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[Footnote 25: See _Trans. Entomological Society of London_, 1887, p.

24.]

[Footnote 26: _Nature_, vol. xix. p. 554.]

[Footnote 27: _Nature_, vol. xvi. p. 163; and vol. xi. p. 227.]

[Footnote 28: _Ibid._, vol. x.x.xi. (1885), p. 533.]

[Footnote 29: _Nature_, vol. xxvi. p. 81.]

[Footnote 30: _Animals and Plants under Domestication_, vol. ii. p.

258.]

CHAPTER IV

VARIATION OF DOMESTICATED ANIMALS AND CULTIVATED PLANTS

The facts of variation and artificial selection--Proofs of the generality of variation--Variations of apples and melons--Variations of flowers--Variations of domestic animals--Domestic pigeons--Acclimatisation--Circ.u.mstances favourable to selection by man--Conditions favourable to variation--Concluding remarks.

Having so fully discussed variation under nature it will be unnecessary to devote so much s.p.a.ce to domesticated animals and cultivated plants, especially as Mr. Darwin has published two remarkable volumes on the subject where those who desire it may obtain ample information. A general sketch of the more important facts will, however, be given, for the purpose of showing how closely they correspond with those described in the preceding chapter, and also to point out the general principles which they ill.u.s.trate. It will also be necessary to explain how these variations have been increased and acc.u.mulated by artificial selection, since we are thereby better enabled to understand the action of natural selection, to be discussed in the succeeding chapter.

_The facts of Variation and Artificial Selection._

Every one knows that in each litter of kittens or of puppies no two are alike. Even in the case in which several are exactly alike in colours, other differences are always perceptible to those who observe them closely. They will differ in size, in the proportions of their bodies and limbs, in the length or texture of their hairy covering, and notably in their disposition. They each possess, too, an individual countenance, almost as varied when closely studied as that of a human being; not only can a shepherd distinguish every sheep in his flock, but we all know that each kitten in the successive families of our old favourite cat has a face of its own, with an expression and individuality distinct from all its brothers and sisters. Now this individual variability exists among all creatures whatever, which we can closely observe, even when the two parents are very much alike and have been matched in order to preserve some special breed. The same thing occurs in the vegetable kingdom. All plants raised from seed differ more or less from each other. In every bed of flowers or of vegetables we shall find, if we look closely, that there are countless small differences, in the size, in the mode of growth, in the shape or colour of the leaves, in the form, colour, or markings of the flowers, or in the size, form, colour, or flavour of the fruit. These differences are usually small, but are yet easily seen, and in their extremes are very considerable; and they have this important quality, that they have a tendency to be reproduced, and thus by careful breeding any particular variation or group of variations can be increased to an enormous extent--apparently to any extent not incompatible with the life, growth, and reproduction of the plant or animal.

The way this is done is by artificial selection, and it is very important to understand this process and its results. Suppose we have a plant with a small edible seed, and we want to increase the size of that seed. We grow as large a quant.i.ty of it as possible, and when the crop is ripe we carefully choose a few of the very largest seeds, or we may by means of a sieve sort out a quant.i.ty of the largest seeds. Next year we sow only these large seeds, taking care to give them suitable soil and manure, and the result is found to be that the _average_ size of the seeds is larger than in the first crop, and that the largest seeds are now somewhat larger and more numerous. Again sowing these, we obtain a further slight increase of size, and in a very few years we obtain a greatly improved race, which will always produce larger seeds than the unimproved race, even if cultivated without any special care. In this way all our fine sorts of vegetables, fruits, and flowers have been obtained, all our choice breeds of cattle or of poultry, our wonderful race-horses, and our endless varieties of dogs. It is a very common but mistaken idea that this improvement is due to crossing and feeding in the case of animals, and to improved cultivation in the case of plants.

Crossing is occasionally used in order to obtain a combination of qualities found in two distinct breeds, and also because it is found to increase the const.i.tutional vigour; but every breed possessing any exceptional quality is the result of the selection of variations occurring year after year and acc.u.mulated in the manner just described.

Purity of breed, with repeated selection of the best varieties of that breed, is the foundation of all improvement in our domestic animals and cultivated plants.

_Proofs of the Generality of Variation._

Another very common error is, that variation is the exception, and rather a rare exception, and that it occurs only in one direction at a time--that is, that only one or two of the numerous possible modes of variation occur at the same time. The experience of breeders and cultivators, however, proves that variation is the rule instead of the exception, and that it occurs, more or less, in almost every direction.

This is shown by the fact that different species of plants and animals have required different _kinds_ of modification to adapt them to our use, and we have never failed to meet with variation _in that particular direction_, so as to enable us to acc.u.mulate it and so to produce ultimately a large amount of change in the required direction. Our gardens furnish us with numberless examples of this property of plants.

In the cabbage and lettuce we have found variation in the size and mode of growth of the leaf, enabling us to produce by selection the almost innumerable varieties, some with solid heads of foliage quite unlike any plant in a state of nature, others with curiously wrinkled leaves like the savoy, others of a deep purple colour used for pickling. From the very same species as the cabbage (Bra.s.sica oleracea) have arisen the broccoli and cauliflower, in which the leaves have undergone little alteration, while the branching heads of flowers grow into a compact ma.s.s forming one of our most delicate vegetables. The brussels sprouts are another form of the same plant, in which the whole mode of growth has been altered, numerous little heads of leaves being produced on the stem. In other varieties the ribs of the leaves are thickened so as to become themselves a culinary vegetable; while, in the Kohlrabi, the stem grows into a turnip-like ma.s.s just above ground. Now all these extraordinarily distinct plants come from one original species which still grows wild on our coasts; and it must have varied in all these directions, otherwise variations could not have been acc.u.mulated to the extent we now see them. The flowers and seeds of all these plants have remained nearly stationary, because no attempt has been made to acc.u.mulate the slight variations that no doubt occur in them.

If now we turn to another set of plants, the turnips, radishes, carrots, and potatoes, we find that the roots or underground tubers have been wonderfully enlarged and improved, and also altered in shape and colour, while the stems, leaves, flowers, and fruits have remained almost unchanged. In the various kinds of peas and beans it is the pod or fruit and the seed that has been subjected to selection, and therefore greatly modified; and it is here very important to notice that while all these plants have undergone cultivation in a great variety of soils and climates, with different manures and under different systems, yet the flowers have remained but little altered, those of the broad bean, the scarlet-runner, and the garden-pea, being nearly the same in all the varieties. This shows us how little change is produced by mere cultivation, or even by variety of soil and climate, if there is no _selection_ to preserve and acc.u.mulate the small variations that are continually occurring. When, however, a great amount of modification has been effected in one country, change to another country produces a decided effect. Thus it has been found that some of the numerous varieties of maize produced and cultivated in the United States change considerably, not only in their size and colour, but even in the shape of the seed when grown for a few successive years in Germany.[31] In all our cultivated fruit trees the fruits vary immensely in shape, size, colour, flavour, time of ripening, and other qualities, while the leaves and flowers usually differ so little that they are hardly distinguishable except to a very close observer.

_Variations of Apples and of Melons._

The most remarkable varieties are afforded by the apple and the melon, and some account of these will be given as ill.u.s.trating the effects of slight variations acc.u.mulated by selection. All our apples are known to have descended from the common crab of our hedges (Pyrus malus), and from this at least a thousand distinct varieties have been produced.

These differ greatly in the size and form of the fruit, in its colour, and in the texture of the skin. They further differ in the time of ripening, in their flavour, and in their keeping properties; but apple trees also differ in many other ways. The foliage of the different varieties can often be distinguished by peculiarities of form and colour, and it varies considerably in the time of its appearance; in some hardly a leaf appears till the tree is in full bloom, while others produce their leaves so early as almost to hide the flowers. The flowers differ in size and colour, and in one case in structure also, that of the St. Valery apple having a double calyx with ten divisions, and fourteen styles with oblique stigmas, but without stamens or corolla.

The flowers, therefore, have to be fertilised with the pollen from other varieties in order to produce fruit. The pips or seeds differ also in shape, size, and colour; some varieties are liable to canker more than others, while the Winter Majetin and one or two others have the strange const.i.tutional peculiarity of never being attacked by the mealy bug even when all the other trees in the same orchard are infested with it.

All the cuc.u.mbers and gourds vary immensely, but the melon (Cuc.u.mis melo) exceeds them all. A French botanist, M. Naudin, devoted six years to their study. He found that previous botanists had described thirty distinct species, as they thought, which were really only varieties of melons. They differ chiefly in their fruits, but also very much in foliage and mode of growth. Some melons are only as large as small plums, others weigh as much as sixty-six pounds. One variety has a scarlet fruit. Another is not more than an inch in diameter, but sometimes more than a yard in length, twisting about in all directions like a serpent. Some melons are exactly like cuc.u.mbers; and an Algerian variety, when ripe, cracks and falls to pieces, just as occurs in a wild gourd (C. momordica).[32]

_Variations of Flowers._

Turning to flowers, we find that in the same genus as our currant and gooseberry, which we have cultivated for their fruits, there are some ornamental species, as the Ribes sanguinea, and in these the flowers have been selected so as to produce deep red, pink, or white varieties.

When any particular flower becomes fashionable and is grown in large quant.i.ties, variations are always met with sufficient to produce great varieties of tint or marking, as shown by our roses, auriculas, and geraniums. When varied leaves are required, it is found that a number of plants vary sufficiently in this direction also, and we have zonal geraniums, variegated ivies, gold and silver marked hollies, and many others.

_Variations of Domestic Animals._

Coming now to our domesticated animals, we find still more extraordinary cases; and it appears as if any special quality or modification in an animal can be obtained if we only breed it in sufficient quant.i.ty, watch carefully for the required variations, and carry on selection with patience and skill for a sufficiently long period. Thus, in sheep we have enormously increased the wool, and have obtained the power of rapidly forming flesh and fat; in cows we have increased the production of milk; in horses we have obtained strength, endurance, or speed, and have greatly modified size, form, and colour; in poultry we have secured various colours of plumage, increase of size, and almost perpetual egg-laying. But it is in dogs and pigeons that the most marvellous changes have been effected, and these require our special attention.

Our various domestic dogs are believed to have originated from several distinct wild species, because in every part of the world the native dogs resemble some wild dogs or wolves of the same country. Thus perhaps several species of wolves and jackals were domesticated in very early times, and from breeds derived from these, crossed and improved by selection, our existing dogs have descended. But this intermixture of distinct species will go a very little way in accounting for the peculiarities of the different breeds of dogs, many of which are totally unlike any wild animal. Such is the case with greyhounds, bloodhounds, bulldogs, Blenheim spaniels, terriers, pugs, turnspits, pointers, and many others; and these differ so greatly in size, shape, colour, and habits, as well as in the form and proportions of all the different parts of the body, that it seems impossible that they could have descended from any of the known wild dogs, wolves, or allied animals, none of which differ nearly so much in size, form, and proportions. We have here a remarkable proof that variation is not confined to superficial characters--to the colour, hair, or external appendages, when we see how the entire skeletons of such forms as the greyhound and the bulldog have been gradually changed in opposite directions till they are both completely unlike that of any known wild animal, recent or extinct. These changes have been the result of some thousands of years of domestication and selection, different breeds being used and preserved for different purposes; but some of the best breeds are known to have been improved and perfected in modern times. About the middle of the last century a new and improved kind of foxhound was produced; the greyhound was also greatly improved at the end of the last century, while the true bulldog was brought to perfection about the same period.

The Newfoundland dog has been so much changed since it was first imported that it is now quite unlike any existing native dog in that island.[33]

_Domestic Pigeons._

The most remarkable and instructive example of variation produced by human selection is afforded by the various races and breeds of domestic pigeons, not only because the variations produced are often most extraordinary in amount and diverse in character, but because in this case there is no doubt whatever that all have been derived from one wild species, the common rock-pigeon (Columba livia). As this is a very important point it is well to state the evidence on which the belief is founded. The wild rock-pigeon is of a slaty-blue colour, the tail has a dark band across the end, the wings have two black bands, and the outer tail-feathers are edged with white at the base. No other wild pigeon in the world has this combination of characters. Now in every one of the domestic varieties, even the most extreme, all the above marks, even to the white edging of the outer tail-feathers, are sometimes found perfectly developed. When birds belonging to two distinct breeds are crossed one or more times, neither of the parents being blue, or having any of the above-named marks, the mongrel offspring are very apt to acquire some of these characters. Mr. Darwin gives instances which he observed himself. He crossed some white fantails with some black barbs, and the mongrels were black, brown, or mottled. He also crossed a barb with a spot, which is a white bird with a red tail and red spot on the forehead, and the mongrel offspring were dusky and mottled. On now crossing these two sets of mongrels with each other, he obtained a bird of a beautiful blue colour, with the barred and white edged tail, and double-banded wings, so as almost exactly to resemble a wild rock-pigeon. This bird was descended in the second generation from a pure white and pure black bird, both of which when unmixed breed their kind remarkably true. These facts, well known to experienced pigeon-fanciers, together with the habits of the birds, which all like to nest in holes, or dovecots, not in trees like the great majority of wild pigeons, have led to the general belief in the single origin of all the different kinds.

In order to afford some idea of the great differences which exist among domesticated pigeons, it will be well to give a brief abstract of Mr.

Darwin's account of them. He divides them into eleven distinct races, most of which have several sub-races.

RACE I. _Pouters_.--These are especially distinguished by the enormously enlarged crop, which can be so inflated in some birds as almost to conceal the beak. They are very long in the body and legs and stand almost upright, so as to present a very distinct appearance. Their skeleton has become modified, the ribs being broader and the vertebrae more numerous than in other pigeons.

RACE II. _Carriers_.--These are large, long-necked birds, with a long pointed beak, and the eyes surrounded with a naked carunculated skin or wattle, which is also largely developed at the base of the beak. The opening of the mouth is unusually wide. There are several sub-races, one being called Dragons.

RACE III. _Runts_.--These are very large-bodied, long-beaked pigeons, with naked skin round the eyes. The wings are usually very long, the legs long, and the feet large, and the skin of the neck is often red.

There are several sub-races, and these differ very much, forming a series of links between the wild rock-pigeon and the carrier.

RACE IV. _Barbs_.--These are remarkable for their very short and thick beak, so unlike that of most pigeons that fanciers compare it with that of a bullfinch. They have also a naked carunculated skin round the eyes, and the skin over the nostrils swollen.

RACE V. _Fantails_.--Short-bodied and rather small-beaked pigeons, with an enormously developed tail, consisting usually of from fourteen to forty feathers instead of twelve, the regular number in all other pigeons, wild and tame. The tail spreads out like a fan and is usually carried erect, and the bird bends back its slender neck, so that in highly-bred varieties the head touches the tail. The feet are small, and they walk stiffly.

RACE VI. _Turbits and Owls_.--These are characterised by the feathers of the middle of neck and breast in front spreading out irregularly so as to form a frill. The Turbits also have a crest on the head, and both have the beak exceedingly short.

RACE VII. _Tumblers_.--- These have a small body and short beak, but they are specially distinguished by the singular habit of tumbling over backwards during flight. One of the sub-races, the Indian Lotan or Ground tumbler, if slightly shaken and placed on the ground, will immediately begin tumbling head over heels until taken up and soothed.

If not taken up, some of them will go on tumbling till they die. Some English tumblers are almost equally persistent. A writer, quoted by Mr.

Darwin, says that these birds generally begin to tumble almost as soon as they can fly; "at three months old they tumble well, but still fly strong; at five or six months they tumble excessively; and in the second year they mostly give up flying, on account of their tumbling so much and so close to the ground. Some fly round with the flock, throwing a clean summersault every few yards till they are obliged to settle from giddiness and exhaustion. These are called Air-tumblers, and they commonly throw from twenty to thirty summersaults in a minute, each clear and clean. I have one red c.o.c.k that I have on two or three occasions timed by my watch, and counted forty summersaults in the minute. At first they throw a single summersault, then it is double, till it becomes a continuous roll, which puts an end to flying, for if they fly a few yards over they go, and roll till they reach the ground.

Thus I had one kill herself, and another broke his leg. Many of them turn over only a few inches from the ground, and will tumble two or three times in flying across their loft. These are called House-tumblers from tumbling in the house. The act of tumbling seems to be one over which they have no control, an involuntary movement which they seem to try to prevent. I have seen a bird sometimes in his struggles fly a yard or two straight upwards, the impulse forcing him backwards while he struggles to go forwards."[34]

The Short-faced tumblers are an improved sub-race which have almost lost the power of tumbling, but are valued for possessing some other characteristics in an extreme degree. They are very small, have almost globular heads, and a very minute beak, so that fanciers say the head of a perfect bird should resemble a cherry with a barleycorn stuck in it.

Some of these weigh less than seven ounces, whereas the wild rock-pigeon weighs about fourteen ounces. The feet, too, are very short and small, and the middle toe has twelve or thirteen instead of fourteen or fifteen scutellae. They have often only nine primary wing-feathers instead of ten as in all other pigeons.

RACE VIII. _Indian Frill-back_.--In these birds the beak is very short, and the feathers of the whole body are reversed or turn backwards.

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