Part 6

Like the bill, the legs vary in regard to their length. The pouters (No. 1) stand on their long legs as on stilts, while the legs of the 'Nürnberger swallow' are strikingly small. Remarkable, too, and very different from the wild species, is the thick growth of feathers on the feet and toes of the pouters and trumpeters (Fig. 1, No. 1), as well as of some other breeds, which suggests the arrangement of feathers on a wing.

Furthermore, the number and size of wing and tail-feathers in the different breeds often deviate considerably from the normal. The fantail (No. 7) in its most perfect form possesses forty tail-feathers, instead of the twelve usual in the wild rock-pigeon, and they are carried upright like a fan, while the head and neck of the bird are bent sharply backwards. In the hen-like pigeons the tail-feathers are few and short, so that they show an upright tail like that of a hen. I have already referred to the extraordinary carunculated skin-growth on the bill of many breeds; such folds also often surround the eye, and, as in the Indian barb (No. 3), are developed into well-formed thick circular ridges, while in the English carrier (No. 6) they lie about the bill as a formless mass of flesh.

Even the skull has undergone many variations, as can be observed even in the living bird in many of the breeds with short forehead. Differences are to be found, too, in the number and breadth of the ribs, the length of the breast-bone, the number and size of the tail-vertebræ in different breeds. Of the internal organs, the crop in many breeds, but particularly in the pouters (No. 1), has attained an enormous size, and with this size is usually associated the habit of blowing it out with air, and assuming the characteristically upright position.

That variations have taken place, too, in the most delicate structure of the brain, is shown by certain new instincts, such as the trumpeting of the trumpeters, the cooing of others, and the silence of yet other breeds, as well as by the curious habit of the tumblers of ascending quickly and vertically to a considerable height, and then turning over once, or even several times, in the course of their descent. In contrast to this, other breeds like the fantails have altogether given up the habit of flying high, and usually remain close to the dove-cot.

Lastly, let me mention that the unusual development of individual feathers, or of groups of feathers, has become a race-character, upon which depend such remarkable structures as the feather-mantle turned over the head in the Jacobins (No. 9), the cap or plume on the head of various breeds, the white beard in the bearded tumbler, the collars which lie like a shirt-collar on the breast, or run down the sides of the neck (Nos. 8 and 10), and the circle of feathers which marks the root of the bill in the Bucharest trumpeter (No. 3).

After what has been said, it is hardly necessary to add that the size of the whole body differs in different races. But the differences are very considerable, for, according to Darwin, one of the largest runt-pigeons weighed exactly five times as much as one of the smallest tumblers with short forehead, and in the illustration (Fig. 1) the pouter looks a giant beside the little barb to its left.

Thus we see that nearly every part of the body of the pigeon has varied under domestication in the most diverse ways, and to a high degree; and the same is true of several other domesticated animals, poultry, horses, sheep, cattle, pigs, and so on, though the matter is not altogether so clear in their case, since descent from a single wild species cannot be proved, and is in many cases improbable. But in the case of pigeons this common descent is certain, and we have now to inquire in what manner all these variations from the parent form have been brought about.

The answering of this question is rendered easier by the fact that new breeds arise even now, and that, to some extent at least, they can be caused to arise, consciously and intentionally. In England, as well as in Germany and France, there are associations for the breeding of birds, and in England especially pigeon and poultry clubs are numerous and highly developed. These by no means confine themselves to simply preserving the purity of existing breeds, they are continually striving to improve them, by increasing and accentuating their characters, or even by introducing quite new qualities, and in many cases they succeed even in this last. Prizes are offered for particular new variations, and thus a spirit of rivalry is fostered among the breeders, and each strives to produce the desired character as quickly as possible. Darwin says: 'The English judges decided that the comb of the Spanish cock, which had previously hung limply down, should stand erect, and in five years this end was achieved; they ordained that hens should have beards, and six years later fifty-seven of the groups of hens exhibited at the Crystal Palace in London were bearded.' The transformation does not always come about so quickly, however; thus, for instance, it required thirteen years before a certain breed of tumblers was furnished with a white head. But the breeders cause every visible part of the body to vary as seems good to them, and within the last fifty years they have really brought about very considerable changes in many breeds. Their method of procedure is carefully to select for breeding those birds which already possess a faint beginning of the desired character. Domesticated animals have on the whole a higher degree of variability than wild species, and the breeder takes advantage of this. Suppose it is a question of adding a crown of feathers to a smooth-headed breed, a bird is chosen which has the feathers on the back of the head a little longer than usual, and mated for breeding. Among its descendants there will probably be some which also exhibit these slightly prominent feathers, and possibly there may be one or other of them which has these feathers considerably lengthened. This one is then used for breeding, and by continually proceeding thus, and selecting for breeding, from generation to generation, only the individuals which approach most nearly to the desired end, the wished-for character is at last secured.

Thus it is not by crossing of different breeds, but by a patient accumulating of insignificant little variations through many generations, that the desired transformations are brought about. That is the magic wand by means of which the expert breeder produces his different breeds, we might almost say, as the sculptor moulds and remoulds his clay model according to his fancy. Quite according to his fancy the breeder has brought about all the fantastic forms we are familiar with among pigeons, mere variations which are of no use either to the bird itself or to man, which simply gratify man's whim without in many cases even satisfying his sense of beauty. For many of the existing breeds of pigeons, hens, and other domesticated animals, are anything but beautiful, the body being often unharmonious in structure and sometimes actually monstrous.

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.