposition that there has been no great change in the external conditions of life during this period. But before Man began to interfere with nature, these external conditions would remain uniform for much longer periods than we have assumed. Let us call the average number of individuals occurring on such a uniform area, _the normal number_ of the species; this number will be determined in the first instance by the number of offspring that are annually brought forth, and secondly by the number that annually perish before reaching maturity. As the fertility of a species is a definite quantity, so also will its elimination be definite, or, as we may say, when the normal number under uniform conditions of life remains constant, the ratio of elimination will also remain constant. Each species is therefore subject to a perfectly definite ratio of elimination which remains on the average constant, and this is the reason why a species does not multiply beyond its normal number notwithstanding the great excess of the food-supply, and notwithstanding the fertility which, in all species, is sufficient to lead to boundless multiplication.

It is not difficult to calculate the ratio of elimination for a particular species, if one knows its rate of multiplication; for if the normal number remains constant, it follows that only two of all the offspring which a pair brings forth in the course of its life can attain to reproductive maturity, and that all the rest must perish.

Suppose, for instance, a pair of storks produced four young ones annually for twenty years, of these eighty young ones which are born within this period, on an average seventy-eight must perish, and only two can become mature animals. If more than two attained maturity the total number of storks would increase, and this is against the presupposition of constancy in the normal number. It is important, in reference to the fact on which we are now focusing our attention, that we should consider some other illustrations from the same point of view. The female trout yearly produces about 600 eggs; let us assume that it remains capable of reproduction for only ten years, then the elimination-number of the species will be 6,000 less two, that is, 5,998, for of the 6,000 eggs only two can become mature animals. But in the majority of fishes the ratio of extermination is enormously greater than this. Thus a female herring brings forth 40,000 eggs annually, the duration of life is estimated at ten years, and this means an elimination number of 400,000 less two, that is, 399,998. The carp produces 200,000 eggs a year, and the sturgeon two millions, and both species live long, and remain capable of reproduction for at least fifty years. But of all the 100 million eggs which are produced by the sturgeon, only two reach their full development and reproduce; all others perish prematurely.

But even with these examples we have not reached the highest elimination number, for many of the lower animals--not to speak of many plants--produce an even greater number of offspring. Leuwenhoek calculated the fertility of a thread-worm at sixty million eggs, and a tape-worm produces hardly less than 100 millions.

There exists, therefore, a constant relation between fertility and the ratio of elimination; the higher the latter is, the greater must the former be, if the species is to survive at all. The example of the tape-worm makes this very obvious, for here we can readily understand why the fertility must be so enormous, as we are aware of the long chain of chances on which the successful development of this animal depends. The common tape-worm of Man, _Tænia solium_, does not lay its eggs, they remain enclosed within one of the liberated joints or 'proglottides.' Only if this liberated joint or one of the embryos within it happens to be fortuitously eaten by a pig or other mammal can there be successful development, and even then under difficulties and possible failures, and not right away into adult animals, but first into microscopically minute larvæ which may bore their way into the walls of the intestine, or, if they are fortunate enough, may get into the blood-stream and be carried by it to a remote part of the body. There they develop into 'measles,' the so-called bladder-worms, within which the head of the tape-worm arises. But in order that this may become a complete and reproductive adult worm the pig must die, and the next step necessary is that a piece of the flesh of the infected first host must happen to be swallowed raw by a man or other mammal! Only then does the fortunate bladder-worm--swallowed with the flesh--attain the goal of its life, that is, a suitable place to mature in, the food-canal of a human being. It is obvious that countless eggs must be lost for one that succeeds in getting through the whole course of a development depending so greatly on chance. Hence the necessity for such enormous productivity of eggs.

In many cases the causes of elimination, which keep a species within due bounds, are very difficult to determine. Enemies, that is to say, other species which use the species in question as food, play an important rôle; often, however, the cause lies in the unfavourableness of external conditions, in chance, which is favourable only to one of a thousand. The oak would only require to produce one seed in the 500 years of its life, if it were certain that that one would grow into an oak-tree; but most of the little acorns are eaten up by pigs, squirrels, insects, &c., before they have had time to sprout, thousands fall on ground already thickly covered with growth where they cannot take root, and even if they do succeed in finding an unoccupied space in which to germinate, the young plants are still surrounded by a thousand dangers--the possibility of being devoured by many animals large and small, of being suffocated by the surrounding vegetation, and so on. We can thus understand, to some extent, though only approximately, why it is that the oak must year by year produce thousands of seeds in order that the species may maintain its normal number, and not be exterminated; for it is obvious that a constant, even though slow diminution of the normal number, a regular deficit, so to speak, can end in nothing else than the gradual extinction of the species.

But even this prodigality of seeds is not the greatest reach of fertility that we meet with in nature; it is, perhaps, amongst the simpler flowerless plants that we find the climax. It has been calculated that a single frond of the beautiful fern so common in our woods, _Aspidium filix mas_, produces about fourteen million spores. They serve to distribute the species, and are carried as motes by the wind, but comparatively few of the millions ever get the length of germinating at all, much less of attaining to full development into adult plants. Thus we see that the apparent prodigality of nature is a real necessity, an indispensable condition of the maintenance of the species; the fertility of each species is related to the actualities of elimination to which it is exposed. This is clearly seen when a species is placed under new and more favourable conditions of life, in which it has an abundant food-supply and few enemies. This was the case, for instance, with the horses introduced from Europe into South America, where they reverted to a feral state, and are now represented by herds of many thousands roaming the great grassy plains. If the small singing-birds of a region diminish in number, there is a great increase of caterpillars and other injurious insects which form part of their food-supply. The colossal destruction which the much-dreaded nun-moth from time to time brings about in our woods probably depends in part on the diminution of one or another of the many animals inimical to insects; but the occurrence of several years of weather-conditions favourable to the larvæ must also be taken into account. How enormously, indeed almost inconceivably, the number of larvæ may increase under favourable conditions is shown by such devastations as that in Prussia in 1856, when many square miles of forest were absolutely eaten up. The caterpillars were so numerous that even from some distance the falling excrement could be heard rustling like rain, and ten hundredweights of the eggs were collected, with an average of 20,000 eggs to the half-ounce!

But it would be a great mistake to conclude, from this enormous and sudden increase in the number of individuals, that the normal number of individuals is determined by the number of enemies alone. The average number of individuals in a species depends on many other conditions, especially on the extent of the available area, and on the amount of the food-supply in relation to the size of body in the species. I cannot dwell on this now, but I wish to point out that, for the continuance of a species, it is indifferent whether it is 'frequent' or 'rare,' if we presuppose that its normal number remains on an average constant for centuries, that is, that its fertility suffices to make good the continual losses through enemies and other causes of elimination. One would be inclined to conclude from such cases of sudden and enormous increase in the number of individuals as these caterpillar-blights, that enemies and other causes of destruction played the major part in the regulation of the normal number of the species. But this is only apparently the case. Enemies necessitate a certain fertility in the species on which they prey, so that the elimination in each generation may be made good; but the number of pairs capable of reproduction is not thereby decisively determined. We must not forget that the number of enemies is also, on the other hand, dependent on the number of victims, and that the normal number of enemies must rise and fall with that of the species preyed upon.

For this reason, such an enormous increase as that of the caterpillars cannot last long; it carries its corrective in itself. The appearance of the caterpillars in such enormous numbers in itself increases the host of their enemies; singing-birds, ichneumon-flies, beetle-grubs, and predaceous beetles find abundant and available food, and therefore reproduce and multiply so rapidly, that, with the help of the caterpillar's plant-enemies, especially the insect-destroying fungi, they soon reduce the caterpillars to their normal number, or even below it. But then the reverse process begins; the enemies of the caterpillars diminish because their food has become scarce, and their normal number is lowered, while that of the caterpillars gradually rises again.

When the number of foxes in a hunting district increases, the number of the hares that they prey upon diminishes, and, on the other hand, the decimating of the foxes by Man brings about an increase in the number of hares in the district. Under natural conditions, that is, without the intervention of Man, there would be a constant balancing of the numbers of hares and foxes, for every noteworthy increase of the hares would be followed by a similar increase of foxes, and this, in its turn, would diminish the number of hares, so that they would no longer suffice for the support of so many foxes, and these would decrease in number again, until the number of hares had again increased because of the lessened persecution and elimination. In nature the case is not quite so simple, because the fox does not live on hares alone, and the hare is not preyed upon only by the fox; but the illustration may serve to elucidate the point that a moving equilibrium is maintained between the species of a district, between persecutors and persecuted, in such a way that the number of individuals in the two species is always varying a little up and down, and that each influences the other so that a regulative process results. Throughout periods of considerable length the average remains the same; that is to say, a _normal number_ is established. This normal strength of population is the mean above and below which the number of individuals is constantly varying. It is, of course, seldom that the mutual influences and regulations are so simple as in the example given; usually several or even many species interact upon each other, and not beasts of prey and their victims alone, but the most diverse species of animals and plants, which do not stand in any obvious relation to one another at all. Moreover, the physical, and especially the climatic conditions, also cause the normal number of the species to rise and fall.

The inter-relations between species living together on the same area are so intricate that I should like to give two other illustrations. Let us first take Darwin's famous instance of the fertility of clover, which depends on the number of cats. It is of course only an imaginary one, but the facts it is based upon are quite correct. The number of cats living in a village to a certain extent determines the number of field-mice in the neighbourhood. These again destroy the nests of the humble-bees, which live in holes in the ground, and thus the number of humble-bees depends on that of the field-mice and cats. But the clover must be pollinated by insects if it is to produce fertile seed, and only the humble-bee has a proboscis long enough to effect the pollination. Therefore the quantity of clover-seed annually produced depends on the number of humble-bees, and ultimately upon the number of cats. And, as a matter of fact, humble-bees were introduced into New Zealand from England, because without them the clover would produce no fertile seeds.

On the grassy plains of Paraguay there are no wild cattle and horses, because of the presence of a fly which has a predilection for laying its eggs in the navel of the newly-born calves and foals, with the result that the calves or foals are killed by the emerging maggots. We may reasonably assume that the numerical strength of this fly-species depends on the distribution of insect-eating birds, whose numbers in turn are determined by certain beasts of prey. These again vary in number in relation to the extent of the forest-land, and this is determined by the number of ruminants which browse on the young growth of the woods (Darwin).

That forests can actually be totally destroyed by ruminants is proved by the case of the island of St. Helena among others. On its discovery the island was covered with thick wood, but in the course of 200 years it was transformed into a bare rock by goats and pigs, which devoured the young growth so completely that trees which were felled or which died were not replaced.

This point is vividly illustrated by Darwin's observation of a wide heath on which stood only a few groups of old pine-trees. The mere fencing in of a portion of the heath sufficed to call forth a thick growth of young seedling pines within the enclosure, and an examination of the open part of the heath revealed that the grazing cattle had eaten up all the young pine-trees which sprang from seed, and that again and again. In one small space thirty-two little trees stood concealed in the grass, and several of these showed as many as twenty-six yearly rings.

How definitely the number of individuals in different species living on the same area mutually limit and thereby regulate each other, Darwin sought to illustrate also by the case of the primitive forest, where the numerous species of plants occur, not mixed together irregularly, but in a definite proportion. We can find examples of the same kind wherever the plant-growth of a district has been left to itself. If we walk along the banks of our little river, the Dreisam, we see a wild confusion of the most diverse trees, shrubs and herbaceous plants. But, even though it cannot be demonstrated, we may be certain that these are represented in definite numerical proportions, dependent on the natural qualities and requirements of each species, on the number of their seeds and the facilities for their distribution, on the favourable or unfavourable season at which they ripen, and on their varying capacity for taking root in the worst ground, and springing quickly up, &c. They limit each other mutually, so that the whole flora of the river-bank will be made up of one per cent. of this species, one per cent. of that, and, it may be, five per cent. of a third, and the same combination will repeat itself in the same proportions on the banks of other rivers of our country in as far as the external conditions are the same. The same must be true of the fauna of such a plant-thicket; the animal species also limit one another mutually, and thereby regulate the number of individuals, which becomes relatively stable over any area on which the conditions remain the same. That is to say, a 'normal number' is attained and persists.

Thus we see that the capacity for boundless multiplication inherent in every species is limited by the co-existence of other species; there is, metaphorically speaking, a continuous struggle going on between species, plant and animal alike; each seeks as far as possible to multiply, and each is hemmed in by the others and as far as possible prevented from doing so. The 'struggle' is by no means only the _direct_ limitation of the number of individuals, which consists in the use of one species by another as food, as in beasts of prey and their victims, or locusts and plants; it is much more the indirect limitation--figuratively speaking, the struggle for space, for light, for moisture among plants, for food among animals. But all this, important as it is, does not yet exhaust the content of that 'struggle for existence' to which Darwin and Wallace ascribe the rôle of the breeder in the process of natural selection. The struggle, that is, the mutual limiting of species, may indeed restrict a species in its distribution, and may reduce its normal number possibly to nil. In other words, it may bring about extinction, but it cannot make a species other than it is. This can only be done by a struggle within the limits of the species itself, and this struggle is due to the fact that of the numerous offspring, on an average those survive--that is, attain to reproduction--which are the most fit, whose constitution makes it most possible for them to overcome the difficulties and dangers of life, and so to reach maturity. We see, in fact, that a large percentage of each generation in all species always perishes before attaining maturity. If, then, the decision as to which is to perish and which is to reach maturity is _not a matter of chance alone_, but is in part due to the constitution of the growing individual; if the 'fittest' do _on the average_ survive, and the 'least fit' are on the average eliminated, we have here a process of selection entirely comparable to that of artificial selection, and one whose result must be the 'improvement' of the species, whether that depends on one set of characters or on another. The victorious qualities, which earlier were peculiar to certain individuals, must gradually become the common property of the species, if in each generation the individuals which attained to reproduction all possessed them, and thus could transmit them to their progeny. But those of the descendants which did not inherit them would again be at a disadvantage in the struggle for existence, or rather for reaching maturity, if in each generation a higher percentage of individuals which possess these characters reach maturity than of those which do not possess them. This percentage must increase in each generation, because, in each, natural selection again chooses out the fittest, and it must finally rise to 100 per cent., that is to say, none but individuals of this fittest type will be left surviving.

This does not yet exhaust the process, however, for we can infer from the results of artificial breed-forming that the selected characters may intensify from generation to generation, and that they will continue to do so as long as it gives them any advantage in the struggle for existence, for so long will it lead to the more frequent survival of its possessors. The increase will only stop when it has reached the highest degree of usefulness, and in this way new characters may be formed, just as, in artificial selection, the short upward-turning feathers of the Jacobin pigeon have been intensified into the peruke, a feather canopy covering the head.

A few examples of natural selection will make the process clearer. Our hare is well secured from discovery by his fur of mixed brown, yellow, white, and black, when he cowers in his form among the dry leaves of the underwood. It is easy to pass close to him without seeing him. But if the ground and the bushes are covered with snow, he contrasts conspicuously with them. Suppose, now, that our climate became colder, and that the winter brought lasting snow, the hares which had the largest mixture of white in their fur would have an advantage in their 'struggle for existence' over their darker fellows; they would be less easily discovered by their enemies--the fox, the badger, the horned owl, and the wild cat. Of the numerous hares which would annually become the prey of these enemies, there would be, on an average, more dark than light individuals. The percentage of light-coloured hares would, therefore, increase from generation to generation, and the longer the winter the keener would be the selection between dark and light hares, until finally none but light ones would remain. At the same time, the colour of the hares would become increasingly light, first, because it would happen more and more frequently that two light hares would pair, and secondly, because, after a time, the struggle for existence would no longer be between light and dark hares, but between light hares and still lighter ones. Thus ultimately a race of white hares would arise, as has actually happened in the Arctic regions and on the Alps.

Or let us think of a herbaceous plant, in appearance something like a belladonna, rich in leaves and very juicy, but not poisonous. It would doubtless be a favourite food with the animals of the forest, and it would not, therefore, attain to more than a sparse occurrence, since few of the individuals would be able to form seeds. But now let us assume that a stuff of very unpleasant taste develops in the stem and leaves of some of the individuals, as may easily happen through very slight changes in the chemical metabolism of the plant, what, then, could result but that such individuals would be less readily eaten than the others? A process of selection must, therefore, ensue, and the unpleasant-tasting specimens of the plant would be much more frequently spared, and consequently would bear seed much oftener than the palatable ones. Thus the number of unpalatable plants would increase from year to year. If the stuff in question were not only unpalatable but poisonous, or gradually became so, a plant would in time be evolved which would be absolutely safe from being devoured by animals, just as the deadly nightshade (_Atropa belladonna_) actually is.

Or let us suppose that a stretch of water is inhabited by a species of carp, which have hitherto had no large enemy, and so have become lazy and slow, and that there migrates from the sea into this stretch of water a large species of pike. At first numerous carp will fall victims to the pike, and the pike will rapidly increase in number. But if all the carp were not equally lazy and dull-witted, if some of them were quicker and more intelligent, these would, on an average, become more rarely the victims of the pike, and numerous individuals with these better qualities would survive in each generation, till ultimately there were no others, and the useful characters would gradually become intensified, and so a more active and wary race of carp would arise.

Let us suppose, however, that the increased activity and wariness would not alone suffice to preserve the colony from extinction; it might require also an increased fertility to prevent the normal number from being permanently lowered; but even this could eventually be brought about by natural selection, if the nature of the species and the general conditions of its life permitted. For there are variations of fertility in every species, and if the chance of seeing some of its eggs become mature animals were greater for the more fertile female than for the less fertile, _ceteris paribus_, a process of selection must take place, which would result in an increase of fertility as far as that was possible.

Obviously, such processes of natural selection can affect all parts and characters--size and form of the body, as well as isolated parts, the external skin and its colour, every internal organ--and not bodily characters alone, but psychical ones as well, such as intelligence and instincts. According to this principle, it is only characters which are biologically indifferent that cannot be altered through natural selection.

Natural selection can also bring about changes at every age, for the elimination of individuals begins from the egg, and any kind of egg which is in some way better able to escape elimination will transmit its useful characters to its descendants, because the resulting young animals will thus more frequently reach full development than the young from other eggs. In the same way, at every succeeding stage of development, every character favourable to the preservation of the individual will be maintained and intensified.

We see from all this that natural selection is vastly more powerful than artificial selection by Man. In the latter, only one character at a time can be caused to change, while natural selection may influence a whole group of characters at the same time, as well as all the stages of development. Through the weeding out of the individuals which are annually exterminated, it is always on an average the 'fittest' which survive, that is to say, those which have the greatest number of bodily parts and rudiments of parts in the fittest possible condition of development at every stage. The longer this process of selection continues, the smaller will be the deviations of the individual from this standard, and the more minute will be the differences of fitness determining which is to be eliminated and which is to survive to reproduce its characteristics. In the immeasurable periods of time which are at the disposal of natural selection, and in the inestimable numbers of individuals on which it may operate, lie the essential causes of superiority of natural selection over the artificial selection of Man.

To sum up briefly: Natural selection depends essentially on the cumulative augmentation of the most minute useful variations in the direction of their utility; only the useful is developed and increased, and great effects are brought about slowly through the summing up of many very minute steps. Natural selection is a self-regulation of the species which secures its preservation; its result is the ceaseless adaptation of the species to its life-conditions. As soon as these vary, natural selection changes its mode of action, for what was previously the best is now no longer so; parts that before had to be large must now perhaps be small, or vice versa; muscle-groups which were weak must now become strong, and so on. The conditions of life are, so to speak, the mould into which natural selection is continually pouring the species anew.

But the philosophical significance of natural selection lies in the fact, that it shows us how to explain the origin of useful, well-adapted structures purely by mechanical forces and without having to fall back on a _directive_ force. We are thus for the first time in a position to understand, in some degree, the marvellous adaptation of the organism to an end, without having to call to our aid any supernaturally intrusive force on the part of the Creator. We understand now how, in a purely mechanical way, through the forces always at work in nature, all forms of life must conform to, and adapt themselves precisely to the conditions of their life, since only the best possible is preserved, and everything less good is continually being rejected.

Before I go on to expound in detail the phenomena which we refer to natural selection, I must briefly state that Darwin did not ascribe to natural selection by any means all the changes which have taken place in organisms in the course of time. On the one hand, he ascribed a not inconsiderable importance to the correlated variations we have already mentioned; still more, however, he relied on the direct influence of altered conditions of life, whether these consist in climatic and other changes in the environment, or in the assumption of new habits, and the increased or diminished use of individual parts and organs thereby induced. He recognized the principle so strongly emphasized by Lamarck, of use and disuse as a cause of heritable increase or decrease of the exercised or neglected part, though he did so with a certain reserve. I shall return later to these factors of modification, and shall then attempt to show that these too are to be referred to processes of selection, which are, however, of a different order from the phenomena which the Darwin-Wallace principle of natural selection serves to interpret. But, in the first instance, it appears to me to be necessary to show how far the Darwin-Wallace interpretation will suffice, and in the next lectures we shall occupy ourselves with this question exclusively.

LECTURE IV

THE COLORATION OF ANIMALS AND ITS RELATION TO THE PROCESSES OF SELECTION

Biological significance of colours--Protective colours of eggs--Animals of the snow-region--Animals of the desert--Transparent animals--Green animals--Nocturnal animals--Double colour-adaptation--Protective marking of caterpillars--Warning markings--Dimorphism of colouring in caterpillars--Shunting back of colouring in ontogeny--'Sympathetic' colouring in diurnal Lepidoptera--In nocturnal Lepidoptera--Theoretical considerations--The influence of illumination in the production of protective colouring, _Tropidoderus_--Harmony of protective colouring in minute details--_Notodonta_--Objections--Imitation of Strange objects, _Xylina_--Leaf-butterflies, _Kallima_--_Hebomoja_--Nocturnal Lepidoptera with leaf-markings--Orthoptera resembling leaves--Caterpillars of the Geometridæ.

WE have seen what Darwin meant by natural selection, and we understand that this process really implies a transformation of organisms by slow degrees, in the direction of adaptive fitness--a transformation which must ensue as necessarily as when a human selector, prompted by conscious intention, tries to improve an animal in a particular direction, by always selecting the 'fittest' animals for breeding. In nature, too, there is selection, because in every generation the majority succumb in the struggle for life, while on an average those which survive, attain to reproductive maturity, and transmit their characters to their descendants, are those which are best adapted to the conditions of their life--that is, which possess those variations of most advantage in overcoming the dangers of life. Since individuals are always variable in some degree, since their variations can be inherited by their progeny, and since the continually repeated elimination of the majority of those descendants is a fact, the inference from these premisses must be correct; there must be a 'natural selection' in the direction of a gradually increasing fitness and effectiveness of the forms of life.

We cannot, however, directly observe this process of natural selection; it goes on too slowly, and our powers of observation are neither comprehensive nor fine enough. How could we set about investigating the millions of individuals which constitute the numerical strength of a species on a given area, to find out whether they possess some variable character in a definite percentage, and whether this percentage increases in the course of decades or centuries? And there is, furthermore, the difficulty of estimating the biological importance of any variation that may occur. Even in cases where we know its significance quite well in a general way, we cannot estimate its relative value in reference to the variation of some other character, though that other may also be quite intelligible. Later on, we shall speak of protective colouring, and in so doing we shall discuss the caterpillars of one of the Sphingidæ, which occur in two protective colours, some being brown, others green. From the greater frequency of the brown form we may conclude that brown is here a better adaptation than green, but how could we infer this from the character itself, or from our merely approximate knowledge of the mode of life of the species, its habits, and the dangers which threaten it? A direct estimation of the relative protective value of the two colours is altogether out of the question. The survival of the fittest cannot be proved in nature, simply because we are not in a position to decide, _a priori_, what the fittest is. For this reason I was forced to try to make the process of natural selection clear by means of imagined examples, rather than observed ones.

But though we cannot directly follow the uninterrupted process of natural selection which is going on under natural conditions, there is another kind of proof for this hypothesis, besides that which consists in logically deducing a process from correct premisses; I should like to call this the practical proof. If a hypothesis can be made to explain a great number of otherwise unintelligible facts, it thereby gains a high degree of probability, and this is increased when there are no facts to be found which are in contradiction to it.

Both of these criteria are fulfilled by the selection-hypothesis, and indeed the phenomena which may be explained by it, and are intelligible in no other way, present themselves to us in such enormous numbers, that there can be no doubt whatever as to the correctness of the principle; all that can be still disputed is, how far it reaches.

Let us now turn our attention to this practical way of proving the theory by the facts which it serves to interpret, beginning with a consideration of the external appearance of organisms, their colour and form.

_The Colour and Form of Organisms._

Erasmus Darwin had in many cases already rightly recognized the biological significance of the colouring of an animal species, and we may be sure that many of the numerous good observers of earlier times had similar ideas. I can even state definitely that Rösel von Rosenhof, the famous miniature-painter and naturalist of Nürnberg in the middle of the eighteenth century, recognized clearly, and gave beautiful descriptions of what we now call colour-adaptation. It is true that he gave them only as isolated instances, and was far from recognizing the phenomenon of colour-adaptation in general, or even from inquiring into its causes. From the time of Linné, the endeavour to establish new species overshadowed all the finer observation of life-habits and inter-relations, and, later on, after Blumenbach, Kielmeyer, Cuvier, and others, the eager investigation of the internal structure of animals also tended to divert attention from these œcological relations. In systematic zoology, colour ranked only as a diagnostic character of subordinate value, because it is often not very stable, and indeed is sometimes very variable; it was therefore found preferable to keep to such relatively stable differences as are to be found in the form, size, and number of parts.

Charles Darwin was the first to redirect attention to the fact that the colouring of animals is anything but an unimportant matter; that, on the contrary, in many cases it is of use to the animal, e.g. in making it inconspicuous; a green insect is not readily seen on green leaves, nor a grey-brown one on the bark of a tree.