The Variation of Animals and Plants under Domestication — Volume 2

by Charles Darwin

As almost every part of the organisation becomes highly variable under domestication, and as variations are easily selected both consciously and unconsciously, it is very difficult to distinguish between the effects of the selection of indefinite variations and the direct action of the conditions of life. For instance, it is possible that the feet of our water-dogs and of the American dogs which have to travel much over the snow, may have become partially webbed from the stimulus of widely extending their toes; but it is more probable that the webbing, like the membrane between the toes of certain pigeons, spontaneously appeared and was afterwards increased by the best swimmers and the best snow-travellers being preserved during many generations. A fancier who wished to decrease the size of his bantams or tumbler-pigeons would never think of starving them, but would select the smallest individuals which spontaneously appeared. Quadrupeds are sometimes born destitute of hair and hairless breeds have been formed, but there is no reason to believe that this is caused by a hot climate. Within the tropics heat often causes sheep to lose their fleeces; on the other hand, wet and cold act as a direct stimulus to the growth of hair; but who will pretend to decide how far the thick fur of arctic animals, or their white colour, is due to the direct action of a severe climate, and how far to the preservation of the best-protected individuals during a long succession of generations?

Of all the laws governing variability, that of correlation is one of the most important. In many cases of slight deviations of structure as well as of grave monstrosities, we cannot even conjecture what is the nature of the bond of connexion. But between homologous parts—between the fore and hind limbs— between the hair, hoofs, horns, and teeth—which are closely similar during their early development and which are exposed to similar conditions, we can see that they would be eminently liable to be modified in the same manner. Homologous parts, from having the same nature, are apt to blend together, and, when many exist, to vary in number.

Although every variation is either directly or indirectly caused by some change in the surrounding conditions, we must never forget that the nature of the organisation which is acted on, is by far the more important factor in the result. We see this in different organisms, which when placed under similar conditions vary in a different manner, whilst closely-allied organisms under dissimilar conditions often vary in nearly the same manner. We see this, in the same modification frequently reappearing in the same variety at long intervals of time, and likewise in the several striking cases given of analogous or parallel variations. Although some of these latter cases are due to reversion, others cannot thus be accounted for.

From the indirect action of changed conditions on the organisation, owing to the reproductive organs being thus affected—from the direct action of such conditions, and these will cause the individuals of the same species either to vary in the same manner, or differently in accordance with slight differences in their constitution—from the effects of the increased or decreased use of parts—and from correlation,—the variability of our domesticated productions is complicated to an extreme degree. The whole organisation becomes slightly plastic. Although each modification must have its own exciting cause, and though each is subjected to law, yet we can so rarely trace the precise relation between cause and effect, that we are tempted to speak of variations as if they arose spontaneously. We may even call them accidental, but this must be only in the sense in which we say that a fragment of rock dropped from a height owes its shape to accident.

It may be worth while briefly to consider the result of the exposure to unnatural conditions of a large number of animals of the same species and allowed to cross freely with no selection of any kind, and afterwards to consider the result when selection is brought into play. Let us suppose that 500 wild rock-pigeons were confined in their native land in an aviary and fed in the same manner as pigeons usually are; and that they were not allowed to increase in number. As pigeons propagate so rapidly, I suppose that a thousand or fifteen hundred birds would have to be annually killed. After several generations had been thus reared, we may feel sure that some of the young birds would vary, and the variations would tend to be inherited; for at the present day slight deviations of structure often occur and are inherited. It would be tedious even to enumerate the multitude of points which still go on varying or have recently varied. Many variations would occur in correlation with one another, as the length of the wing and tail feathers—the number of the primary wing-feathers, as well as the number and breadth of the ribs, in correlation with the size and form of the body—the number of the scutellae with the size of the feet—the length of the tongue with the length of the beak—the size of the nostrils and eyelids and the form of lower jaw in correlation with the development of wattle—the nakedness of the young with the future colour of the plumage—the size of the feet with that of the beak, and other such points. Lastly, as our birds are supposed to be confined in an aviary, they would use their wings and legs but little, and certain parts of the skeleton, such as the sternum, scapulae and feet, would in consequence become slightly reduced in size.

As in our assumed case many birds have to be indiscriminately killed every year, the chances are against any new variety surviving long enough to breed. And as the variations which arise are of an extremely diversified nature, the chances are very great against two birds pairing which have varied in the same manner; nevertheless, a varying bird even when not thus paired would occasionally transmit its character to its young; and these would not only be exposed to the same conditions which first caused the variation in question to appear, but would in addition inherit from their modified parent a tendency again to vary in the same manner. So that, if the conditions decidedly tended to induce some particular variation, all the birds might in the course of time become similarly modified. But a far commoner result would be, that one bird would vary in one way and another bird in another way; one would be born with a beak a little longer, and another with a shorter beak; one would gain some black feathers, another some white or red feathers. And as these birds would be continually intercrossing, the final result would be a body of individuals differing from each other in many ways, but only slightly; yet more than did the original rock-pigeons. But there would not be the least tendency towards the formation of several distinct breeds.

If two separate lots of pigeons were treated in the manner just described, one in England and the other in a tropical country, the two lots being supplied with different kinds of food, would they after many generations differ? When we reflect on the cases given in the twenty-third chapter, and on such facts as the difference in former times between the breeds of cattle, sheep, etc., in almost every district of Europe, we are strongly inclined to admit that the two lots would be differently modified through the influence of climate and food. But the evidence on the definite action of changed conditions is in most cases insufficient; and, with respect to pigeons, I have had the opportunity of examining a large collection of domesticated kinds, sent to me by Sir W. Elliot from India, and they varied in a remarkably similar manner with our European birds.

If two distinct breeds were mingled together in equal numbers, there is reason to suspect that they would to a certain extent prefer pairing with their own kind; but they would often intercross. From the greater vigour and fertility of the crossed offspring, the whole body would by this means become interblended sooner than would otherwise have occurred. From certain breeds being prepotent over others, it does not follow that the interblended progeny would be strictly intermediate in character. I have, also, proved that the act of crossing in itself gives a strong tendency to reversion, so that the crossed offspring would tend to revert to the state of the aboriginal rock- pigeon; and in the course of time they would probably be not much more heterogeneous in character than in our first case, when birds of the same breed were confined together.

I have just said that the crossed offspring would gain in vigour and fertility. From the facts given in the seventeenth chapter there can be no doubt of this fact; and there can be little doubt, though the evidence on this head is not so easily acquired, that long-continued close interbreeding leads to evil results. With hermaphrodites of all kinds, if the sexual elements of the same individual habitually acted on each other, the closest possible interbreeding would be perpetual. But we should bear in mind that the structure of all hermaphrodite animals, as far as I can learn, permits and frequently necessitates a cross with a distinct individual. With hermaphrodite plants we incessantly meet with elaborate and perfect contrivances for this same end. It is no exaggeration to assert that, if the use of the talons and tusks of a carnivorous animal, or of the plumes and hooks on a seed, may be safely inferred from their structure, we may with equal safety infer that many flowers are constructed for the express purpose of ensuring a cross with a distinct plant. From these various considerations, not to mention the result of a long series of experiments which I have tried, the conclusion arrived at in the chapter just referred to—namely, that great good of some kind is derived from the sexual concourse of distinct individuals—must be admitted.

To return to our illustration: we have hitherto assumed that the birds were kept down to the same number by indiscriminate slaughter; but if the least choice be permitted in their preservation, the whole result will be changed. Should the owner observe any slight variation in one of his birds, and wish to obtain a breed thus characterised, he would succeed in a surprisingly short time by careful selection. As any part which has once varied generally goes on varying in the same direction, it is easy, by continually preserving the most strongly marked individuals, to increase the amount of difference up to a high, predetermined standard of excellence. This is methodical selection.