Darwinism (Wallace)/Chapter IV

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Darwinism (Wallace) by Alfred Russel Wallace
Chapter IV - VARIATION OF DOMESTICATED ANIMALS AND CULTIVATED PLANTS

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—Circumstances 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 space 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 illustrate. It will also be necessary to explain how these variations have been increased and accumulated 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 quantity 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 quantity 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 constitutional vigour; but every breed possessing any exceptional quality is the result of the selection of variations occurring year after year and accumulated 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 accumulate 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 (Brassica 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 mass 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 mass 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 accumulated 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 accumulate 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 accumulate 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.[1] 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 illustrating the effects of slight variations accumulated 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 constitutional 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 cucumbers and gourds vary immensely, but the melon (Cucumis 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 cucumbers; and an Algerian variety, when ripe, cracks and falls to pieces, just as occurs in a wild gourd (C. momordica).[2]

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 quantities, 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 quantity, 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.[3]

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 cock 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."[4]

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.

RACE IX. Jacobin.—These curious birds have a hood of feathers almost enclosing the head and meeting in front of the neck. The wings and tail are unusually long.

RACE X. Trumpeter.—Distinguished by a tuft of feathers curling forwards over the beak, and the feet very much feathered. They obtain their name from the peculiar voice unlike that of any other pigeon. The coo is rapidly repeated, and is continued for several minutes. The feet are covered with feathers so large as often to appear like little wings.

RACE XI. comprises Laughers, Frill-backs, Nuns, Spots, and Swallows.—They are all very like the common rock-pigeon, but have each some slight peculiarity. The Laughers have a peculiar voice, supposed to resemble a laugh. The Nuns are white, with the head, tail, and primary wing-feathers black or red. The Spots are white, with the tail and a spot on the forehead red. The Swallows are slender, white in colour, with the head and wings of some darker colour.

Besides these races and sub-races a number of other kinds have been described, and about one hundred and fifty varieties can be distinguished. It is interesting to note that almost every part of the bird, whose variations can be noted and selected, has led to variations of a considerable extent, and many of these have necessitated changes in the plumage and in the skeleton quite as great as any that occur in the numerous distinct species of large genera. The form of the skull and beak varies enormously, so that the skulls of the Short-faced tumbler and some of the Carriers differ more than any wild pigeons, even those classed in distinct genera. The breadth and number of the ribs vary, as well as the processes on them; the number of the vertebrae and the length of the sternum also vary; and the perforations in the sternum vary in size and shape. The oil gland varies in development, and is sometimes absent. The number of the wing-feathers varies, and those of the tail to an enormous extent. The proportions of the leg and feet and the number of the scutellae also vary. The eggs also vary somewhat in size and shape; and the amount of downy clothing on the young bird, when first hatched, differs very considerably. Finally, the attitude of the body, the manner of walking, the mode of flight, and the voice, all exhibit modifications of the most remarkable kind.[5]

Acclimatisation.

A very important kind of variation is that constitutional change termed acclimatisation, which enables any organism to become gradually adapted to a different climate from the parent stock. As closely allied species often inhabit different countries possessing very different climates, we should expect to find cases illustrating this change among our domesticated animals and cultivated plants. A few examples will therefore be adduced showing that such constitutional variation does occur.

Among animals the cases are not numerous, because no systematic attempt has been made to select varieties for this special quality. It has, however, been observed that, though no European dogs thrive well in India, the Newfoundland dog, originating from a severe climate, can hardly be kept alive. A better case, perhaps, is furnished by merino sheep, which, when imported directly from England, do not thrive, while those which have been bred in the intermediate climate of the Cape of Good Hope do much better. When geese were first introduced into Bogota, they laid few eggs at long intervals, and few of the young survived. By degrees, however, the fecundity improved, and in about twenty years became equal to what it is in Europe. According to Garcilaso, when fowls were first introduced into Peru they were not fertile, whereas now they are as much so as in Europe.

Plants furnish much more important evidence. Our nurserymen distinguish in their catalogues varieties of fruit-trees which are more or less hardy, and this is especially the case in America, where certain varieties only will stand the severe climate of Canada. There is one variety of pear, the Forelle, which both in England and France withstood frosts that killed the flowers and buds of all other kinds of pears. Wheat, which is grown over so large a portion of the world, has become adapted to special climates. Wheat imported from India and sown in good wheat soil in England produced the most meagre ears; while wheat taken from France to the West Indian Islands produced either wholly barren spikes or spikes furnished with two or three miserable seeds, while West Indian seed by its side yielded an enormous harvest. The orange was very tender when first introduced into Italy, and continued so as long as it was propagated by grafts, but when trees were raised from seed many of these were found to be hardier, and the orange is now perfectly acclimatised in Italy. Sweet-peas (Lathyrus odoratus) imported from England to the Calcutta Botanic Gardens produced few blossoms and no seed; those from France flowered a little better, but still produced no seed, but plants raised from seed brought from Darjeeling in the Himalayas, but originally derived from England, flower and seed profusely in Calcutta.[6]

An observation by Mr. Darwin himself is perhaps even more instructive. He says: "On 24th May 1864 there was a severe frost in Kent, and two rows of scarlet runners (Phaseolus multiflorus) in my garden, containing 390 plants of the same age and equally exposed, were all blackened and killed except about a dozen plants. In an adjoining row of Fulmer's dwarf bean (Phaseolus vulgaris) one single plant escaped. A still more severe frost occurred four days afterwards, and of the dozen plants which had previously escaped only three survived; these were not taller or more vigorous than the other young plants, but they escaped completely, with not even the tips of their leaves browned. It was impossible to behold these three plants, with their blackened, withered, and dead brethren all around them, and not see at a glance that they differed widely in their constitutional power of resisting frost."

The preceding sketch of the variation that occurs among domestic animals and cultivated plants shows how wide it is in range and how great in amount; and we have good reason to believe that similar variation extends to all organised beings. In the class of fishes, for example, we have one kind which has been long domesticated in the East, the gold and silver carps; and these present great variation, not only of colour but in the form and structure of the fins and other external organs. In like manner, the only domesticated insects, hive bees and silkworm moths, present numbers of remarkable varieties which have been produced by the selection of chance variations just as in the case of plants and the higher animals.

Circumstances favourable to Selection by Man.

It may be supposed, that the systematic selection which has been employed for the purpose of improving the races of animals or plants useful to man is of comparatively recent origin, though some of the different races are known to have been in existence in very early times. But Mr. Darwin has pointed out, that unconscious selection must have begun to produce an effect as soon as plants were cultivated or animals domesticated by man. It would have been very soon observed that animals and plants produced their like, that seed of early wheat produced early wheat, that the offspring of very swift dogs were also swift, and as every one would try to have a good rather than a bad sort this would necessarily lead to the slow but steady improvement of all useful plants and animals subject to man's care. Soon there would arise distinct breeds, owing to the varying uses to which the animals and plants were put. Dogs would be wanted chiefly to hunt one kind of game in one part of the country and another kind elsewhere; for one purpose scent would be more important, for another swiftness, for another strength and courage, for yet another watchfulness and intelligence, and this would soon lead to the formation of very distinct races. In the case of vegetables and fruits, different varieties would be found to succeed best in certain soils and climates; some might be preferred on account of the quantity of food they produced, others for their sweetness and tenderness, while others might be more useful on account of their ripening at a particular season, and thus again distinct varieties would be established. An instance of unconscious selection leading to distinct results in modern times is afforded by two flocks of Leicester sheep which both originated from the same stock, and were then bred pure for upwards of fifty years by two gentlemen, Mr. Buckley and Mr. Burgess. Mr. Youatt, one of the greatest authorities on breeding domestic animals, says: "There is not a suspicion existing in the mind of any one at all acquainted with the subject that the owner of either of them has deviated in any one instance from the pure blood of Mr. Bakewell's original flock, and yet the difference between the sheep possessed by these two gentlemen is so great that they have the appearance of being quite different varieties." In this case there was no desire to deviate from the original breed, and the difference must have arisen from some slight difference of taste or judgment in selecting, each year, the parents for the next year's stock, combined perhaps with some direct effect of the slight differences of climate and soil on the two farms.

Most of our domesticated animals and cultivated plants have come to us from the earliest seats of civilisation in Western Asia or Egypt, and have therefore been the subjects of human care and selection for some thousands of years, the result being that, in many cases, we do not know the wild stock from which they originally sprang. The horse, the camel, and the common bull and cow are nowhere found in a wild state, and they have all been domesticated from remote antiquity. The original of the domestic fowl is still wild in India and the Malay Islands, and it was domesticated in India and China before 1400 B.C. It was introduced into Europe about 600 B.C. Several distinct breeds were known to the Romans about the commencement of the Christian era, and they have since spread all over the civilised world and been subjected to a vast amount of conscious and unconscious selection, to many varieties of climate and to differences of food; the result being seen in the wonderful diversity of breeds which differ quite as remarkably as do the different races of pigeons already described.

In the vegetable kingdom, most of the cereals—wheat, barley, etc.—are unknown as truly wild plants; and the same is the case with many vegetables, for De Candolle states that out of 157 useful cultivated plants thirty-two are quite unknown in a wild state, and that forty more are of doubtful origin. It is not improbable that most of these do exist wild, but they have been so profoundly changed by thousands of years of cultivation as to be quite unrecognisable. The peach is unknown in a wild state, unless it is derived from the common almond, on which point there is much difference of opinion among botanists and horticulturists.

The immense antiquity of most of our cultivated plants sufficiently explains the apparent absence of such useful productions in Australia and the Cape of Good Hope, notwithstanding that they both possess an exceedingly rich and varied flora. These countries having been, until a comparatively recent period, inhabited only by uncivilised men, neither cultivation nor selection has been carried on for a sufficiently long time. In North America, however, where there was evidently a very ancient if low form of civilisation, as indicated by the remarkable mounds, earthworks, and other prehistoric remains, maize was cultivated, though it was probably derived from Peru; and the ancient civilisation of that country and of Mexico has given rise to no fewer than thirty-three useful cultivated plants.

Conditions favourable to the production of Variations.

In order that plants and animals may be improved and modified to any considerable extent, it is of course essential that suitable variations should occur with tolerable frequency. There seem to be three conditions which are especially favourable to the production of variations: (1) That the particular species or variety should be kept in very large numbers; (2) that it should be spread over a wide area and thus subjected to a considerable diversity of physical conditions; and (3) that it should be occasionally crossed with some distinct but closely allied race. The first of these conditions is perhaps the most important, the chance of variations of any particular kind being increased in proportion to the quantity of the original stock and of its annual offspring. It has been remarked that only those breeders who keep large flocks can effect much improvement; and it is for the same reason that pigeons and fowls, which can be so easily and rapidly increased, and which have been kept in such large numbers by so great a number of persons, have produced such strange and numerous varieties. In like manner, nurserymen who grow fruit and flowers in large quantities have a great advantage over private amateurs in the production of new varieties.

Although I believe, for reasons which will be given further on, that some amount of variability is a constant and necessary property of all organisms, yet there appears to be good evidence to show that changed conditions of life tend to increase it, both by a direct action on the organisation and by indirectly affecting the reproductive system. Hence the extension of civilisation, by favouring domestication under altered conditions, facilitates the process of modification. Yet this change does not seem to be an essential condition, for nowhere has the production of extreme varieties of plants and flowers been carried farther than in Japan, where careful selection continued for many generations must have been the chief factor. The effect of occasional crosses often results in a great amount of variation, but it also leads to instability of character, and is therefore very little employed in the production of fixed and well-marked races. For this purpose, in fact, it has to be carefully avoided, as it is only by isolation and pure breeding that any specially desired qualities can be increased by selection. It is for this reason that among savage peoples, whose animals run half wild, little improvement takes place; and the difficulty of isolation also explains why distinct and pure breeds of cats are so rarely met with. The wide distribution of useful animals and plants from a very remote epoch has, no doubt, been a powerful cause of modification, because the particular breed first introduced into each country has often been kept pure for many years, and has also been subjected to slight differences of conditions. It will also usually have been selected for a somewhat different purpose in each locality, and thus very distinct races would soon originate.

The important physiological effects of crossing breeds or strains, and the part this plays in the economy of nature, will be explained in a future chapter.

Concluding Remarks.

The examples of variation now adduced—and these might have been almost indefinitely increased—will suffice to show that there is hardly an organ or a quality in plants or animals which has not been observed to vary; and further, that whenever any of these variations have been useful to man he has been able to increase them to a marvellous extent by the simple process of always preserving the best varieties to breed from. Along with these larger variations others of smaller amount occasionally appear, sometimes in external, sometimes in internal characters, the very bones of the skeleton often changing slightly in form, size, or number; but as these secondary characters have been of no use to man, and have not been specially selected by him, they have, usually, not been developed to any great amount except when they have been closely dependent on those external characters which he has largely modified.

As man has considered only utility to himself, or the satisfaction of his love of beauty, of novelty, or merely of something strange or amusing, the variations he has thus produced have something of the character of monstrosities. Not only are they often of no use to the animals or plants themselves, but they are not unfrequently injurious to them. In the Tumbler pigeons, for instance, the habit of tumbling is sometimes so excessive as to injure or kill the bird; and many of our highly-bred animals have such delicate constitutions that they are very liable to disease, while their extreme peculiarities of form or structure would often render them quite unfit to live in a wild state. In plants, many of our double flowers, and some fruits, have lost the power of producing seed, and the race can thus be continued only by means of cuttings or grafts. This peculiar character of domestic productions distinguishes them broadly from wild species and varieties, which, as will be seen by and by, are necessarily adapted in every part of their organisation to the conditions under which they have to live. Their importance for our present inquiry depends on their demonstrating the occurrence of incessant slight variations in all parts of an organism, with the transmission to the offspring of the special characteristics of the parents; and also, that all such slight variations are capable of being accumulated by selection till they present very large and important divergencies from the ancestral stock.

We thus see, that the evidence as to variation afforded by animals and plants under domestication strikingly accords with that which we have proved to exist in a state of nature. And it is not at all surprising that it should be so, since all the species were in a state of nature when first domesticated or cultivated by man, and whatever variations occur must be due to purely natural causes. Moreover, on comparing the variations which occur in any one generation of domesticated animals with those which we know to occur in wild animals, we find no evidence of greater individual variation in the former than in the latter. The results of man's selection are more striking to us because we have always considered the varieties of each domestic animal to be essentially identical, while those which we observe in a wild state are held to be essentially diverse. The greyhound and the spaniel seem wonderful, as varieties of one animal produced by man's selection ; while we think little of the diversities of the fox and the wolf, or the horse and the zebra, because we have been accustomed to look upon them as radically distinct animals, not as the results of nature's selection of the varieties of a common ancestor.

  1. Darwin, Animals and Plants under Domestication, vol. i. p. 322.
  2. These facts are taken from Darwin's Domesticated Animals and Cultivated Plants, vol. i. pp. 359, 360, 392-401; vol. ii. pp. 231, 275, 330.
  3. See Darwin's Animals and Plants under Domestication, vol. i. pp. 40-42.
  4. Mr. Brent in Journal of Horticulture, 1861, p. 76; quoted by Darwin, Animals and Plants under Domestication, vol. i. p. 151.
  5. This account of domestic pigeons is greatly condensed from Mr. Darwin's work already referred to.
  6. Animals and Plants under Domestication, vol. ii. pp. 307-311.