XX. SELECTION BY MAN…

XX. SELECTION BY MAN

SELECTION A DIFFICULT ART—METHODICAL, UNCONSCIOUS, AND NATURAL SELECTION—RESULTS OF METHODICAL SELECTION—CARE TAKEN IN SELECTION—SELECTION WITH PLANTS—SELECTION CARRIED ON BY THE ANCIENTS AND BY SEMI-CIVILISED PEOPLE—UNIMPORTANT CHARACTERS OFTEN ATTENDED TO—UNCONSCIOUS SELECTION—AS CIRCUMSTANCES SLOWLY CHANGE, SO HAVE OUR DOMESTICATED ANIMALS CHANGED THROUGH THE ACTION OF UNCONSCIOUS SELECTION—INFLUENCE OF DIFFERENT BREEDERS ON THE SAME SUB-VARIETY—PLANTS AS AFFECTED BY UNCONSCIOUS SELECTION—EFFECTS OF SELECTION AS SHOWN BY THE GREAT AMOUNT OF DIFFERENCE IN THE PARTS MOST VALUED BY MAN.

The power of Selection, whether exercised by man, or brought into play under nature through the struggle for existence and the consequent survival of the fittest, absolutely depends on the variability of organic beings. Without variability nothing can be effected; slight individual differences, however, suffice for the work, and are probably the chief or sole means in the production of new species. Hence our discussion on the causes and laws of variability ought in strict order to have preceded the present subject, as well as inheritance, crossing, etc.; but practically the present arrangement has been found the most convenient. Man does not attempt to cause variability; though he unintentionally effects this by exposing organisms to new conditions of life, and by crossing breeds already formed. But variability being granted, he works wonders. Unless some degree of selection be exercised, the free commingling of the individuals of the same variety soon obliterates, as we have previously seen, the slight differences which arise, and gives uniformity of character to the whole body of individuals. In separated districts, long-continued exposure to different conditions of life may produce new races without the aid of selection; but to this subject of the direct action of the conditions of life I shall recur in a future chapter.

When animals or plants are born with some conspicuous and firmly inherited new character, selection is reduced to the preservation of such individuals, and to the subsequent prevention of crosses; so that nothing more need be said on the subject. But in the great majority of cases a new character, or some superiority in an old character, is at first faintly pronounced, and is not strongly inherited; and then the full difficulty of selection is experienced. Indomitable patience, the finest powers of discrimination, and sound judgment must be exercised during many years. A clearly predetermined object must be kept steadily in view. Few men are endowed with all these qualities, especially with that of discriminating very slight differences; judgment can be acquired only by long experience; but if any of these qualities be wanting, the labour of a life may be thrown away. I have been astonished when celebrated breeders, whose skill and judgment have been proved by their success at exhibitions, have shown me their animals, which appeared all alike, and have assigned their reasons for matching this and that individual. The importance of the great principle of Selection mainly lies in this power of selecting scarcely appreciable differences, which nevertheless are found to be transmissible, and which can be accumulated until the result is made manifest to the eyes of every beholder.

The principle of selection may be conveniently divided into three kinds. Methodical selection is that which guides a man who systematically endeavours to modify a breed according to some predetermined standard. Unconscious selection is that which follows from men naturally preserving the most valued and destroying the less valued individuals, without any thought of altering the breed; and undoubtedly this process slowly works great changes. Unconscious selection graduates into methodical, and only extreme cases can be distinctly separated; for he who preserves a useful or perfect animal will generally breed from it with the hope of getting offspring of the same character; but as long as he has not a predetermined purpose to improve the breed, he may be said to be selecting unconsciously.[1] Lastly, we have Natural selection, which implies that the individuals which are best fitted for the complex, and in the course of ages changing conditions to which they are exposed, generally survive and procreate their kind. With domestic productions, natural selection comes to a certain extent into action, independently of, and even in opposition to, the will of man.

Methodical Selection.—What man has effected within recent times in England by methodical selection is clearly shown by our exhibitions of improved quadrupeds and fancy birds. With respect to cattle, sheep, and pigs, we owe their great improvement to a long series of well-known names—Bakewell, Coiling, Ellman, Bates, Jonas Webb, Lords Leicester and Western, Fisher Hobbs, and others. Agricultural writers are unanimous on the power of selection: any number of statements to this effect could be quoted; a few will suffice. Youatt, a sagacious and experienced observer, writes[2] the principle of selection is “that which enables the agriculturist, not only to modify the character of his flock, but to change it altogether.” A great breeder of Shorthorns[3] says, “In the anatomy of the shoulder modern breeders have made great improvement on the Ketton shorthorns by correcting the defect in the knuckle or shoulder-joint, and by laying the top of the shoulder more snugly in the crop, and thereby filling up the hollow behind it . . . The eye has its fashion at different periods: at one time the eye high and outstanding from the head, and at another time the sleepy eye sunk into the head; but these extremes have merged into the medium of a full, clear and prominent eye with a placid look.”

Again, hear what an excellent judge of pigs[4] says: “The legs should be no longer than just to prevent the animal’s belly from trailing on the ground. The leg is the least profitable portion of the hog, and we therefore require no more of it than is absolutely necessary for the support of the rest.” Let any one compare the wild-boar with any improved breed, and he will see how effectually the legs have been shortened.

Few persons, except breeders, are aware of the systematic care taken in selecting animals, and of the necessity of having a clear and almost prophetic vision into futurity. Lord Spencer’s skill and judgment were well known; and he writes,[5] “It is therefore very desirable, before any man commences to breed either cattle or sheep, that he should make up his mind to the shape and qualities he wishes to obtain, and steadily pursue this object.” Lord Somerville, in speaking of the marvellous improvement of the New Leicester sheep, effected by Bakewell and his successors, says, “It would seem as if they had first drawn a perfect form, and then given it life.” Youatt[6] urges the necessity of annually drafting each flock, as many animals will certainly degenerate “from the standard of excellence which the breeder has established in his own mind.” Even with a bird of such little importance as the canary, long ago (1780-1790) rules were established, and a standard of perfection was fixed according to which the London fanciers tried to breed the several sub-varieties.[7] A great winner of prizes at the Pigeon-shows,[8] in describing the short-faced Almond Tumbler, says, “There are many first-rate fanciers who are particularly partial to what is called the goldfinch-beak, which is very beautiful; others say, take a full-size round cherry then take a barleycorn, and judiciously placing and thrusting it into the cherry, form as it were your beak; and that is not all, for it will form a good head and beak, provided, as I said before, it is judiciously done; others take an oat; but as I think the goldfinch-beak the handsomest, I would advise the inexperienced fancier to get the head of a goldfinch, and keep it by him for his observation.” Wonderfully different as are the beaks of the rock pigeon and goldfinch, the end has undoubtedly been nearly gained, as far as external shape and proportions are concerned.

Not only should our animals be examined with the greatest care whilst alive, but, as Anderson remarks[9] their carcases should be scrutinised, “so as to breed from the descendants of such only as, in the language of the butcher, cut up well.” The “grain of the meat” in cattle, and its being well marbled with fat,[10] and the greater or less accumulation of fat in the abdomen of our sheep, have been attended to with success. So with poultry, a writer,[11] speaking of Cochin-China fowls, which are said to differ much in the quality of their flesh, says, “the best mode is to purchase two young brother-cocks, kill, dress, and serve up one; if he be indifferent, similarly dispose of the other, and try again; if, however, he be fine and well-flavoured, his brother will not be amiss for breeding purposes for the table.”

The great principle of the division of labour has been brought to bear on selection. In certain districts[12] “the breeding of bulls is confined to a very limited number of persons, who by devoting their whole attention to this department, are able from year to year to furnish a class of bulls which are steadily improving the general breed of the district.” The rearing and letting of choice rams has long been, as is well known, a chief source of profit to several eminent breeders. In parts of Germany this principle is carried with merino sheep to an extreme point.[13] So “important is the proper selection of breeding animals considered, that the best flock-masters do not trust to their own judgment or to that of their shepherds, but employ persons called ‘sheep-classifiers’ who make it their special business to attend to this part of the management of several flocks, and thus to preserve, or if possible to improve, the best qualities of both parents in the lambs.” In Saxony, “when the lambs are weaned, each in his turn is placed upon a table that his wool and form may be minutely observed. The finest are selected for breeding and receive a first mark. When they are one year old, and prior to shearing them, another close examination of those previously marked takes place: those in which no defect can be found receive a second mark, and the rest are condemned. A few months afterwards a third and last scrutiny is made; the prime rams and ewes receive a third and final mark, but the slightest blemish is sufficient to cause the rejection of the animal.” These sheep are bred and valued almost exclusively for the fineness of their wool; and the result corresponds with the labour bestowed on their selection. Instruments have been invented to measure accurately the thickness of the fibres; and “an Austrian fleece has been produced of which twelve hairs equalled in thickness one from a Leicester sheep.”

Throughout the world, wherever silk is produced, the greatest care is bestowed on selecting the cocoons from which the moths for breeding are to be reared. A careful cultivator[14] likewise examines the moths themselves, and destroys those that are not perfect. But what more immediately concerns us is that certain families in France devote themselves to raising eggs for sale.[15] In China, near Shanghai, the inhabitants of two small districts have the privilege of raising eggs for the whole surrounding country, and that they may give up their whole time to this business, they are interdicted by law from producing silk.[16]

The care which successful breeders take in matching their birds is surprising. Sir John Sebright, whose fame is perpetuated by the “Sebright Bantam,” used to spend “two and three days in examining, consulting, and disputing with a friend which were the best of five or six birds.”[17] Mr. Bult, whose pouter-pigeons won so many prizes, and were exported to North America under the charge of a man sent on purpose, told me that he always deliberated for several days before he matched each pair. Hence we can understand the advice of an eminent fancier, who writes[18] “I would here particularly guard you against having too great a variety of pigeons, otherwise you will know a little of all, but nothing about one as it ought to be known.” Apparently it transcends the power of the human intellect to breed all kinds: “it is possible that there may be a few fanciers that have a good general knowledge of fancy pigeons; but there are many more who labour under the delusion of supposing they know what they do not.” The excellence of one sub-variety, the Almond Tumbler, lies in the plumage, carriage, head, beak, and eye; but it is too presumptuous in the beginner to try for all these points. The great judge above quoted says, “There are some young fanciers who are over-covetous, who go for all the above five properties at once; they have their reward by getting nothing.” We thus see that breeding even fancy pigeons is no simple art: we may smile at the solemnity of these precepts, but he who laughs will win no prizes.

What methodical selection has effected for our animals is sufficiently proved, as already remarked, by our Exhibitions. So greatly were the sheep belonging to some of the earlier breeders, such as Bakewell and Lord Western, changed, that many persons could not be persuaded that they had not been crossed. Our pigs, as Mr. Corringham remarks[19] during the last twenty years have undergone, through rigorous selection together with crossing, a complete metamorphosis. The first exhibition for poultry was held in the Zoological Gardens in 1845; and the improvement effected since that time has been great. As Mr. Bailey, the great judge, remarked to me, it was formerly ordered that the comb of the Spanish cock should be upright, and in four or five years all good birds had upright combs; it was ordered that the Polish cock should have no comb or wattles, and now a bird thus furnished would be at once disqualified; beards were ordered, and out of fifty-seven pens lately (1860) exhibited at the Crystal Palace, all had beards. So it has been in many other cases. But in all cases the judges order only what is occasionally produced and what can be improved and rendered constant by selection. The steady increase in weight during the last few years in our fowls, turkeys, ducks, and geese is notorious; “six-pound ducks are now common, whereas four pounds was formerly the average.” As the time required to make a change has not often been recorded, it may be worth mentioning that it took Mr. Wicking thirteen years to put a clean white head on an almond tumbler’s body, “a triumph,” says another fancier, “of which he may be justly proud.”[20]

Mr. Tollet, of Betley Hall, selected cows, and especially bulls, descended from good milkers, for the sole purpose of improving his cattle for the production of cheese; he steadily tested the milk with the lactometer, and in eight years he increased, as I was informed by him, the product in proportion of four to three. Here is a curious case[21] of steady but slow progress, with the end not as yet fully attained: in 1784 a race of silkworms was introduced into France, in which one hundred in the thousand failed to produce white cocoons; but now after careful selection during sixty-five generations, the proportion of yellow cocoons has been reduced to thirty-five in the thousand.

With plants selection has been followed with the same good result as with animals. But the process is simpler, for plants in the great majority of cases bear both sexes. Nevertheless, with most kinds it is necessary to take as much care to prevent crosses as with animals or unisexual plants; but with some plants, such as peas, this care is not necessary. With all improved plants, excepting of course those which are propagated by buds, cuttings, etc., it is almost indispensable to examine the seedlings and destroy those which depart from the proper type. This is called “roguing,” and is, in fact, a form of selection, like the rejection of inferior animals. Experienced horticulturists and agriculturists incessantly urge every one to preserve the finest plants for the production of seed.

Although plants often present much more conspicuous variations than animals, yet the closest attention is generally requisite to detect each slight and favourable change. Mr. Masters relates[22] how “many a patient hour was devoted,” whilst he was young, to the detection of differences in peas intended for seed. Mr. Barnet[23] remarks that the old scarlet American strawberry was cultivated for more than a century without producing a single variety; and another writer observes how singular it was that when gardeners first began to attend to this fruit it began to vary; the truth no doubt being that it had always varied, but that, until slight variations were selected and propagated by seed, no conspicuous result was obtained. The finest shades of difference in wheat have been discriminated and selected with almost as much care as, in the case of the higher animals, for instance by Col. Le Couteur and more especially by Major Hallett.

It may be worth while to give a few examples of methodical selection with plants; but in fact the great improvement of all our anciently cultivated plants may be attributed to selection long carried on, in part methodically, and in part unconsciously. I have shown in a former chapter how the weight of the gooseberry has been increased by systematic selection and culture. The flowers of the Heartsease have been similarly increased in size and regularity of outline. With the Cineraria, Mr. Glenny[24] “was bold enough when the flowers were ragged and starry and ill defined in colour, to fix a standard which was then considered outrageously high and impossible, and which, even if reached, it was said, we should be no gainers by, as it would spoil the beauty of the flowers. He maintained that he was right; and the event has proved it to be so.” The doubling of flowers has several times been effected by careful selection: the Rev. W. Williamson,[25] after sowing during several years seed of Anemone coronaria, found a plant with one additional petal; he sowed the seed of this, and by perseverance in the same course obtained several varieties with six or seven rows of petals. The single Scotch rose was doubled, and yielded eight good varieties in nine or ten years.[26] The Canterbury bell (Campanula medium) was doubled by careful selection in four generations.[27] In four years Mr. Buckman,[28] by culture and careful selection, converted parsnips, raised from wild seed, into a new and good variety. By selection during a long course of years, the early maturity of peas has been hastened by between ten and twenty-one days.[29] A more curious case is offered by the beet plant, which since its cultivation in France, has almost exactly doubled its yield of sugar. This has been effected by the most careful selection; the specific gravity of the roots being regularly tested, and the best roots saved for the production of seed.[30]

Selection by Ancient and Semi-civilised People.

In attributing so much importance to the selection of animals and plants, it may be objected, that methodical selection would not have been carried on during ancient times. A distinguished naturalist considers it as absurd to suppose that semi-civilised people should have practised selection of any kind. Undoubtedly the principle has been systematically acknowledged and followed to a far greater extent within the last hundred years than at any former period, and a corresponding result has been gained; but it would be a greater error to suppose, as we shall immediately see, that its importance was not recognised and acted on during the most ancient times, and by semi-civilised people. I should premise that many facts now to be given only show that care was taken in breeding; but when this is the case, selection is almost sure to be practised to a certain extent. We shall hereafter be enabled better to judge how far selection, when only occasionally carried on, by a few of the inhabitants of a country, will slowly produce a great effect.

In a well-known passage in the thirtieth chapter of Genesis, rules are given for influencing, as was then thought possible, the colour of sheep; and speckled and dark breeds are spoken of as being kept separate. By the time of David the fleece was likened to snow. Youatt,[31] who has discussed all the passages in relation to breeding in the Old Testament, concludes that at this early period “some of the best principles of breeding must have been steadily and long pursued.” It was ordered, according to Moses, that “Thou shalt not let thy cattle gender with a diverse kind;” but mules were purchased[32] so that at this early period other nations must have crossed the horse and ass. It is said[33] that Erichthonius, some generations before the Trojan war, had many brood-mares, “which by his care and judgment in the choice of stallions produced a breed of horses superior to any in the surrounding countries.” Homer (Book 5) speaks of Aeneas’ horses as bred from mares which were put to the steeds of Laomedon. Plato, in his ‘Republic’ says to Glaucus, “I see that you raise at your house a great many dogs for the chase. Do you take care about breeding and pairing them? Among animals of good blood, are there not always some which are superior to the rest?” To which Glaucus answers in the affirmative.[34] Alexander the Great selected the finest Indian cattle to send to Macedonia to improve the breed.[35] According to Pliny,[36] King Pyrrhus had an especially valuable breed of oxen: and he did not suffer the bulls and cows to come together till four years old, that the breed might not degenerate. Virgil, in his Georgics (lib. 3), gives as strong advice as any modern agriculturist could do, carefully to select the breeding stock; “to note the tribe, the lineage, and the sire; whom to reserve for husband of the herd;”—to brand the progeny;—to select sheep of the purest white, and to examine if their tongues are swarthy. We have seen that the Romans kept pedigrees of their pigeons, and this would have been a senseless proceeding had not great care been taken in breeding them. Columella gives detailed instructions about breeding fowls: “Let the breeding hens therefore be of a choice colour, a robust body, square-built, full-breasted, with large heads, with upright and bright-red combs. Those are believed to be the best bred which have five toes.”[37] According to Tacitus, the Celts attended to the races of their domestic animals; and Caesar states that they paid high prices to merchants for fine imported horses.[38] In regard to plants, Virgil speaks of yearly culling the largest seeds; and Celsus says, “where the corn and crop is but small, we must pick out the best ears of corn, and of them lay up our seed separately by itself.”[39]

Coming down the stream of time, we may be brief. At about the beginning of the ninth century Charlemagne expressly ordered his officers to take great care of his stallions; and if any proved bad or old, to forewarn him in good time before they were put to the mares.[40] Even in a country so little civilised as Ireland during the ninth century, it would appear from some ancient verses,[41] describing a ransom demanded by Cormac, that animals from particular places, or having a particular character, were valued. Thus it is said,—

Two pigs of the pigs of Mac Lir, A ram and ewe both round and red, I brought with me from Aengus. I brought with me a stallion and a mare From the beautiful stud of Manannan, A bull and a white cow from Druim Cain.

Athelstan, in 930, received running-horses as a present from Germany; and he prohibited the exportation of English horses. King John imported “one hundred chosen stallions from Flanders.”[42] On June 16th, 1305, the Prince of Wales wrote to the Archbishop of Canterbury, begging for the loan of any choice stallion, and promising its return at the end of the season.[43] There are numerous records at ancient periods in English history of the importation of choice animals of various kinds, and of foolish laws against their exportation. In the reigns of Henry VII. and VIII. it was ordered that the magistrates, at Michaelmas, should scour the heaths and commons, and destroy all mares beneath a certain size.[44] Some of our earlier kings passed laws against the slaughtering rams of any good breed before they were seven years old, so that they might have time to breed. In Spain Cardinal Ximenes issued, in 1509, regulations on the selection of good rams for breeding.[45]

The Emperor Akbar Khan before the year l600 is said to have “wonderfully improved” his pigeons by crossing the breeds; and this necessarily implies careful selection. About the same period the Dutch attended with the greatest care to the breeding of these birds. Belon in 1555 says that good managers in France examined the colour of their goslings in order to get geese of a white colour and better kinds. Markham in 1631 tells the breeder “to elect the largest and goodliest conies,” and enters into minute details. Even with respect to seeds of plants for the flower-garden, Sir J. Hanmer writing about the year 1660[46] says, in “choosing seed, the best seed is the most weighty, and is had from the lustiest and most vigorous stems;” and he then gives rules about leaving only a few flowers on plants for seed; so that even such details were attended to in our flower-gardens two hundred years ago. In order to show that selection has been silently carried on in places where it would not have been expected, I may add that in the middle of the last century, in a remote part of North America, Mr. Cooper improved by careful selection all his vegetables, “so that they were greatly superior to those of any other person. When his radishes, for instance, are fit for use, he takes ten or twelve that he most approves, and plants them at least 100 yards from others that blossom at the same time. In the same manner he treats all his other plants, varying the circumstances according to their nature.”[47]

In the great work on China published in the last century by the Jesuits, and which is chiefly compiled from ancient Chinese encyclopaedias, it is said that with sheep “improving the breed consists in choosing with particular care the lambs which are destined for propagation, in nourishing them well, and in keeping the flocks separate.” The same principles were applied by the Chinese to various plants and fruit-trees.[48] An imperial edict recommends the choice of seed of remarkable size; and selection was practised even by imperial hands, for it is said that the Ya-mi, or imperial rice, was noticed at an ancient period in a field by the Emperor Khang-hi, was saved and cultivated in his garden, and has since become valuable from being the only kind which will grow north of the Great Wall.[49] Even with flowers, the tree paeony (P. moutan) has been cultivated, according to Chinese traditions, for 1400 years; between 200 and 300 varieties have been raised, which are cherished like tulips formerly were by the Dutch.[50]

Turning now to semi-civilised people and to savages: it occurred to me, from what I had seen of several parts of South America, where fences do not exist, and where the animals are of little value, that there would be absolutely no care in breeding or selecting them; and this to a large extent is true. Roulin,[51] however, describes in Columbia a naked race of cattle, which are not allowed to increase, on account of their delicate constitution. According to Azara[52] horses are often born in Paraguay with curly hair; but, as the natives do not like them, they are destroyed. On the other hand, Azara states that a hornless bull, born in 1770, was preserved and propagated its race. I was informed of the existence in Banda Oriental of a breed with reversed hair; and the extraordinary niata cattle first appeared and have since been kept distinct in La Plata. Hence certain conspicuous variations have been preserved, and others have been habitually destroyed, in these countries, which are so little favourable for careful selection. We have also seen that the inhabitants sometimes introduce fresh cattle on their estates to prevent the evil effects of close interbreeding. On the other hand, I have heard on reliable authority that the Gauchos of the Pampas never take any pains in selecting the best bulls or stallions for breeding; and this probably accounts for the cattle and horses being remarkably uniform in character throughout the immense range of the Argentine republic.

Looking to the Old World, in the Sahara Desert “The Touareg is as careful in the selection of his breeding Mahari (a fine race of the dromedary) as the Arab is in that of his horse. The pedigrees are handed down, and many a dromedary can boast a genealogy far longer than the descendants of the Darley Arabian.”[53] According to Pallas the Mongolians endeavour to breed the Yaks or horse-tailed buffaloes with white tails, for these are sold to the Chinese mandarins as fly-flappers; and Moorcroft, about seventy years after Pallas, found that white-tailed animals were still selected for breeding.[54]

We have seen in the chapter on the Dog that savages in different parts of North America and in Guiana cross their dogs with wild Canidæ, as did the ancient Gauls, according to Pliny. This was done to give their dogs strength and vigour, in the same way as the keepers in large warrens now sometimes cross their ferrets (as I have been informed by Mr. Yarrell) with the wild polecat, “to give them more devil.” According to Varro, the wild ass was formerly caught and crossed with the tame animal to improve the breed, in the same manner as at the present day the natives of Java sometimes drive their cattle into the forests to cross with the wild Banteng (Bos sondaicus).[55] In Northern Siberia, among the Ostyaks, the dogs vary in markings in different districts, but in each place they are spotted black and white in a remarkably uniform manner;[56] and from this fact alone we may infer careful breeding, more especially as the dogs of one locality are famed throughout the country for their superiority. I have heard of certain tribes of Esquimaux who take pride in their teams of dogs being uniformly coloured. In Guiana, as Sir H. Schomburgk informs me,[57] the dogs of the Turuma Indians are highly valued and extensively bartered: the price of a good one is the same as that given for a wife: they are kept in a sort of cage, and the Indians “take great care when the female is in season to prevent her uniting with a dog of an inferior description.” The Indians told Sir Robert that, if a dog proved bad or useless, he was not killed, but was left to die from sheer neglect. Hardly any nation is more barbarous than the Fuegians, but I hear from Mr. Bridges, the Catechist to the Mission, that, “when these savages have a large, strong, and active bitch, they take care to put her to a fine dog, and even take care to feed her well, that her young may be strong and well favoured.”

In the interior of Africa, negroes, who have not associated with white men, show great anxiety to improve their animals; they “always choose the larger and stronger males for stock;” the Malakolo were much pleased at Livingstone’s promise to send them a bull, and some Bakalolo carried a live cock all the way from Loanda into the interior.[58] At Falaba Mr. Winwood Reade noticed an unusually fine horse, and the negro King informed him that “the owner was noted for his skill in breeding horses.” Further south on the same continent, Andersson states that he has known a Damara give two fine oxen for a dog which struck his fancy. The Damaras take great delight in having whole droves of cattle of the same colour, and they prize their oxen in proportion to the size of their horns. “The Namaquas have a perfect mania for a uniform team; and almost all the people of Southern Africa value their cattle next to their women, and take a pride in possessing animals that look high-bred. They rarely or never make use of a handsome animal as a beast of burden.”[59] The power of discrimination which these savages possess is wonderful, and they can recognise to which tribe any cattle belong. Mr. Andersson further informs me that the natives frequently match a particular bull with a particular cow.

The most curious case of selection by semi-civilised people, or indeed by any people, which I have found recorded, is that given by Garcilazo de la Vega, a descendant of the Incas, as having been practised in Peru before the country was subjugated by the Spaniards.[60] The Incas annually held great hunts, when all the wild animals were driven from an immense circuit to a central point. The beasts of prey were first destroyed as injurious. The wild Guanacos and Vicunas were sheared; the old males and females killed, and the others set at liberty. The various kinds of deer were examined; the old males and females were likewise killed, “but the young females, with a certain number of males, selected from the most beautiful and strong,” were given their freedom. Here, then, we have selection by man aiding natural selection. So that the Incas followed exactly the reverse system of that which our Scottish sportsman are accused of following, namely, of steadily killing the finest stags, thus causing the whole race to degenerate.[61] In regard to the domesticated llamas and alpacas, they were separated in the time of the Incas according to colour: and if by chance one in a flock was born of the wrong colour, it was eventually put into another flock.

In the genus Auchenia there are four forms,—the Guanaco and Vicuna, found wild and undoubtedly distinct species; the Llama and Alpaca, known only in a domesticated condition. These four animals appear so different, that most naturalists, especially those who have studied these animals in their native country, maintain that they are specifically distinct, notwithstanding that no one pretends to have seen a wild llama or alpaca. Mr. Ledger, however, who has closely studied these animals both in Peru and during their exportation to Australia, and who has made many experiments on their propagation, adduces arguments[62] which seem to me conclusive, that the llama is the domesticated descendant of the guanaco, and the alpaca of the vicuna. And now that we know that these animals were systematically bred and selected many centuries ago, there is nothing surprising in the great amount of change which they have undergone.

It appeared to me at one time probable that, though ancient and semi-civilised people might have attended to the improvement of their more useful animals in essential points, yet that they would have disregarded unimportant characters. But human nature is the same throughout the world: fashion everywhere reigns supreme, and man is apt to value whatever he may chance to possess. We have seen that in South America the niata cattle, which certainly are not made useful by their shortened faces and upturned nostrils, have been preserved. The Damaras of South Africa value their cattle for uniformity of colour and enormously long horns. And I will now show that there is hardly any peculiarity in our most useful animals which, from fashion, superstition, or some other motive, has not been valued, and consequently preserved. With respect to cattle, “an early record,” according to Youatt[63] “speaks of a hundred white cows with red ears being demanded as a compensation by the princes of North and South Wales. If the cattle were of a dark or black colour, 150 were to be presented.” So that colour was attended to in Wales before its subjugation by England. In Central Africa, an ox that beats the ground with its tail is killed; and in South Africa some of the Damaras will not eat the flesh of a spotted ox. The Kaffirs value an animal with a musical voice; and “at a sale in British Kaffraria the low of a heifer excited so much admiration that a sharp competition sprung up for her possession, and she realised a considerable price.”[64] With respect to sheep, the Chinese prefer rams without horns; the Tartars prefer them with spirally wound horns, because the hornless are thought to lose courage.[65] Some of the Damaras will not eat the flesh of hornless sheep. In regard to horses, at the end of the fifteenth century animals of the colour described as liart pomme were most valued in France. The Arabs have a proverb, “Never buy a horse with four white feet, for he carries his shroud with him”;[66] the Arabs also, as we have seen, despise dun-coloured horses. So with dogs, Xenophon and others at an ancient period were prejudiced in favour of certain colours; and “white or slate-coloured hunting dogs were not esteemed.”[67]

Turning to poultry, the old Roman gourmands thought that the liver of a white goose was the most savoury. In Paraguay black-skinned fowls are kept because they are thought to be more productive, and their flesh the most proper for invalids.[68] In Guiana, as I am informed by Sir R. Schomburgk, the aborigines will not eat the flesh or eggs of the fowl, but two races are kept distinct merely for ornament. In the Philippines, no less than nine sub-varieties of the game-cock are kept and named, so that they must be separately bred.

At the present time in Europe, the smallest peculiarities are carefully attended to in our most useful animals, either from fashion, or as a mark of purity of blood. Many examples could be given; two will suffice. “In the Western counties of England the prejudice against a white pig is nearly as strong as against a black one in Yorkshire.” In one of the Berkshire sub-breeds, it is said, “the white should be confined to four white feet, a white spot between the eyes, and a few white hairs behind each shoulder.” Mr. Saddler possessed “three hundred pigs, every one of which was marked in this manner.”[69] Marshall, towards the close of the last century, in speaking of a change in one of the Yorkshire breeds of cattle, says the horns have been considerably modified, as “a clean, small, sharp horn has been fashionable for the last twenty years.”[70] In a part of Germany the cattle of the Race de Gfoehl are valued for many good qualities, but they must have horns of a particular curvature and tint, so much so that mechanical means are applied if they take a wrong direction; but the inhabitants “consider it of the highest importance that the nostrils of the bull should be flesh-coloured, and the eyelashes light; this is an indispensable condition. A calf with blue nostrils would not be purchased, or purchased at a very low price.”[71] Therefore let no man say that any point or character is too trifling to be methodically attended to and selected by breeders.

Unconscious Selection.—By this term I mean, as already more than once explained, the preservation by man of the most valued, and the destruction of the least valued individuals, without any conscious intention on his part of altering the breed. It is difficult to offer direct proofs of the results which follow from this kind of selection; but the indirect evidence is abundant. In fact, except that in the one case man acts intentionally, and in the other unintentionally, there is little difference between methodical and unconscious selection. In both cases man preserves the animals which are most useful or pleasing to him, and destroys or neglects the others. But no doubt a far more rapid result follows from methodical than from unconscious selection. The “roguing” of plants by gardeners, and the destruction by law in Henry VIII.’s reign of all under-sized mares, are instances of a process the reverse of selection in the ordinary sense of the word, but leading to the same general result. The influence of the destruction of individuals having a particular character is well shown by the necessity of killing every lamb with a trace of black about it, in order to keep the flock white; or again, by the effects on the average height of the men of France of the destructive wars of Napoleon, by which many tall men were killed, the short ones being left to be the fathers of families. This at least is the conclusion of some of those who have closely studied the effects of the conscription; and it is certain that since Napoleon’s time the standard for the army has been lowered two or three times.

Unconscious selection blends with methodical, so that it is scarcely possible to separate them. When a fancier long ago first happened to notice a pigeon with an unusually short beak, or one with the tail-feathers unusually developed, although he bred from these birds with the distinct intention of propagating the variety, yet he could not have intended to make a short-faced tumbler or a fantail, and was far from knowing that he had made the first step towards this end. If he could have seen the final result, he would have been struck with astonishment, but, from what we know of the habits of fanciers, probably not with admiration. Our English carriers, barbs, and short-faced tumblers have been greatly modified in the same manner, as we may infer both from the historical evidence given in the chapters on the Pigeon, and from the comparison of birds brought from distant countries.

So it has been with dogs; our present fox-hounds differ from the old English hound; our greyhounds have become lighter: the Scotch deer-hound has been modified, and is now rare. Our bulldogs differ from those which were formerly used for baiting bulls. Our pointers and Newfoundlands do not closely resemble any native dog now found in the countries whence they were brought. These changes have been effected partly by crosses; but in every case the result has been governed by the strictest selection. Nevertheless, there is no reason to suppose that man intentionally and methodically made the breeds exactly what they now are. As our horses became fleeter, and the country more cultivated and smoother, fleeter fox-hounds were desired and produced, but probably without any one distinctly foreseeing what they would become. Our pointers and setters, the latter almost certainly descended from large spaniels, have been greatly modified in accordance with fashion and the desire for increased speed. Wolves have become extinct, and so has the wolf-dog; deer have become rarer, bulls are no longer baited, and the corresponding breeds of the dog have answered to the change. But we may feel almost sure that when, for instance, bulls were no longer baited, no man said to himself, I will now breed my dogs of smaller size, and thus create the present race. As circumstances changed, men unconsciously and slowly modified their course of selection.

With racehorses selection for swiftness has been followed methodically, and our horses now easily surpass their progenitors. The increased size and different appearance of the English racehorse led a good observer in India to ask, “Could any one in this year of 1856, looking at our racehorses, conceive that they were the result of the union of the Arab horse and the African mare?”[72] This change has, it is probable, been largely effected through unconscious selection, that is, by the general wish to breed as fine horses as possible in each generation, combined with training and high feeding, but without any intention to give to them their present appearance. According to Youatt,[73] the introduction in Oliver Cromwell’s time of three celebrated Eastern stallions speedily affected the English breed; “so that Lord Harleigh, one of the old school, complained that the great horse was fast disappearing.” This is an excellent proof how carefully selection must have been attended to; for without such care, all traces of so small an infusion of Eastern blood would soon have been absorbed and lost. Notwithstanding that the climate of England has never been esteemed particularly favourable to the horse, yet long-continued selection, both methodical and unconscious, together with that practised by the Arabs during a still longer and earlier period, has ended in giving us the best breed of horses in the world. Macaulay[74] remarks, “Two men whose authority on such subjects was held in great esteem, the Duke of Newcastle and Sir John Fenwick, pronounced that the meanest hack ever imported from Tangier would produce a finer progeny than could be expected from the best sire of our native breed. They would not readily have believed that a time would come when the princes and nobles of neighbouring lands would be as eager to obtain horses from England as ever the English had been to obtain horses from Barbary.”

The London dray-horse, which differs so much in appearance from any natural species, and which from its size has so astonished many Eastern princes, was probably formed by the heaviest and most powerful animals having been selected during many generations in Flanders and England, but without the least intention or expectation of creating a horse such as we now see. If we go back to an early period of history, we behold in the antique Greek statues, as Schaaffhausen has remarked,[75] a horse equally unlike a race or dray horse, and differing from any existing breed.

The results of unconscious selection, in an early stage, are well shown in the difference between the flocks descended from the same stock, but separately reared by careful breeders. Youatt gives an excellent instance of this fact in the sheep belonging to Messrs. Buckley and Burgess, which “have been purely bred from the original stock of Mr. Bakewell for upwards of fifty years. There is not a suspicion existing in the mind of any one at all acquainted with the subject that the owner of either flock has deviated in any one instance from the pure blood of Mr. Bakewell’s flock; yet the difference between the sheep possessed by these two gentlemen is so great, that they have the appearance of being quite different varieties.”[76] I have seen several analogous and well marked cases with pigeons: for instance, I had a family of barbs descended from those long bred by Sir J. Sebright, and another family long bred by another fancier, and the two families plainly differed from each other. Nathusius—and a more competent witness could not be cited—observes that, though the Shorthorns are remarkably uniform in appearance (except in colour), yet the individual character and wishes of each breeder become impressed on his cattle, so that different herds differ slightly from one another.[77] The Hereford cattle assumed their present well-marked character soon after the year 1769, through careful selection by Mr. Tomkins[78] and the breed has lately split into two strains—one strain having a white face, and differing slightly, it is said,[79] in some other points: but there is no reason to believe that this split, the origin of which is unknown, was intentionally made; it may with much more probability be attributed to different breeders having attended to different points. So again, the Berkshire breed of swine in the year 1810 had greatly changed from what it was in 1780; and since 1810 at least two distinct sub-breeds have arisen bearing the same name.[80] Keeping in mind how rapidly all animals increase, and that some must be annually slaughtered and some saved for breeding, then, if the same breeder during a long course of years deliberately settles which shall be saved and which shall be killed, it is almost inevitable that his individual turn of mind will influence the character of his stock, without his having had any intention to modify the breed.

Unconscious selection in the strictest sense of the word, that is, the saving of the more useful animals and the neglect or slaughter of the less useful, without any thought of the future, must have gone on occasionally from the remotest period and amongst the most barbarous nations. Savages often suffer from famines, and are sometimes expelled by war from their own homes. In such cases it can hardly be doubted that they would save their most useful animals. When the Fuegians are hard pressed by want, they kill their old women for food rather than their dogs; for, as we were assured, “old women no use—dogs catch otters.” The same sound sense would surely lead them to preserve their more useful dogs when still harder pressed by famine. Mr. Oldfield, who has seen so much of the aborigines of Australia, informs me that “they are all very glad to get a European kangaroo dog, and several instances have been known of the father killing his own infant that the mother might suckle the much-prized puppy.” Different kinds of dogs would be useful to the Australian for hunting opossums and kangaroos, and to the Fuegian for catching fish and otters; and the occasional preservation in the two countries of the most useful animals would ultimately lead to the formation of two widely distinct breeds.

With plants, from the earliest dawn of civilisation, the best variety which was known would generally have been cultivated at each period and its seeds occasionally sown; so that there will have been some selection from an extremely remote period, but without any prefixed standard of excellence or thought of the future. We at the present day profit by a course of selection occasionally and unconsciously carried on during thousands of years. This is proved in an interesting manner by Oswald Heer’s researches on the lake-inhabitants of Switzerland, as given in a former chapter; for he shows that the grain and seed of our present varieties of wheat, barley, oats, peas, beans, lentils, and poppy, exceed in size those which were cultivated in Switzerland during the Neolithic and Bronze periods. These ancient people, during the Neolithic period, possessed also a crab considerably larger than that now growing wild on the Jura.[81] The pears described by Pliny were evidently extremely inferior in quality to our present pears. We can realise the effects of long-continued selection and cultivation in another way, for would any one in his senses expect to raise a first-rate apple from the seed of a truly wild crab, or a luscious melting pear from the wild pear? Alphonse de Candolle informs me that he has lately seen on an ancient mosaic at Rome a representation of the melon; and as the Rotnans, who were such gourmands, are silent on this fruit, he infers that the melon has been greatly ameliorated since the classical period.

Coming to later times, Buffon[82] on comparing the flowers, fruit, and vegetables which were then cultivated with some excellent drawings made a hundred and fifty years previously, was struck with surprise at the great improvement which had been effected; and remarks that these ancient flowers and vegetables would now be rejected, not only by a florist but by a village gardener. Since the time of Buffon the work of improvement has steadily and rapidly gone on. Every florist who compares our present flowers with those figured in books published not long since, is astonished at the change. A well-known amateur,[83] in speaking of the varieties of Pelargonium raised by Mr. Garth only twenty-two years before, remarks, “What a rage they excited: surely we had attained perfection, it was said; and now not one of the flowers of those days will be looked at. But none the less is the debt of gratitude which we owe to those who saw what was to be done, and did it.” Mr. Paul, the well-known horticulturist, in writing of the same flower,[84] says he remembers when young being delighted with the portraits in Sweet’s work; “but what are they in point of beauty compared with the Pelargoniums of this day? Here again nature did not advance by leaps; the improvement was gradual, and if we had neglected those very gradual advances, we must have foregone the present grand results.” How well this practical horticulturist appreciates and illustrates the gradual and accumulative force of selection! The Dahlia has advanced in beauty in a like manner; the line of improvement being guided by fashion, and by the successive modifications which the flower slowly underwent.[85] A steady and gradual change has been noticed in many other flowers: thus an old florist,[86] after describing the leading varieties of the Pink which were grown in 1813 adds, “the pinks of those days would now be scarcely grown as border-flowers.” The improvement of so many flowers and the number of the varieties which have been raised is all the more striking when we hear that the earliest known flower-garden in Europe, namely at Padua, dates only from the year 1545.[87]

Effects of Selection, as shown by the parts most valued by man presenting the greatest amount of difference.—The power of long-continued selection, whether methodical or unconscious, or both combined, is well shown in a general way, namely, by the comparison of the differences between the varieties of distinct species, which are valued for different parts, such as for the leaves, or stems, or tubers, the seed, or fruit, or flowers. Whatever part man values most, that part will be found to present the greatest amount of difference. With trees cultivated for their fruit, Sageret remarks that the fruit is larger than in the parent-species, whilst with those cultivated for the seed, as with nuts, walnuts, almonds, chestnuts, etc., it is the seed itself which is larger; and he accounts for this fact by the fruit in the one case, and by the seed in the other, having been carefully attended to and selected during many ages. Gallesio has made the same observation. Godron insists on the diversity of the tuber in the potato, of the bulb in the onion, and of the fruit in the melon; and on the close similarity of the other parts in these same plants.[88]

In order to judge how far my own impression on this subject was correct, I cultivated numerous varieties of the same species close to one another. The comparison of the amount of difference between widely different organs is necessarily vague; I will therefore give the results in only a few cases. We have previously seen in the ninth chapter how greatly the varieties of the cabbage differ in their foliage and stems, which are the selected parts, and how closely they resemble one another in their flowers, capsules, and seeds. In seven varieties of the radish, the roots differed greatly in colour and shape, but no difference whatever could be detected in their foliage, flowers, or seeds. Now what a contrast is presented, if we compare the flowers of the varieties of these two plants with those of any species cultivated in our flower-gardens for ornament; or if we compare their seeds with those of the varieties of maize, peas, beans, etc., which are valued and cultivated for their seeds. In the ninth chapter it was shown that the varieties of the pea differ but little except in the tallness of the plant, moderately in the shape of the pod, and greatly in the pea itself, and these are all selected points. The varieties, however, of the Pois sans parchemin differ much more in their pods, and these are eaten and valued. I cultivated twelve varieties of the common bean; one alone, the Dwarf Fan, differed considerably in general appearance; two differed in the colour of their flowers, one being an albino, and the other being wholly instead of partially purple; several differed considerably in the shape and size of the pod, but far more in the bean itself, and this is the valued and selected part. Toker’s bean, for instance, is twice-and-a-half as long and broad as the horse-bean, and is much thinner and of a different shape.

The varieties of the gooseberry, as formerly described, differ much in their fruit, but hardly perceptibly in their flowers or organs of vegetation. With the plum, the differences likewise appear to be greater in the fruit than in the flowers or leaves. On the other hand, the seed of the strawberry, which corresponds with the fruit of the plum, differs hardly at all; whilst every one knows how greatly the fruit—that is, the enlarged receptacle—differs in several varieties. In apples, pears, and peaches the flowers and leaves differ considerably, but not, as far as I can judge, in proportion with the fruit. The Chinese double-flowering peaches, on the other hand, show that varieties of this tree have been formed, which differ more in flower than in fruit. If, as is highly probable, the peach is the modified descent of the almond, a surprising amount of change has been effected in the same species, in the fleshy covering of the former and in the kernels of the latter.

When parts stand in close relationship to each other, such as the seed and the fleshy covering of the fruit (whatever its homological nature may be), changes in the one are usually accompanied by modifications in the other, though not necessarily to the same degree. With the plum-tree, for instance, some varieties produce plums which are nearly alike, but include stones extremely dissimilar in shape; whilst conversely other varieties produce dissimilar fruit with barely distinguishable stones; and generally the stones, though they have never been subjected to selection, differ greatly in the several varieties of the plum. In other cases organs which are not manifestly related, through some unknown bond vary together, and are consequently liable, without any intention on man’s part, to be simultaneously acted on by selection. Thus the varieties of the stock (Matthiola) have been selected solely for the beauty of their flowers, but the seeds differ greatly in colour and somewhat in size. Varieties of the lettuce have been selected solely on account of their leaves, yet produce seeds which likewise differ in colour. Generally, through the law of correlation, when a variety differs greatly from its fellow-varieties in any one character, it differs to a certain extent in several other characters. I observed this fact when I cultivated together many varieties of the same species, for I used first to make a list of the varieties which differed most from each other in their foliage and manner of growth, afterwards of those that differed most in their flowers, then in their seed-capsules, and lastly in their mature seed; and I found that the same names generally occurred in two, three, or four of the successive lists. Nevertheless the greatest amount of difference between the varieties was always exhibited, as far as I could judge, by that part or organ for which the plant was cultivated.

When we bear in mind that each plant was at first cultivated because useful to man, and that its variation was a subsequent, often a long subsequent, event, we cannot explain the greater amount of diversity in the valuable parts by supposing that species endowed with an especial tendency to vary in any particular manner were originally chosen. We must attribute the result to the variations in these parts having been successively preserved, and thus continually augmented; whilst other variations, excepting such as inevitably appeared through correlation, were neglected and lost. We may therefore infer that most plants might be made, through long-continued selection, to yield races as different from one another in any character as they now are in those parts for which they are valued and cultivated.

With animals we see nothing of the same kind; but a sufficient number of species have not been domesticated for a fair comparison. Sheep are valued for their wool, and the wool differs much more in the several races than the hair in cattle. Neither sheep, goats, European cattle, nor pigs are valued for their fleetness or strength; and we do not possess breeds differing in these respects like the racehorse and dray-horse. But fleetness and strength are valued in camels and dogs; and we have with the former the swift dromedary and heavy camel; with the latter the greyhound and mastiff. But dogs are valued even in a higher degree for their mental qualities and senses; and every one knows how greatly the races differ in these respects. On the other hand, where the dog is kept solely to serve for food, as in the Polynesian islands and China, it is described as an extremely stupid animal.[89] Blumenbach remarks that “many dogs, such as the badger-dog, have a build so marked and so appropriate for particular purposes, that I should find it very difficult to persuade myself that this astonishing figure was an accidental consequence of degeneration.”[90] Had Blumenbach reflected on the great principle of selection, he would not have used the term degeneration, and he would not have been astonished that dogs and other animals should become excellently adapted for the service of man.

On the whole we may conclude that whatever part or character is most valued—whether the leaves, stems, tubers, bulbs, flowers, fruit, or seed of plants, or the size, strength, fleetness, hairy covering, or intellect of animals—that character will almost invariably be found to present the greatest amount of difference both in kind and degree. And this result may be safely attributed to man having preserved during a long course of generations the variations which were useful to him, and neglected the others.

I will conclude this chapter by some remarks on an important subject. With animals such as the giraffe, of which the whole structure is admirably co-ordinated for certain purposes, it has been supposed that all the parts must have been simultaneously modified; and it has been argued that, on the principle of natural selection, this is scarcely possible. But in thus arguing, it has been tacitly assumed that the variations must have been abrupt and great. No doubt, if the neck of a ruminant were suddenly to become greatly elongated, the fore limbs and back would have to be simultaneously strengthened and modified; but it cannot be denied that an animal might have its neck, or head, or tongue, or fore-limbs elongated a very little without any corresponding modification in other parts of the body; and animals thus slightly modified would, during a dearth, have a slight advantage, and be enabled to browse on higher twigs, and thus survive. A few mouthfuls more or less every day would make all the difference between life and death. By the repetition of the same process, and by the occasional intercrossing of the survivors, there would be some progress, slow and fluctuating though it would be, towards the admirably coordinated structure of the giraffe. If the short-faced tumbler-pigeon, with its small conical beak, globular head, rounded body, short wings, and small feet—characters which appear all in harmony—had been a natural species, its whole structure would have been viewed as well fitted for its life; but in this case we know that inexperienced breeders are urged to attend to point after point, and not to attempt improving the whole structure at the same time. Look at the greyhound, that perfect image of grace, symmetry, and vigour; no natural species can boast of a more admirably co-ordinated structure, with its tapering head, slim body, deep chest, tucked-up abdomen, rat-like tail, and long muscular limbs, all adapted for extreme fleetness, and for running down weak prey. Now, from what we see of the variability of animals, and from what we know of the method which different men follow in improving their stock—some chiefly attending to one point, others to another point, others again correcting defects by crosses, and so forth—we may feel assured that if we could see the long line of ancestors of a first-rate greyhound up to its wild wolf-like progenitor, we should behold an infinite number of the finest gradations, sometimes in one character and sometimes in another, but all leading towards our present perfect type. By small and doubtful steps such as these, nature, as we may confidently believe, has progressed, on her grand march of improvement and development.

A similar line of reasoning is as applicable to separate organs as to the whole organisation. A writer[91] has recently maintained that “it is probably no exaggeration to suppose that in order to improve such an organ as the eye at all, it must be improved in ten different ways at once. And the improbability of any complex organ being produced and brought to perfection in any such way is an improbability of the same kind and degree as that of producing a poem or a mathematical demonstration by throwing letters at random on a table.” If the eye were abruptly and greatly modified, no doubt many parts would have to be simultaneously altered, in order that the organ should remain serviceable.

But is this the case with smaller changes? There are persons who can see distinctly only in a dull light, and this condition depends, I believe, on the abnormal sensitiveness of the retina, and is known to be inherited. Now if a bird, for instance, receive some great advantage from seeing well in the twilight, all the individuals with the most sensitive retina would succeed best and be the most likely to survive; and why should not all those which happened to have the eye itself a little larger, or the pupil capable of greater dilatation, be likewise preserved, whether or not these modifications were strictly simultaneous? These individuals would subsequently intercross and blend their respective advantages. By such slight successive changes, the eye of a diurnal bird would be brought into the condition of that of an owl, which has often been advanced as an excellent instance of adaptation. Short-sight, which is often inherited, permits a person to see distinctly a minute object at so near a distance that it would be indistinct to ordinary eyes; and here we have a capacity which might be serviceable under certain conditions, abruptly gained. The Fuegians on board the Beagle could certainly see distant objects more distinctly than our sailors with all their long practice; I do not know whether this depends upon sensitiveness or on the power of adjustment in the focus; but this capacity for distant vision might, it is probable, be slightly augmented by successive modifications of either kind. Amphibious animals which are enabled to see both in the water and in the air, require and possess, as M. Plateau has shown,[92] eyes constructed on the following plan: “the cornea is always flat, or at least much flattened in the front of the crystalline and over a space equal to the diameter of that lens, whilst the lateral portions may be much curved.” The crystalline is very nearly a sphere, and the humours have nearly the same density as water. Now as a terrestrial animal became more and more aquatic in its habits, very slight changes, first in the curvature of the cornea or crystalline, and then in the density of the humours, or conversely, might successively occur, and would be advantageous to the animal whilst under water, without serious detriment to its power of vision in the air. It is of course impossible to conjecture by what steps the fundamental structure of the eye in the Vertebrata was originally acquired, for we know nothing about this organ in the first progenitors of the class. With respect to the lowest animals in the scale, the transitional states through which the eye at first probably passed, can by the aid of analogy be indicated, as I have attempted to show in my ‘Origin of Species.’[93]

REFERENCES

[1] The term unconscious selection has been objected to as a contradiction; but see some excellent observations on this head by Prof. Huxley (‘Nat. Hist. Review,’ Oct. 1864, p. 578), who remarks that when the wind heaps up sand-dunes it sifts and unconsciously selects from the gravel on the beach grains of sand of equal size.

[2] ‘On Sheep,’ 1838, p. 60.

[3] Mr. J. Wright on Shorthorn Cattle, in ‘Journal of Royal Agricult. Soc.,’ vol. vii. pp. 208, 209.

[4] H. D. Richardson ‘On Pigs,’ 1847, p. 44.

[5] ‘Journal of Royal Agricult. Soc.,’ vol. i. p. 24.

[6] ‘On Sheep,’ pp. 520, 319.

[7] Loudon’s ‘Mag. of Nat. Hist.,’ vol. viii., 1835, p. 618.

[8] ‘A treatise on the Art of Breeding the Almond Tumbler.’ 1851. p. 9.

[9] ‘Recreations in Agriculture,’ vol. ii. p. 409.

[10] Youatt on Cattle, pp. 191, 227.

[11] Ferguson, ‘Prize Poultry,’ 1854, p. 208.

[12] Wilson, in ‘Transact. Highland Agricult. Soc.,’ quoted in ‘Gardener’s Chronicle,’ 1844, p. 29.

[13] Simmonds, quoted in ‘Gardener’s Chronicle,’ 1855, p. 637. And for the second quotation, see Youatt on Sheep, p. 171.

[14] Robinet, ‘Vers à Soie,’ 1848, p. 271.

[15] Quatrefages, ‘Les Maladies du Ver à Soie,’ 1859, p. 101.

[16] M. Simon, in ‘Bull. de la Soc. d’Acclimat.,’ tom. ix., 1862, p. 221.

[17] ‘The Poultry Chronicle,’ vol. i., 1854, p. 607.

[18] J. M. Eaton, ‘A Treatise on Fancy Pigeons,’ 1852, p. xiv., and ‘A Treatise on the Almond Tumbler,’ 1851, p. 11.

[19] ‘Journal Royal Agricultural Soc.,’ vol. vi., p. 22.

[20] ‘Poultry Chronicle,’ vol. ii., 1855, p. 596.

[21] Isid. Geoffroy St.-Hilaire, ‘Hist. Nat. Gén.,’ tom. iii. p. 254.

[22] ‘Gardener’s Chronicle,’ 1850, p. 198.

[23] ‘Transact. Hort. Soc.,’ vol. vi. p. 152.

[24] ‘Journal of Horticulture,’ 1862, p. 369.

[25] ‘Transact. Hort. Soc.,’ vol. iv. p. 381.

[26] ‘Transact. Hort. Soc.,’ vol. iv. p. 285.

[27] Rev. W. Bromehead, in ‘Gardener’s Chronicle,’ 1857, p. 550.

[28] ‘Gardener’s Chronicle,’ 1862, p. 721.

29[] Dr. Anderson, in ‘The Bee,’ vol. vi. p. 96; Mr. Barnes in ‘Gardener’s Chronicle,’ 1844, p. 476.

[30] Godron, ‘De l’Espèce,’ 1859, tom. ii. p. 69; ‘Gardener’s Chronicle,’ 1854, p. 258.Return to passage

[31] On Sheep, p. 18.Return to passage

[32] Volz, ‘Beiträge zur Kulturgeschichte,’ 1852, s. 47.Return to passage

[33] Mitford’s ‘History of Greece,’ vol. i. p. 73.Return to passage

[34] Dr. Dally, translated in ‘Anthropological Review,’ May 1864, p. 101.Return to passage

[35] Volz, ‘Beiträge,’ etc., 1852, s. 80.Return to passage

[36] ‘History of the World,’ ch. 45.Return to passage

[37] ‘Gardener’s Chronicle,’ 1848, p. 323.Return to passage

[38] Reynier, ‘De l’Économie des Celtes,’ 1818, pp. 487, 503.Return to passage

[39] Le Couteur on Wheat, p. 15.Return to passage

[40] Michel, ‘Des Haras,’ 1861, p. 84.Return to passage

[41] Sir W. Wilde, an ‘Essay on Unmanufactured Animal Remains,’ etc., 1860, p. 11.Return to passage

[42] Col. Hamilton Smith, ‘Nat. Library,’ vol. xii., Horses, pp. 135, 140.Return to passage

[43] Michel, ‘Des Haras,’ p. 90.Return to passage

[44] Mr. Baker, ‘History of the Horse,’ ‘Veterinary,’ vol. xiii. p. 423.Return to passage

[45] M. l’Abbé Carlier, in ‘Journal de Physique,’ vol. xxiv., 1784, p. 181; this memoir contains much information on the ancient selection of sheep; and is my authority for rams not being killed young in England.Return to passage

[46] ‘Gardener’s Chronicle,’ 1843, p. 389.Return to passage

[47] ‘Communications to Board of Agriculture’ quoted in Dr. Darwin’s ‘Phytologia,’ 1800, p. 451.Return to passage

[48] ‘Mémoire sur les Chinois,’ 1786, tom. xi. p. 55; tom. v. p. 507.Return to passage

[49] ‘Recherches sur l’Agriculture des Chinois,’ par L. D’Hervey Saint-Denys, 1850, p. 229. With respect to Khang-hi see Huc’s ‘Chinese Empire,’ p. 311.Return to passage

[50] Anderson, in ‘Linn. Transact.,’ vol. xii. p. 253.Return to passage

[51] ‘Mém. de l’Acad.’ (divers savants), tom. vi., 1835, p. 333.Return to passage

[52] ‘Des Quadrupèdes du Paraguay,’ 1801, tom. ii. pp. 333, 371.Return to passage

[53] ‘The Great Sahara,’ by the Rev. H. B. Tristram, 1860, p. 238.Return to passage

[54] Pallas, ‘Act. Acad. St. Petersburg,’ 1777, p. 249; Moorcroft and Trebeck, ‘Travels in the Himalayan Provinces,’ 1841.Return to passage

[55] Quoted from Raffles, in the ‘Indian Field,’ 1859, p. 196: for Varro, see Pallas ut supra.Return to passage

[56] Erman’s ‘Travels in Siberia,’ Eng. translat., vol. i. p. 453.Return to passage

[57] See also ‘Journal of R. Geograph. Soc.,’ vol. xiii. part i. p. 65.Return to passage

[58] Livingstone’s ‘First Travels,’ pp. 191, 439, 565; see also ‘Expedition to the Zambesi,’ 1865, p. 495, for an analogous case respecting a good breed of goats.Return to passage

[59] Andersson’s ‘Travels in South Africa,’ pp. 232, 318, 319.Return to passage

[60] Dr. Vavasseur, in ‘Bull. de La Soc. d’Acclimat.,’ tom. viii. 1861, p. 136.Return to passage

[61] ‘The Natural History of Dee Side,’ 1855, p. 476.Return to passage

[62] ‘Bull. de la Soc. d’Acclimat.,’ tom. vii. 1860, p. 457.Return to passage

[63] ‘Cattle,’ p. 48.Return to passage

[64] Livingstone’s Travels, p. 576; Andersson, ‘Lake Ngami,’ 1856, p. 222. With respect to the sale in Kaffraria, see ‘Quarterly Review,’ 1860, p. 139.Return to passage

[65] ‘Mémoire sur les Chinois’ (by the Jesuits), 1786, tom. xi. p. 57.Return to passage

[66] F. Michel, ‘Des Haras,’ pp. 47, 50.Return to passage

[67] Col. Hamilton Smith, Dogs, in ‘Nat. Lib.,’ vol. x. p. 103.Return to passage

[68] Azara, ‘Quadrupèdes du Paraguay,’ tom. ii. p. 324.Return to passage

[69] Sidney’s edit. of Youatt, 1860, pp. 24, 25.Return to passage

[70] ‘Rural Economy of Yorkshire,’ vol. ii. p. 182.Return to passage

[71] Moll et Gayot, ‘Du Boeuf,’ 1860, p. 547.Return to passage

[72] ‘The India Sporting Review,’ vol. ii. p. 181; ‘The Stud Farm,’ by Cecil, p. 58.Return to passage

[73] ‘The Horse,’ p. 22.Return to passage

[74] ‘History of England,’ vol. i. p. 316.Return to passage

[75] ‘Ueber Beständigkeit der Arten.’Return to passage

[76] Youatt on Sheep, p. 315.Return to passage

[77] ‘Ueber Shorthorn Rindvieh,’ 1857, s. 51.Return to passage

[78] Low, ‘Domesticated Animals,’ 1845, p. 363.Return to passage

[79] ‘Quarterly Review,’ 1849, p. 392.Return to passage

[80] H. von Nathusius, ‘Vorstudien . . . Schweineschädel,’ 1864, s 140.Return to passage

[81] See also Dr. Christ, in Rütimeyer’s ‘Pfahlbauten,’ 1861, s. 226.Return to passage

[82] The passage is given, ‘Bull. Soc. d’Acclimat.,’ 1858, p. 11.Return to passage

[83] ‘Journal of Horticulture,’ 1862, p. 394.Return to passage

[84] ‘Gardener’s Chronicle,’ 1857, p. 85.Return to passage

[85] See Mr. Wildman’s address to the Floricult. Soc. in ‘Gardener’s Chronicle,’ 1843, p. 86.Return to passage

[86] ‘Journal of Horticulture,’ Oct. 24th, 1865, p. 239.Return to passage

[87] Prescott’s ‘Hist. of Mexico,’ vol. ii. p. 61.Return to passage

[88] Sagaret, ‘Pomologie Physiologique,’ 1830, p. 47; Gallesio, ‘Teoria della Riproduzione,’ 1816, p. 88; Godron, ‘De l’Espèce,’ 1859, tom. 2 pp. 63, 67, 70. In my tenth and eleventh chapters I have given details on the potato; and I can confirm similar remarks with respect to the onion. I have also shown how far Naudin concurs in regard to the varieties of the melon.Return to passage

[89] Godron, ‘De l’Espèce,’ tom. ii. p. 27.Return to passage

[90] ‘The Anthropological Treatises of Blumenbach,’ 1856, p. 292.Return to passage

[91] Mr. J. J. Murphy, in his opening address to the Belfast Nat. Hist. Soc., as given in the ‘Belfast Northern Whig,’ Nov. 19th, 1866. Mr. Murphy here follows the line of argument against my views previously and more cautiously given by the Rev. C. Pritchard, Pres. Royal Astronomical Soc., in his sermon (Appendix, p. 33) preached before the British Association at Nottingham, 1866.Return to passage

[92] On the Vision of Fishes and Amphibia, translated in ‘Annals and Mag. of Nat. Hist.,’ vol. xviii., 1866, p. 469.Return to passage

[93] Sixth edition, 1872, p. 144.Return to passage

XXI. SELECTION, continued.

NATURAL SELECTION AS AFFECTING DOMESTIC PRODUCTIONS—CHARACTERS WHICH APPEAR OF TRIFLING VALUE OFTEN OF REAL IMPORTANCE—CIRCUMSTANCES FAVOURABLE TO SELECTION BY MAN—FACILITY IN PREVENTING CROSSES, AND THE NATURE OF THE CONDITIONS—CLOSE ATTENTION AND PERSEVERANCE INDISPENSABLE—THE PRODUCTION OF A LARGE NUMBER OF INDIVIDUALS ESPECIALLY FAVOURABLE—WHEN NO SELECTION IS APPLIED, DISTINCT RACES ARE NOT FORMED—HIGHLY-BRED ANIMALS LIABLE TO DEGENERATION—TENDENCY IN MAN TO CARRY THE SELECTION OF EACH CHARACTER TO AN EXTREME POINT, LEADING TO DIVERGENCE OF CHARACTER, RARELY TO CONVERGENCE—CHARACTERS CONTINUING TO VARY IN THE SAME DIRECTION IN WHICH THEY HAVE ALREADY VARIED—DIVERGENCE OF CHARACTER, WITH THE EXTINCTION OF INTERMEDIATE VARIETIES, LEADS TO DISTINCTNESS IN OUR DOMESTIC RACES—LIMIT TO THE POWER OF SELECTION—LAPSE OF TIME IMPORTANT—MANNER IN WHICH DOMESTIC RACES HAVE ORIGINATED—SUMMARY.

Natural Selection, or the Survival of the Fittest, as affecting domestic productions.—We know little on this head. But as animals kept by savages have to provide throughout the year their own food either entirely or to a large extent, it can hardly be doubted that in different countries, varieties differing in constitution and in various characters would succeed best, and so be naturally selected. Hence perhaps it is that the few domesticated animals kept by savages partake, as has been remarked by more than one writer, of the wild appearance of their masters, and likewise resemble natural species. Even in long-civilised countries, at least in the wilder parts, natural selection must act on our domestic races. It is obvious that varieties having very different habits, constitution, and structure, would succeed best on mountains and on rich lowland pastures. For example, the improved Leicester sheep were formerly taken to the Lammermuir Hills; but an intelligent sheep-master reported that “our coarse lean pastures were unequal to the task of supporting such heavy-bodied sheep; and they gradually dwindled away into less and less bulk: each generation was inferior to the preceding one; and when the spring was severe, seldom more than two-thirds of the lambs survived the ravages of the storms.”[1] So with the mountain cattle of North Wales and the Hebrides, it has been found that they could not withstand being crossed with the larger and more delicate lowland breeds. Two French naturalists, in describing the horses of Circassia, remark that, subjected as they are to extreme vicissitudes of climate, having to search for scanty pasture, and exposed to constant danger from wolves, the strongest and most vigorous alone survive.[2]

Every one must have been struck with the surpassing grace, strength, and vigour of the Game-cock, with its bold and confident air, its long, yet firm neck, compact body, powerful and closely pressed wings, muscular thighs, strong beak massive at the base, dense and sharp spurs set low on the legs for delivering the fatal blow, and its compact, glossy, and mail-like plumage serving as a defence. Now the English game-cock has not only been improved during many years by man’s careful selection, but in addition, as Mr. Tegetmeier has remarked,[3] by a kind of natural selection, for the strongest, most active and courageous birds have stricken down their antagonists in the cockpit, generation after generation, and have subsequently served as the progenitors of their race. The same kind of double selection has come into play with the carrier pigeon, for during their training the inferior birds fail to return home and are lost, so that even without selection by man only the superior birds propagate their race.

In Great Britain, in former times, almost every district had its own breed of cattle and sheep; “they were indigenous to the soil, climate, and pasturage of the locality on which they grazed: they seemed to have been formed for it and by it.”[4] But in this case we are quite unable to disentangle the effects of the direct action of the conditions of life,—of use or habit—of natural selection—and of that kind of selection which we have seen is occasionally and unconsciously followed by man even during the rudest periods of history.

Let us now look to the action of natural selection on special characters. Although nature is difficult to resist, yet man often strives against her power, and sometimes with success. From the facts to be given, it will also be seen that natural selection would powerfully affect many of our domestic productions if left unprotected. This is a point of much interest, for we thus learn that differences apparently of very slight importance would certainly determine the survival of a form when forced to struggle for its own existence. It may have occurred to some naturalists, as it formerly did to me, that, though selection acting under natural conditions would determine the structure of all important organs, yet that it could not affect characters which are esteemed by us of little importance; but this is an error to which we are eminently liable, from our ignorance of what characters are of real value to each living creature.

When man attempts to make a breed with some serious defect in structure, or in the mutual relation of the several parts, he will partly or completely fail, or encounter much difficulty; he is in fact resisted by a form of natural selection. We have seen that an attempt was once made in Yorkshire to breed cattle with enormous buttocks, but the cows perished so often in bringing forth their calves, that the attempt had to be given up. In rearing short-faced tumblers, Mr. Eaton says,[5] “I am convinced that better head and beak birds have perished in the shell than ever were hatched; the reason being that the amazingly short-faced bird cannot reach and break the shell with its beak, and so perishes.” Here is a more curious case, in which natural selection comes into play only at long intervals of time: during ordinary seasons the Niata cattle can graze as well as others, but occasionally, as from 1827 to 1830 the plains of La Plata suffer from long-continued droughts and the pasture is burnt up; at such times common cattle and horses perish by the thousand, but many survive by browsing on twigs, reeds, etc.; this the Niata cattle cannot so well effect from their upturned jaws and the shape of their lips; consequently, if not attended to, they perish before the other cattle. In Columbia, according to Roulin, there is a breed of nearly hairless cattle, called Pelones; these succeed in their native hot district, but are found too tender for the Cordillera; in this case, however, natural selection determines only the range of the variety. It is obvious that a host of artificial races could never survive in a state of nature;—such as Italian greyhounds,—hairless and almost toothless Turkish dogs,—fantail pigeons, which cannot fly well against a strong wind,—barbs and Polish fowls, with their vision impeded by their eye wattles and great topknots,—hornless bulls and rams, which consequently cannot cope with other males, and thus have a poor chance of leaving offspring,—seedless plants, and many other such cases.

Colour is generally esteemed by the systematic naturalist as unimportant: let us, therefore, see how far it indirectly affects our domestic productions, and how far it would affect them if they were left exposed to the full force of natural selection. In a future chapter I shall have to show that constitutional peculiarities of the strangest kind, entailing liability to the action of certain poisons, are correlated with the colour of the skin. I will here give a single case, on the high authority of Professor Wyman; he informs me that, being surprised at all the pigs in a part of Virginia being black, he made inquiries, and ascertained that these animals feed on the roots of the Lachnanthes tinctoria, which colours their bones pink, and, excepting in the case of the black varieties, causes the hoofs to drop off. Hence, as one of the squatters remarked, “we select the black members of the litter for raising, as they alone have a good chance of living.” So that here we have artificial and natural selection working hand in hand. I may add that in the Tarentino the inhabitants keep black sheep alone, because the Hypericum crispum abounds there; and this plant does not injure black sheep, but kills the white ones in about a fortnight’s time.[6]

Complexion, and liability to certain diseases, are believed to run together in man and the lower animals. Thus white terriers suffer more than those of any other colour from the fatal distemper.[7] In North America plum-trees are liable to a disease which Downing[8] believes is not caused by insects; the kinds bearing purple fruit are most affected, “and we have never known the green or yellow fruited varieties infected until the other sorts had first become filled with the knots.” On the other hand, peaches in North America suffer much from a disease called the yellows, which seems to be peculiar to that continent, and more than nine-tenths of the victims, “when the disease first appeared, were the yellow-fleshed peaches. The white-fleshed kinds are much more rarely attacked; in some parts of the country never.” In Mauritius, the white sugar-canes have of late years been so severely attacked by a disease, that many planters have been compelled to give up growing this variety (although fresh plants were imported from China for trial), and cultivate only red canes.[9] Now, if these plants had been forced to struggle with other competing plants and enemies, there cannot be a doubt that the colour of the flesh or skin of the fruit, unimportant as these characters are considered, would have rigorously determined their existence.

Liability to the attacks of parasites is also connected with colour. White chickens are certainly more subject than dark-coloured chickens to the “gapes,” which is caused by a parasitic worm in the trachea.[10] On the other hand, experience has shown that in France the caterpillars which produce white cocoons resist the deadly fungus better than those producing yellow cocoons.[11] Analogous facts have been observed with plants: a new and beautiful white onion, imported from France, though planted close to other kinds, was alone attacked by a parasitic fungus.[12] White verbenas are especially liable to mildew.[13] Near Malaga, during an early period of the vine-disease, the green sorts suffered most; “and red and black grapes, even when interwoven with the sick plants, suffered not at all.” In France whole groups of varieties were comparatively free, and others, such as the Chasselas, did not afford a single fortunate exception; but I do not know whether any correlation between colour and liability to disease was here observed.[14] In a former chapter it was shown how curiously liable one variety of the strawberry is to mildew.

It is certain that insects regulate in many cases the range and even the existence of the higher animals, whilst living under their natural conditions. Under domestication light-coloured animals suffer most: in Thuringia[15] the inhabitants do not like grey, white, or pale cattle, because they are much more troubled by various kinds of flies than the brown, red, or black cattle. An Albino negro, it has been remarked,[16] was peculiarly sensitive to the bites of insects. In the West Indies[17] it is said that “the only horned cattle fit for work are those which have a good deal of black in them. The white are terribly tormented by the insects; and they are weak and sluggish in proportion to the white.”

In Devonshire there is a prejudice against white pigs, because it is believed that the sun blisters them when turned out;[18] and I knew a man who would not keep white pigs in Kent, for the same reason. The scorching of flowers by the sun seems likewise to depend much on colour; thus, dark pelargoniums suffer most; and from various accounts it is clear that the cloth-of-gold variety will not withstand a degree of exposure to sunshine which other varieties enjoy. Another amateur asserts that not only all dark-coloured verbenas, but likewise scarlets, suffer from the sun: “the paler kinds stand better, and pale blue is perhaps the best of all.” So again with the heartsease (Viola tricolor); hot weather suits the blotched sorts, whilst it destroys the beautiful markings of some other kinds.[19] During one extremely cold season in Holland all red-flowered hyacinths were observed to be very inferior in quality. It is believed by many agriculturists that red wheat is hardier in northern climates than white wheat.[20]

With animals, white varieties from being conspicuous are the most liable to be attacked by beasts and birds of prey. In parts of France and Germany where hawks abound, persons are advised not to keep white pigeons; for, as Parmentier says, “it is certain that in a flock the white always first fall victims to the kite.” In Belgium, where so many societies have been established for the flight of carrier-pigeons, white is the one colour which for the same reason is disliked.[21] Prof. G. Jaeger[22] whilst fishing found four pigeons which had been killed by hawks, and all were white; on another occasion he examined the eyrie of a hawk, and the feathers of the pigeons which had been caught were all of a white or yellow colour. On the other hand, it is said that the sea-eagle (Falco ossifragus, Linn.) on the west coast of Ireland picks out the black fowls, so that “the villagers avoid as much as possible rearing birds of that colour.” M. Daudin,[23] speaking of white rabbits kept in warrens in Russia, remarks that their colour is a great disadvantage, as they are thus more exposed to attack, and can be seen during bright nights from a distance. A gentleman in Kent, who failed to stock his woods with a nearly white and hardy kind of rabbit, accounted in the same manner for their early disappearance. Any one who will watch a white cat prowling after her prey will soon perceive under what a disadvantage she lies.

The white Tartarian cherry, “owing either to its colour being so much like that of the leaves, or to the fruit always appearing from a distance unripe,” is not so readily attacked by birds as other sorts. The yellow-fruited raspberry, which generally comes nearly true by seed, “is very little molested by birds, who evidently are not fond of it; so that nets may be dispensed with in places where nothing else will protect the red fruit.”[24] This immunity, though a benefit to the gardener, would be a disadvantage in a state of nature both to the cherry and raspberry, as dissemination depends on birds. I noticed during several winters that some trees of the yellow-berried holly, which were raised from seed from a tree found wild by my father remained covered with fruit, whilst not a scarlet berry could be seen on the adjoining trees of the common kind. A friend informs me that a mountain-ash (Pyrus aucuparia) growing in his garden bears berries which, though not differently coloured, are always devoured by birds before those on the other trees. This variety of the mountain-ash would thus be more freely disseminated, and the yellow-berried variety of the holly less freely, than the common varieties of these two trees.

Independently of colour, trifling differences are sometimes found to be of importance to plants under cultivation, and would be of paramount importance if they had to fight their own battle and to struggle with many competitors. The thin-shelled peas, called pois sans parchemin, are attacked by birds[25] much more commonly than ordinary peas. On the other hand, the purple-podded pea, which has a hard shell, escaped the attacks of tomtits (Parus major) in my garden far better than any other kind. The thin-shelled walnut likewise suffers greatly from the tomtit.[26] These same birds have been observed to pass over and thus favour the filbert, destroying only the other kinds of nuts which grew in the same orchard.[27]

Certain varieties of the pear have soft bark, and these suffer severely from wood-boring beetles; whilst other varieties are known to resist their attacks much better.[28] In North America the smoothness, or absence of down on the fruit, makes a great difference in the attacks of the weevil, “which is the uncompromising foe of all smooth stone-fruits;” and the cultivator “has the frequent mortification of seeing nearly all, or indeed often the whole crop, fall from the trees when half or two-thirds grown.” Hence the nectarine suffers more than the peach. A particular variety of the Morello cherry, raised in North America, is, without any assignable cause, more liable to be injured by this same insect than other cherry-trees.[29] From some unknown cause, certain varieties of the apple enjoy, as we have seen, the great advantage in various parts of the world of not being infested by the coccus. On the other hand, a particular case has been recorded in which aphides confined themselves to the Winter Nelis pear and touched no other kind in an extensive orchard.[30] The existence of minute glands on the leaves of peaches, nectarines, and apricots, would not be esteemed by botanists as a character of the least importance for they are present or absent in closely-related sub-varieties, descended from the same parent-tree; yet there is good evidence[31] that the absence of glands leads to mildew, which is highly injurious to these trees.

A difference either in flavour or in the amount of nutriment in certain varieties causes them to be more eagerly attacked by various enemies than other varieties of the same species. Bullfinches (Pyrrhula vulgaris) injure our fruit-trees by devouring the flower-buds, and a pair of these birds have been seen “to denude a large plum-tree in a couple of days of almost every bud;” but certain varieties[32] of the apple and thorn (Cratægus oxyacantha) are more especially liable to be attacked. A striking instance of this was observed in Mr. Rivers’s garden, in which two rows of a particular variety of plum[33] had to be carefully protected, as they were usually stripped of all their buds during the winter, whilst other sorts growing near them escaped. The root (or enlarged stem) of Laing’s Swedish turnip is preferred by hares, and therefore suffers more than other varieties. Hares and rabbits eat down common rye before St. John’s-day-rye, when both grow together.[34] In the south of France, when an orchard of almond-trees is formed, the nuts of the bitter variety are sown, “in order that they may not be devoured by field-mice”;[35] so we see the use of the bitter principle in almonds.

Other slight differences, which would be thought quite unimportant, are no doubt sometimes of great service both to plants and animals. The Whitesmith’s gooseberry, as formerly stated, produces its leaves later than other varieties, and, as the flowers are thus left unprotected, the fruit often fails. In one variety of the cherry, according to Mr. Rivers,[36] the petals are much curled backwards, and in consequence of this the stigmas were observed to be killed by a severe frost; whilst at the same time, in another variety with incurved petals, the stigmas were not in the least injured. The straw of the Fenton wheat is remarkably unequal in height; and a competent observer believes that this variety is highly productive, partly because the ears from being distributed at various heights above the ground are less crowded together. The same observer maintains that in the upright varieties the divergent awns are serviceable by breaking the shocks when the ears are dashed together by the wind.[37] If several varieties of a plant are grown together, and the seed is indiscriminately harvested, it is clear that the hardier and more productive kinds will, by a sort of natural selection, gradually prevail over the others; this takes place, as Colonel Le Couteur believes,[38] in our wheat-fields, for, as formerly shown, no variety is quite uniform in character. The same thing, as I am assured by nurserymen, would take place in our flower-gardens, if the seed of the different varieties were not separately saved. When the eggs of the wild and tame duck are hatched together, the young wild ducks almost invariably perish, from being of smaller size and not getting their fair share of food.[39]

Facts in sufficient number have now been given showing that natural selection often checks, but occasionally favours, man’s power of selection. These facts teach us, in addition, a valuable lesson, namely, that we ought to be extremely cautious in judging what characters are of importance in a state of nature to animals and plants, which have to struggle for existence from the hour of their birth to that of their death,—their existence depending on conditions, about which we are profoundly ignorant.

Circumstances favourable to Selection by Man.

The possibility of selection rests on variability, and this, as we shall see in the following chapters, mainly depends on changed conditions of life, but is governed by infinitely complex and unknown laws. Domestication, even when long continued, occasionally causes but a small amount of variability, as in the case of the goose and turkey. The slight differences, however, which characterise each individual animal and plant would in most, probably in all, cases suffice for the production of distinct races through careful and prolonged selection. We see what selection, though acting on mere individual differences, can effect when families of cattle, sheep, pigeons, etc., of the same race, have been separately bred during a number of years by different men without any wish on their part to modify the breed. We see the same fact in the difference between hounds bred for hunting in different districts,[40] and in many other such cases.

In order that selection should produce any result, it is manifest that the crossing of distinct races must be prevented; hence facility in pairing, as with the pigeon, is highly favourable for the work; and difficulty in pairing, as with cats, prevents the formation of distinct breeds. On nearly the same principle the cattle of the small island of Jersey have been improved in their milking qualities “with a rapidity that could not have been obtained in a widely extended country like France.”[41] Although free crossing is a danger on the one side which every one can see, too close interbreeding is a hidden danger on the other side. Unfavourable conditions of life overrule the power of selection. Our improved heavy breeds of cattle and sheep could not have been formed on mountainous pastures; nor could dray-horses have been raised on a barren and inhospitable land, such as the Falkland Islands, where even the light horses of La Plata rapidly decrease in size. It seems impossible to preserve several English breeds of sheep in France; for as soon as the lambs are weaned their vigour decays as the heat of the summer increases:[42] it would be impossible to give great length of wool to sheep within the tropics; yet selection has kept the Merino breed nearly true under diversified and unfavourable conditions. The power of selection is so great, that breeds of the dog, sheep, and poultry, of the largest and smallest size, long and short beaked pigeons, and other breeds with opposite characters, have had their characteristic qualities augmented, though treated in every way alike, being exposed to the same climate and fed on the same food. Selection, however, is either checked or favoured by the effects of use or habit. Our wonderfully-improved pigs could never have been formed if they had been forced to search for their own food; the English racehorse and greyhound could not have been improved up to their present high standard of excellence without constant training.

As conspicuous deviations of structure occur rarely, the improvement of each breed is generally the result of the selection of slight individual differences. Hence the closest attention, the sharpest powers of observation, and indomitable perseverance, are indispensable. It is, also, highly important that many individuals of the breed which is to be improved should be raised; for thus there will be a better chance of the appearance of variations in the right direction, and individuals varying in an unfavourable manner may be freely rejected or destroyed. But that a large number of individuals should be raised, it is necessary that the conditions of life should favour the propagation of the species. Had the peacock been reared as easily as the fowl, we should probably ere this have had many distinct races. We see the importance of a large number of plants, from the fact of nursery gardeners almost always beating amateurs in the exhibition of new varieties. In 1845 it was estimated[43] that between 4000 and 5000 pelargoniums were annually raised from seed in England, yet a decidedly improved variety is rarely obtained. At Messrs. Carter’s grounds, in Essex, where such flowers as the Lobelia, Nemophila, Mignonette, etc., are grown by the acre for seed, “scarcely a season passes without some new kinds being raised, or some improvement effected on old kinds.”[44] At Kew, as Mr. Beaton remarks, where many seedlings of common plants are raised, “you see new forms of Laburnums, Spiraeas, and other shrubs.”[45] So with animals: Marshall,[46] in speaking of the sheep in one part of Yorkshire, remarks, “as they belong to poor people, and are mostly in small lots, they never can be improved.” Lord Rivers, when asked how he succeeded in always having first-rate greyhounds, answered, “I breed many, and hang many.” This, as another man remarks, “was the secret of his success; and the same will be found in exhibiting fowls,— successful competitors breed largely, and keep the best.”[47]

It follows from this that the capacity of breeding at an early age and at short intervals, as with pigeons, rabbits, etc., facilitates selection; for the result is thus soon made visible, and perseverance in the work encouraged. It can hardly be an accident that the great majority of the culinary and agricultural plants which have yielded numerous races are annuals or biennials, which therefore are capable of rapid propagation, and thus of improvement. Sea-kale, asparagus, common and Jerusalem artichokes, potatoes, and onions, must be excepted, as they are perennials: but onions are propagated like annuals, and of the other plants just specified, none, with the exception of the potato, have yielded in this country more than one or two varieties. In the Mediterranean region, where artichokes are often raised from seed, there are several kinds, as I hear from Mr. Bentham. No doubt fruit-trees, which cannot be propagated quickly by seed, have yielded a host of varieties, though not permanent races; but these, judging from prehistoric remains, have been produced at a comparatively late period.

A species may be highly variable, but distinct races will not be formed, if from any cause selection be not applied. It would be difficult to select slight variations in fishes from their place of habitation; and though the carp is extremely variable and is much attended to in Germany, only one well-marked race has been formed, as I hear from Lord A. Russell, namely the spiegel-carpe; and this is carefully secluded from the common scaly kind. On the other hand, a closely allied species, the gold-fish, from being reared in small vessels, and from having been carefully attended to by the Chinese, has yielded many races. Neither the bee, which has been semi-domesticated from an extremely remote period, nor the cochineal insect, which was cultivated by the aboriginal Mexicans,[48] has yielded races; and it would be impossible to match the queen-bee with any particular drone, and most difficult to match cochineal insects. Silk-moths, on the other hand, have been subjected to rigorous selection, and have produced a host of races. Cats, which from their nocturnal habits cannot be selected for breeding, do not, as formerly remarked, yield distinct races within the same country. Dogs are held in abomination in the East, and their breeding is neglected; consequently, as Prof. Moritz Wagner[49] remarks, one kind alone exists there. The ass in England varies much in colour and size; but as it is an animal of little value and bred by poor people, there has been no selection, and distinct races have not been formed. We must not attribute the inferiority of our asses to climate, for in India they are of even smaller size than in Europe. But when selection is brought to bear on the ass, all is changed. Near Cordova, as I am informed (Feb. 1860) by Mr. W. E. Webb, C.E., they are carefully bred, as much as 200_l._ having been paid for a stallion ass, and they have been immensely improved. In Kentucky, asses have been imported (for breeding mules) from Spain, Malta, and France; these “seldom averaged more than fourteen hands high: but the Kentuckians, by great care, have raised them up to fifteen hands, and sometimes even to sixteen. The prices paid for these splendid animals, for such they really are, will prove how much they are in request. One male, of great celebrity, was sold for upwards of one thousand pounds sterling.” These choice asses are sent to cattle-shows, a day being given for their exhibition.[50]

Analogous facts have been observed with plants: the nutmeg-tree in the Malay archipelago is highly variable, but there has been no selection, and there are no distinct races.[51] The common mignonette (Reseda odorata), from bearing inconspicuous flowers, valued solely for their fragrance, “remains in the same unimproved condition as when first introduced.”[52] Our common forest-trees are very variable, as may be seen in every extensive nursery-ground; but as they are not valued like fruit-trees, and as they seed late in life, no selection has been applied to them; consequently, as Mr. Patrick Matthews remarks,[53] they have not yielded distinct races, leafing at different periods, growing to different sizes, and producing timber fit for different purposes. We have gained only some fanciful and semi-monstrous varieties, which no doubt appeared suddenly as we now see them.

Some botanists have argued that plants cannot have so strong a tendency to vary as is generally supposed, because many species long grown in botanic gardens, or unintentionally cultivated year after year mingled with our corn crops, have not produced distinct races; but this is accounted for by slight variations not having been selected and propagated. Let a plant which is now grown in a botanic garden, or any common weed, be cultivated on a large scale, and let a sharp-sighted gardener look out for each slight variety and sow the seed, and then, if distinct races are not produced, the argument will be valid.

The importance of selection is likewise shown by considering special characters. For instance, with most breeds of fowls the form of the comb and the colour of the plumage have been attended to, and are eminently characteristic of each race; but in Dorkings fashion has never demanded uniformity of comb or colour; and the utmost diversity in these respects prevails. Rose-combs, double-combs, cup-combs, etc., and colours of all kinds, may be seen in purely bred and closely related Dorking fowls, whilst other points, such as the general form of body, and the presence of an additional toe, have been attended to, and are invariably present. It has also been ascertained that colour can be fixed in this breed, as well as in any other.[54]

During the formation or improvement of a breed, its members will always be found to vary much in those characters to which especial attention is directed, and of which each slight improvement is eagerly sought and selected. Thus, with short-faced tumbler-pigeons, the shortness of the beak, shape of head and plumage,—with carriers, the length of the beak and wattle,—with fantails, the tail and carriage,—with Spanish fowls, the white face and comb,—with long-eared rabbits, the length of ear, are all points which are eminently variable. So it is in every case; and the large price paid for first-rate animals proves the difficulty of breeding them up to the highest standard of excellence. This subject has been discussed by fanciers,[55] and the greater prizes given for highly improved breeds, in comparison with those given for old breeds which are not now undergoing rapid improvement, have been fully justified. Nathusius makes[56] a similar remark when discussing the less uniform character of improved Shorthorn cattle and of the English horse, in comparison, for example, with the unennobled cattle of Hungary, or with the horses of the Asiatic steppes. This want of uniformity in the parts which at the time are undergoing selection chiefly depends on the strength of the principle of reversion; but it likewise depends to a certain extent on the continued variability of the parts which have recently varied. That the same parts do continue varying in the same manner we must admit, for if it were not so, there could be no improvement beyond an early standard of excellence, and we know that such improvement is not only possible, but is of general occurrence.

As a consequence of continued variability, and more especially of reversion, all highly improved races, if neglected or not subjected to incessant selection, soon degenerate. Youatt gives a curious instance of this in some cattle formerly kept in Glamorganshire; but in this case the cattle were not fed with sufficient care. Mr. Baker, in his memoir on the Horse, sums up: “It must have been observed in the preceding pages that, whenever there has been neglect, the breed has proportionally deteriorated.”[57] If a considerable number of improved cattle, sheep, or other animals of the same race, were allowed to breed freely together, with no selection, but with no change in their condition of life, there can be no doubt that after a score or hundred generations they would be very far from excellent of their kind; but, from what we see of the many common races of dogs, cattle, fowls, pigeons, etc., which without any particular care have long retained nearly the same character, we have no grounds for believing that they would altogether depart from their type.

It is a general belief amongst breeders that characters of all kinds become fixed by long-continued inheritance. But I have attempted to show in the fourteenth chapter that this belief apparently resolves itself into the following proposition, namely, that all characters whatever, whether recently acquired or ancient, tend to be transmitted, but that those which have already long withstood all counteracting influences, will, as a general rule, continue to withstand them, and consequently be faithfully transmitted.

Tendency in Man to carry the practice of Selection to an extreme point.

It is an important principle that in the process of selection man almost invariably wishes to go to an extreme point. Thus, there is no limit to his desire to breed certain kinds of horses and dogs as fleet as possible, and others as strong as possible; certain kinds of sheep for extreme fineness, and others for extreme length of wool; and he wishes to produce fruit, grain, tubers, and other useful parts of plants, as large and excellent as possible. With animals bred for amusement, the same principle is even more powerful; for fashion, as we see in our dress, always runs to extremes. This view has been expressly admitted by fanciers. Instances were given in the chapters on the pigeon, but here is another: Mr. Eaton, after describing a comparatively new variety, namely, the Archangel, remarks, “What fanciers intend doing with this bird I am at a loss to know, whether they intend to breed it down to the tumbler’s head and beak, or carry it out to the carrier’s head and beak; leaving it as they found it, is not progressing.” Ferguson, speaking of fowls, says, “their peculiarities, whatever they may be, must necessarily be fully developed: a little peculiarity forms nought but ugliness, seeing it violates the existing laws of symmetry.” So Mr. Brent, in discussing the merits of the sub-varieties of the Belgian canary-bird, remarks, “Fanciers always go to extremes; they do not admire indefinite properties.”[58]

This principle, which necessarily leads to divergence of character, explains the present state of various domestic races. We can thus see how it is that racehorses and dray-horses, greyhounds and mastiffs, which are opposed to each other in every character,—how varieties so distinct as Cochin-china fowls and bantams, or carrier-pigeons with very long beaks, and tumblers with excessively short beaks, have been derived from the same stock. As each breed is slowly improved, the inferior varieties are first neglected and finally lost. In a few cases, by the aid of old records, or from intermediate varieties still existing in countries where other fashions have prevailed, we are enabled partially to trace the graduated changes through which certain breeds have passed. Selection, whether methodical or unconscious, always tending towards an extreme point, together with the neglect and slow extinction of the intermediate and less-valued forms, is the key which unlocks the mystery of how man has produced such wonderful results.

In a few instances selection, guided by utility for a single purpose, has led to convergence of character. All the improved and different races of the pig, as Nathusius has well shown,[59] closely approach each other in character, in their shortened legs and muzzles, their almost hairless, large, rounded bodies, and small tusks. We see some degree of convergence in the similar outline of the body in well-bred cattle belonging to distinct races.[60] I know of no other such cases.

Continued divergence of character depends on, and is indeed a clear proof, as previously remarked, of the same parts continuing to vary in the same direction. The tendency to mere general variability or plasticity of organisation can certainly be inherited, even from one parent, as has been shown by Gärtner and Kölreuter, in the production of varying hybrids from two species, of which one alone was variable. It is in itself probable that, when an organ has varied in any manner, it will again vary in the same manner, if the conditions which first caused the being to vary remain, as far as can be judged, the same. This is either tacitly or expressly admitted by all horticulturists: if a gardener observes one or two additional petals in a flower, he feels confident that in a few generations he will be able to raise a double flower, crowded with petals. Some of the seedlings from the weeping Moccas oak were so prostrate that they only crawled along the ground. A seedling from the fastigiate or upright Irish yew is described as differing greatly from the parent-form “by the exaggeration of the fastigiate habit of its branches.”[61] Mr. Shirreff, who has been highly successful in raising new kinds of wheat, remarks, “A good variety may safely be regarded as the forerunner of a better one.”[62] A great rose-grower, Mr. Rivers, has made the same remark with respect to roses. Sageret,[63] who had large experience, in speaking of the future progress of fruit-trees, observes that the most important principle is “that the more plants have departed from their original type, the more they tend to depart from it.” There is apparently much truth in this remark; for we can in no other way understand the surprising amount of difference between varieties in the parts or qualities which are valued, whilst other parts retain nearly their original character.

The foregoing discussion naturally leads to the question, what is the limit to the possible amount of variation in any part or quality, and, consequently, is there any limit to what selection can effect? Will a racehorse ever be reared fleeter than Eclipse? Can our prize-cattle and sheep be still further improved? Will a gooseberry ever weigh more than that produced by “London” in 1852? Will the beet-root in France yield a greater percentage of sugar? Will future varieties of wheat and other grain produce heavier crops than our present varieties? These questions cannot be positively answered; but it is certain that we ought to be cautious in answering them by a negative. In some lines of variation the limit has probably been reached. Youatt believes that the reduction of bone in some of our sheep has already been carried so far that it entails great delicacy of constitution.[64] But seeing the great improvement within recent times in our cattle and sheep, and especially in our pigs; seeing the wonderful increase in weight in our poultry of all kinds during the last few years; he would be a bold man who would assert that perfection has been reached. It has often been said that Eclipse never was, and never will be, beaten in speed by any other horse; but on making inquiries I find that the best judges believe that our present racehorses are fleeter.[65] The attempt to raise a new variety of wheat more productive than the many old kinds, might have been thought until lately quite hopeless; but this has been effected by Major Hallett, by careful selection. With respect to almost all our animals and plants, those who are best qualified to judge do not believe that the extreme point of perfection has yet been reached even in the characters which have already been carried to a high standard. For instance, the short-faced tumbler-pigeon has been greatly modified; nevertheless, according to Mr. Eaton[66] “the field is still as open for fresh competitors as it was one hundred years ago.” Over and over again it has been said that perfection had been attained with our flowers, but a higher standard has soon been reached. Hardly any fruit has been more improved than the strawberry, yet a great authority remarks,[67] “it must not be concealed that we are far from the extreme limits at which we may arrive.”

No doubt there is a limit beyond which the organisation cannot be modified compatibly with health or life. The extreme degree of fleetness, for instance, of which a terrestrial animal is capable, may have been acquired by our present racehorses; but as Mr. Wallace has well remarked,[68] the question that interests us, “is not whether indefinite and unlimited change in any or all directions is possible, but whether such differences as do occur in nature could have been produced by the accumulation of varieties by selection.” And in the case of our domestic productions, there can be no doubt that many parts of the organisation, to which man has attended, have been thus modified to a greater degree than the corresponding parts in the natural species of the same genera or even families. We see this in the form and size of our light and heavy dogs or horses,—in the beak and many other characters of our pigeons,—in the size and quality of many fruits,—in comparison with the species belonging to the same natural groups.

Time is an important element in the formation of our domestic races, as it permits innumerable individuals to be born, and these when exposed to diversified conditions are rendered variable. Methodical selection has been occasionally practised from an ancient period to the present day, even by semi-civilised people, and during former times will have produced some effect. Unconscious selection will have been still more effective; for during a lengthened period the more valuable individual animals will occasionally have been saved, and the less valuable neglected. In the course of time, different varieties, especially in the less civilised countries, will also have been more or less modified through natural selection. It is generally believed, though on this head we have little or no evidence, that new characters in time become fixed; and after having long remained fixed it seems possible that under new conditions they might again be rendered variable.

How great the lapse of time has been since man first domesticated animals and cultivated plants, we begin dimly to see. When the lake-dwellings of Switzerland were inhabited during the Neolithic period, several animals were already domesticated and various plants cultivated. The science of language tells us that the art of ploughing and sowing the land was followed, and the chief animals had been already domesticated, at an epoch so immensely remote, that the Sanskrit, Greek, Latin, Gothic, Celtic, and Sclavonic languages had not as yet diverged from their common parent-tongue.[69]

It is scarcely possible to overrate the effects of selection occasionally carried on in various ways and places during thousands of generations. All that we know, and, in a still stronger degree, all that we do not know,[70] of the history of the great majority of our breeds, even of our more modern breeds, agrees with the view that their production, through the action of unconscious and methodical selection, has been almost insensibly slow. When a man attends rather more closely than is usual to the breeding of his animals, he is almost sure to improve them to a slight extent. They are in consequence valued in his immediate neighbourhood, and are bred by others; and their characteristic features, whatever these may be, will then slowly but steadily be increased, sometimes by methodical and almost always by unconscious selection. At last a strain, deserving to be called a sub-variety, becomes a little more widely known, receives a local name, and spreads. The spreading will have been extremely slow during ancient and less civilised times, but now is rapid. By the time that the new breed had assumed a somewhat distinct character, its history, hardly noticed at the time, will have been completely forgotten; for, as Low remarks,[71] “we know how quickly the memory of such events is effaced.”

As soon as a new breed is thus formed, it is liable through the same process to break up into new strains and sub-varieties. For different varieties are suited for, and are valued under, different circumstances. Fashion changes, but, should a fashion last for even a moderate length of time, so strong is the principle of inheritance, that some effect will probably be impressed on the breed. Thus varieties go on increasing in number, and history shows us how wonderfully they have increased since the earliest records.[72] As each new variety is produced, the earlier, intermediate, and less valuable forms will be neglected, and perish. When a breed, from not being valued, is kept in small numbers, its extinction almost inevitably follows sooner or later, either from accidental causes of destruction or from close interbreeding; and this is an event which, in the case of well-marked breeds, excites attention. The birth or production of a new domestic race is so slow a process that it escapes notice; its death or destruction is comparatively sudden, is often recorded, and when too late sometimes regretted.

Several authors have drawn a wide distinction between artificial and natural races. The latter are more uniform in character, possessing in a high degree the appearance of natural species, and are of ancient origin. They are generally found in less civilised countries, and have probably been largely modified by natural selection, and only to a small extent by man’s unconscious and methodical selection. They have, also, during a long period, been directly acted on by the physical conditions of the countries which they inhabit. The so-called artificial races, on the other hand, are not so uniform in character; some have a semi-monstrous character, such as “the wry-legged terriers so useful in rabbit-shooting,”[73] turnspit dogs, ancon sheep, niata oxen, Polish fowls, fantail-pigeons, etc.; their characteristic features have generally been acquired suddenly, though subsequently increased by careful selections in many cases. Other races, which certainly must be called artificial, for they have been largely modified by methodical selection and by crossing, as the English racehorse, terrier-dogs, the English game-cock, Antwerp carrier-pigeons, etc., nevertheless cannot be said to have an unnatural appearance; and no distinct line, as it seems to me, can be drawn between natural and artificial races.

It is not surprising that domestic races should generally present a different aspect from natural species. Man selects and propagates modifications solely for his own use or fancy, and not for the creature’s own good. His attention is struck by strongly marked modifications, which have appeared suddenly, due to some great disturbing cause in the organisation. He attends almost exclusively to external characters; and when he succeeds in modifying internal organs,—when for instance he reduces the bones and offal, or loads the viscera with fat, or gives early maturity, etc.-the chances are strong that he will at the same time weaken the constitution. On the other hand, when an animal has to struggle throughout its life with many competitors and enemies, under circumstances inconceivably complex and liable to change, modifications of the most varied nature in the internal organs as well as in external characters, in the functions and mutual relations of parts, will be rigorously tested, preserved, or rejected. Natural selection often checks man’s comparatively feeble and capricious attempts at improvement; and if it were not so, the result of his work, and of nature’s work, would be even still more different. Nevertheless, we must not overrate the amount of difference between natural species and domestic races; the most experienced naturalists have often disputed whether the latter are descended from one or from several aboriginal stocks, and this clearly shows that there is no palpable difference between species and races.

Domestic races propagate their kind far more truly, and endure for munch longer periods, than most naturalists are willing to admit. Breeders feel no doubt on this head: ask a man who has long reared Shorthorn or Hereford cattle, Leicester or Southdown sheep, Spanish or Game poultry, tumbler or carrier-pigeons, whether these races may not have been derived from common progenitors, and he will probably laugh you to scorn. The breeder admits that he may hope to produce sheep with finer or longer wool and with better carcases, or handsomer fowls, or carrier-pigeons with beaks just perceptibly longer to the practised eye, and thus be successful at an exhibition. Thus far he will go, but no farther. He does not reflect on what follows from adding up during a long course of time many slight, successive modifications; nor does he reflect on the former existence of numerous varieties, connecting the links in each divergent line of descent. He concludes, as was shown in the earlier chapters, that all the chief breeds to which he has long attended are aboriginal productions. The systematic naturalist, on the other hand, who generally knows nothing of the art of breeding, who does not pretend to know how and when the several domestic races were formed, who cannot have seen the intermediate gradations, for they do not now exist, nevertheless feels no doubt that these races are sprung from a single source. But ask him whether the closely allied natural species which he has studied may not have descended from a common progenitor, and he in his turn will perhaps reject the notion with scorn. Thus the naturalist and breeder may mutually learn a useful lesson from each other.

Summary on Selection by Man.—There can be no doubt that methodical selection has effected and will effect wonderful results. It was occasionally practised in ancient times, and is still practised by semi-civilised people. Characters of the highest importance, and others of trifling value, have been attended to, and modified. I need not here repeat what has been so often said on the part which unconscious selection has played: we see its power in the difference between flocks which have been separately bred, and in the slow changes, as circumstances have slowly changed, which many animals have undergone in the same country, or when transported into a foreign land. We see the combined effects of methodical and unconscious selection, in the great amount of difference in those parts or qualities which are valued by man in comparison with the parts which are not valued, and consequently have not been attended to. Natural selection often determines man’s power of selection. We sometimes err in imagining that characters, which are considered as unimportant by the systematic naturalist, could not be affected by the struggle for existence, and could not be acted on by natural selection; but striking cases have been given, showing how great an error this is.

The possibility of selection coming into action rests on variability; and this is mainly caused, as we shall hereafter see, by changes in the conditions of life. Selection is sometimes rendered difficult, or even impossible, by the conditions being opposed to the desired character or quality. It is sometimes checked by the lessened fertility and weakened constitution which follow from long-continued close interbreeding. That methodical selection may be successful, the closest attention and discernment, combined with unwearied patience, are absolutely necessary; and these same qualities, though not indispensable, are highly serviceable in the case of unconscious selection. It is almost necessary that a large number of individuals should be reared; for thus there will be a fair chance of variations of the desired nature arising, and of every individual with the slightest blemish or in any degree inferior being freely rejected. Hence length of time is an important element of success. Thus, also, reproduction at an early age and at short intervals favours the work. Facility in pairing animals, or their inhabiting a confined area, is advantageous as a check to free crossing. Whenever and wherever selection is not practised, distinct races are not formed within the same country. When any one part of the body or one quality is not attended to, it remains either unchanged or varies in a fluctuating manner, whilst at the same time other parts and other qualities may become permanently and greatly modified. But from the tendency to reversion and to continued variability, those parts or organs which are now undergoing rapid improvement through selection, are likewise found to vary much. Consequently highly-bred animals when neglected soon degenerate; but we have no reason to believe that the effects of long-continued selection would, if the conditions of life remained the same, be soon and completely lost.

Man always tends to go to an extreme point in the selection, whether methodical or unconscious, of all useful and pleasing qualities. This is an important principle, as it leads to continued divergence, and in some rare cases to convergence of character. The possibility of continued divergence rests on the tendency in each part or organ to go on varying in the same manner in which it has already varied; and that this occurs, is proved by the steady and gradual improvement of many animals and plants during lengthened periods. The principle of divergence of character, combined with the neglect and final extinction of all previous, less-valued, and intermediate varieties, explains the amount of difference and the distinctness of our several races. Although we may have reached the utmost limit to which certain characters can be modified, yet we are far from having reached, as we have good reason to believe, the limit in the majority of cases. Finally, from the difference between selection as carried on by man and by nature, we can understand how it is that domestic races often, though by no means always, differ in general aspect from closely allied natural species.

Throughout this chapter and elsewhere I have spoken of selection as the paramount power, yet its action absolutely depends on what we in our ignorance call spontaneous or accidental variability. Let an architect be compelled to build an edifice with uncut stones, fallen from a precipice. The shape of each fragment may be called accidental; yet the shape of each has been determined by the force of gravity, the nature of the rock, and the slope of the precipice,—events and circumstances, all of which depend on natural laws; but there is no relation between these laws and the purpose for which each fragment is used by the builder. In the same manner the variations of each creature are determined by fixed and immutable laws; but these bear no relation to the living structure which is slowly built up through the power of selection, whether this be natural or artificial selection.

If our architect succeeded in rearing a noble edifice, using the rough wedge-shaped fragments for the arches, the longer stones for the lintels, and so forth, we should admire his skill even in a higher degree than if he had used stones shaped for the purpose. So it is with selection, whether applied by man or by nature; for although variability is indispensably necessary, yet, when we look at some highly complex and excellently adapted organism, variability sinks to a quite subordinate position in importance in comparison with selection, in the same manner as the shape of each fragment used by our supposed architect is unimportant in comparison with his skill.

REFERENCES

[1] Quoted by Youatt on Sheep, p. 325. See also Youatt on Cattle, pp. 62, 69.Return to passage

[2] MM. Lherbette and De Quatrefages, in ‘Bull. Soc. d’Acclimat.,’ tom. viii. 1861, p. 311.Return to passage

[3] ‘The Poultry Book,’ 1866, p. 123. Mr. Tegetmeier, ‘The Homing or Carrier Pigeon,’ 1871, pp. 45-58.Return to passage

[4] Youatt on Sheep, p. 312.Return to passage

[5] ‘Treatise on the Almond Tumbler,’ 1851, p. 33.Return to passage

[6] Dr. Heusinger, ‘Wochenschrift für die Heilkunde,’ Berlin, 1846, s. 279.Return to passage

[7] Youatt on the Dog, p. 232.Return to passage

[8] ‘The Fruit-trees of America,’ 1845, p. 270: for peaches, p. 466.Return to passage

[9] ‘Proc. Royal Soc. of Arts and Sciences of Mauritius,’ 1852, p. 135.Return to passage

[10] ‘Gardener’s Chronicle,’ 1856, p. 379.Return to passage

[11] Quatrefages, ‘Maladies Actuelles du Ver à Soie,’ 1859, pp. 12, 214.Return to passage

[12] ‘Gardener’s Chronicle,’ 1851, p. 595.Return to passage

[13] ‘Journal of Horticulture,’ 1862, p. 476.Return to passage

[14] ‘Gardener’s Chronicle,’ 1852, pp. 435, 691.Return to passage

[15] Bechstein, ‘Naturgesch. Deutschlands,’ 1801, B. 1 s. 310.Return to passage

[16] Prichard, ‘Phys. Hist. of Mankind,’ 1851, vol. i. p. 224.Return to passage

[17] G. Lewis’s ‘Journal of Residence in West Indies,’ ‘Home and Col. Library,’ p. 100.Return to passage

[18] Sidney’s edition of Youatt on the Pig, p. 24. I have given analogous facts in the case of mankind in my ‘Descent of Man,’ 2nd edit., p. 195.Return to passage

[19] ‘Journal of Horticulture,’ 1862, pp. 476, 498; 1865, p. 460. With respect to the heartsease, ‘Gardener’s Chronicle,’ 1863, p. 628.Return to passage

[20] ‘Des Jacinthes, de leur Culture,’ 1768, p. 53: on wheat ‘Gardener’s Chronicle,’ 1846, p. 653.Return to passage

[21] W. B. Tegetmeier, ‘The Field,’ Feb. 25, 1865. With respect to black fowls, see a quotation in Thompson’s ‘Nat. Hist. of Ireland,’ 1849, vol. i. p. 22.Return to passage

[22] ‘In Sachen Darwin’s contra Wigand,’ 1874, p. 70.Return to passage

[23] ‘Bull. de la Soc. d’Acclimat.,’ tom. vii. 1860, p. 359.Return to passage

[24] ‘Transact. Hort. Soc.,’ vol. i. 2nd series, 1835, p. 275. For raspberries, see ‘Gardener’s Chronicle,’ 1855, p. 154, and 1863 p. 245.Return to passage

[25] ‘Gardener’s Chronicle,’ 1843, p. 806.Return to passage

[26] Ibid., 1850, p. 732.Return to passage

[27] Ibid., 1860, p. 956.Return to passage

[28] J. De Jonghe, in ‘Gardener’s Chronicle,’ 1860, p. 120.Return to passage

[29] Downing, ‘Fruit-trees of North America,’ pp. 266, 501: in regard to the cherry, p. 198.Return to passage

[30] ‘Gardener’s Chronicle,’ 1849, p. 755.Return to passage

[31] ‘Journal of Horticulture,’ Sept. 26th, 1865, p. 254; see other references given in chap. x.Return to passage

[32] Mr. Selby, in ‘Mag. of Zoology and Botany,’ Edinburgh, vol. ii. 1838, p. 393.Return to passage

[33] The Reine Claude de Bavay, ‘Journal of Horticulture,’ Dec. 27, 1864, p. 511.Return to passage

[34] Mr. Pusey, in ‘Journal of R. Agricult. Soc.,’ vol. vi. p. 179. For Swedish turnips, see ‘Gardener’s Chronicle,’ 1847, p. 91.Return to passage

[35] Godron, ‘De l’Espèce,’ tom. ii. p. 98.Return to passage

[36] ‘Gardener’s Chronicle,’ 1866, p. 732.Return to passage

[37] ‘Gardener’s Chronicle,’ 1862, pp. 820, 821.Return to passage

[38] ‘On the Varieties of Wheat,’ p. 59.Return to passage

[39] Mr. Hewitt and others, in ‘Journal of Hort.,’ 1862, p. 773.Return to passage

[40] ‘Encyclop. of Rural Sports,’ p. 405.Return to passage

[41] Col. Le Couteur, ‘Journal Roy. Agricult. Soc.,’ vol. iv. p. 43.Return to passage

[42] Malingié-Nouel, ‘Journal R. Agricult. Soc.,’ vol. xiv. 1853, pp. 215, 217.Return to passage

[43] ‘Gardener’s Chronicle,’ 1845, p. 273.Return to passage

[44] ‘Journal of Horticulture,’ 1862, p. 157.Return to passage

[45] ‘Cottage Gardener,’ 1860, p. 368.Return to passage

[46] ‘A Review of Reports,’ 1808, p. 406.Return to passage

[47] ‘Gardener’s Chronicle,’ 1853, p. 45.Return to passage

[48] Isidore Geoffroy Saint-Hilaire, ‘Hist. Nat. Gen.,’ tom. iii. p. 49. ‘On the Cochineal Insect,’ p. 46.Return to passage

[49] ‘Die Darwin’sche Theorie und das Migrationsgesetz der Organismen,’ 1868, p. 19.Return to passage

[50] Capt. Marryat, quoted by Blyth in ‘Journ. Asiatic Soc. of Bengal,’ vol. xxviii. p. 229.Return to passage

[51] Mr. Oxley, ‘Journal of the Indian Archipelago,’ vol. ii. 1848, p. 645.Return to passage

[52] Mr. Abbey, ‘Journal of Horticulture,’ Dec. 1, 1863, p. 430.Return to passage

[53] ‘On Naval Timber,’ 1831, p. 107.Return to passage

[54] Mr. Baily, in ‘The Poultry Chronicle,’ vol. ii. 1854, p. 150. Also vol. i. p. 342; vol. iii. p. 245.Return to passage

[55] ‘Cottage Gardener,’ 1855, December, p. 171; 1856, January, pp. 248, 323.Return to passage

[56] ‘Ueber Shorthorn Rindvieh,’ 1857, s. 51.Return to passage

[57] ‘The Veterinary,’ vol. xiii. p. 720. For the Glamorganshire cattle, see Youatt on Cattle, p. 51.Return to passage

[58] J. M. Eaton, ‘A Treatise on Fancy Pigeons,’ p. 82; Ferguson, on ‘Rare and Prize Poultry,’ p. 162; Mr. Brent, in ‘Cottage Gardener,’ Oct. 1860, p. 13.Return to passage

[59] ‘Die Racen des Schweines,’ 1860, s. 48.Return to passage

[60] See some good remarks on this head by M. de Quatrefages, ‘Unité de l’Espèce Humaine,’ 1861, p. 119.Return to passage

[61] Verlot, ‘Des Variétés,’ 1865, p. 94.Return to passage

[62] Mr. Patrick Shirreff, in ‘Gardener’s Chronicle,’ 1858, p. 771.Return to passage

[63] ‘Pomologie Physiolog.,’ 1830, p. 106.Return to passage

[64] Youatt on Sheep, p. 521.Return to passage

[65] See also Stonehenge, ‘British Rural Sports,’ edition of 1871, p. 384.Return to passage

[66] ‘A Treatise on the Almond Tumbler,’ p. 1.Return to passage

[67] M. J. de Jonghe, in ‘Gardener’s Chronicle,’ 1858, p. 173.Return to passage

[68] ‘Contributions to the Theory of Natural Selection,’ 2nd edit., 1871, p. 292.Return to passage

[69] Max Müller, ‘Science of Language,’ 1861, p. 223.Return to passage

[70] Youatt on Cattle, pp. 116, 128.Return to passage

[71] ‘Domesticated Animals,’ p. 188.Return to passage

[72] Volz, ‘Beiträge zur Kulturgeschichte,’ 1852, s. 99 et passim.Return to passage

[73] Blaine, ‘Encyclop. of Rural Sports,’ p. 213.Return to passage

XXII. CAUSES OF VARIABILITY

VARIABILITY DOES NOT NECESSARILY ACCOMPANY REPRODUCTION—CAUSES ASSIGNED BY VARIOUS AUTHORS—INDIVIDUAL DIFFERENCES—VARIABILITY OF EVERY KIND DUE TO CHANGED CONDITIONS OF LIFE—ON THE NATURE OF SUCH CHANGES—CLIMATE, FOOD, EXCESS OF NUTRIMENT—SLIGHT CHANGES SUFFICIENT—EFFECTS OF GRAFTING ON THE VARIABILITY OF SEEDLING-TREES—DOMESTIC PRODUCTIONS BECOME HABITUATED TO CHANGED CONDITIONS—ON THE ACCUMULATIVE ACTION OF CHANGED CONDITIONS—CLOSE INTERBREEDING AND THE IMAGINATION OF THE MOTHER SUPPOSED TO CAUSE VARIABILITY—CROSSING AS A CAUSE OF THE APPEARANCE OF NEW CHARACTERS—VARIABILITY FROM THE COMMINGLING OF CHARACTERS AND FROM REVERSION—ON THE MANNER AND PERIOD OF ACTION OF THE CAUSES WHICH EITHER DIRECTLY, OR INDIRECTLY THROUGH THE REPRODUCTIVE SYSTEM, INDUCE VARIABILITY.

We will now consider, as far as we can, the causes of the almost universal variability of our domesticated productions. The subject is an obscure one; but it may be useful to probe our ignorance. Some authors, for instance Dr. Prosper Lucas, look at variability as a necessary contingent on reproduction, and as much an aboriginal law as growth or inheritance. Others have of late encouraged, perhaps unintentionally, this view by speaking of inheritance and variability as equal and antagonistic principles. Pallas maintained, and he has had some followers, that variability depends exclusively on the crossing of primordially distinct forms. Other authors attribute variability to an excess of food, and with animals to an excess relatively to the amount of exercise taken, or again to the effects of a more genial climate. That these causes are all effective is highly probable. But we must, I think, take a broader view, and conclude that organic beings, when subjected during several generations to any change whatever in their conditions, tend to vary; the kind of variation which ensues depending in most cases in a far higher degree on the nature or constitution of the being, than on the nature of the changed conditions.

Those authors who believe that it is a law of nature that each individual should differ in some slight degree from every other, may maintain, apparently with truth, that this is the fact, not only with all domesticated animals and cultivated plants, but likewise with all organic beings in a state of nature. The Laplander by long practice knows and gives a name to each reindeer, though, as Linnæus remarks, “to distinguish one from another among such multitudes was beyond my comprehension, for they were like ants on an anthill.” In Germany shepherds have won wagers by recognising each sheep in a flock of a hundred, which they had never seen until the previous fortnight. This power of discrimination, however, is as nothing compared to that which some florists have acquired. Verlot mentions a gardener who could distinguish 150 kinds of camellia, when not in flower; and it has been positively asserted that the famous old Dutch florist Voorhelm, who kept above 1200 varieties of the hyacinth, was hardly ever deceived in knowing each variety by the bulb alone. Hence we must conclude that the bulbs of the hyacinth and the branches and leaves of the camellia, though appearing to an unpractised eye absolutely undistinguishable, yet really differ.[1]

As Linnæus has compared the reindeer in number to ants, I may add that each ant knows its fellow of the same community. Several times I carried ants of the same species (Formica rufa) from one ant-hill to another, inhabited apparently by tens of thousands of ants; but the strangers were instantly detected and killed. I then put some ants taken from a very large nest into a bottle strongly perfumed with assafœtida, and after an interval of twenty-four hours returned them to their home; they were at first threatened by their fellows, but were soon recognised and allowed to pass. Hence each ant certainly recognised, independently of odour, its fellow; and if all the ants of the same community have not some countersign or watchword, they must present to each other’s senses some distinguishable character.

The dissimilarity of brothers or sisters of the same family, and of seedlings from the same capsule, may be in part accounted for by the unequal blending of the characters of the two parents, and by the more or less complete recovery through reversion of ancestral characters on either side; but we thus only push the difficulty further back in time, for what made the parents or their progenitors different? Hence the belief[2] that an innate tendency to vary exists, independently of external differences, seems at first sight probable. But even the seeds nurtured in the same capsule are not subjected to absolutely uniform conditions, as they draw their nourishment from different points; and we shall see in a future chapter that this difference sometimes suffices to affect the character of the future plant. The greater dissimilarity of the successive children of the same family in comparison with twins, which often resemble each other in external appearance, mental disposition, and constitution, in so extraordinary a manner, apparently proves that the state of the parents at the exact period of conception, or the nature of the subsequent embryonic development, has a direct and powerful influence on the character of the offspring. Nevertheless, when we reflect on the individual differences between organic beings in a state of nature, as shown by every wild animal knowing its mate; and when we reflect on the infinite diversity of the many varieties of our domesticated productions, we may well be inclined to exclaim, though falsely as I believe, that Variability must be looked at as an ultimate fact, necessarily contingent on reproduction.

Those authors who adopt this latter view would probably deny that each separate variation has its own proper exciting cause. Although we can seldom trace the precise relation between cause and effect, yet the considerations presently to be given lead to the conclusion that each modification must have its own distinct cause, and is not the result of what we blindly call accident. The following striking case has been communicated to me by Dr. William Ogle. Two girls, born as twins, and in all respects extremely alike, had their little fingers on both hands crooked; and in both children the second bicuspid tooth of the second dentition on the right side in the upper jaw was misplaced; for, instead of standing in a line with the others, it grew from the roof of the mouth behind the first bicuspid. Neither the parents nor any other members of the family were known to have exhibited any similar peculiarity; but a son of one of these girls had the same tooth similarly misplaced. Now, as both the girls were affected in exactly the same manner, the idea of accident is at once excluded: and we are compelled to admit that there must have existed some precise and sufficient cause which, if it had occurred a hundred times, would have given crooked fingers and misplaced bicuspid teeth to a hundred children. It is of course possible that this case may have been due to reversion to some long-forgotten progenitor, and this would much weaken the value of the argument. I have been led to think of the probability of reversion, from having been told by Mr. Galton of another case of twin girls born with their little fingers slightly crooked, which they inherited from their maternal grandmother.

We will now consider the general arguments, which appear to me to have great weight, in favour of the view that variations of all kinds and degrees are directly or indirectly caused by the conditions of life to which each being, and more especially its ancestors, have been exposed.

No one doubts that domesticated productions are more variable than organic beings which have never been removed from their natural conditions. Monstrosities graduate so insensibly into mere variations that it is impossible to separate them; and all those who have studied monstrosities believe that they are far commoner with domesticated than with wild animals and plants;[3] and in the case of plants, monstrosities would be equally noticeable in the natural as in the cultivated state. Under nature, the individuals of the same species are exposed to nearly uniform conditions, for they are rigorously kept to their proper places by a host of competing animals and plants; they have, also, long been habituated to their conditions of life; but it cannot be said that they are subject to quite uniform conditions, and they are liable to a certain amount of variation. The circumstances under which our domestic productions are reared are widely different: they are protected from competition; they have not only been removed from their natural conditions and often from their native land, but they are frequently carried from district to district, where they are treated differently, so that they rarely remain during any considerable length of time exposed to closely similar conditions. In conformity with this, all our domesticated productions, with the rarest exceptions, vary far more than natural species. The hive-bee, which feeds itself and follows in most respects its natural habits of life, is the least variable of all domesticated animals, and probably the goose is the next least variable; but even the goose varies more than almost any wild bird, so that it cannot be affiliated with perfect certainty to any natural species. Hardly a single plant can be named, which has long been cultivated and propagated by seed, that is not highly variable; common rye (Secale cereale) has afforded fewer and less marked varieties than almost any other cultivated plant;[4] but it may be doubted whether the variations of this, the least valuable of all our cereals, have been closely observed.

Bud-variation, which was fully discussed in a former chapter, shows us that variability may be quite independent of seminal reproduction, and likewise of reversion to long-lost ancestral characters. No one will maintain that the sudden appearance of a moss-rose on a Provence-rose is a return to a former state, for mossiness of the calyx has been observed in no natural species; the same argument is applicable to variegated and laciniated leaves; nor can the appearance of nectarines on peach-trees be accounted for on the principle of reversion. But bud-variations more immediately concern us, as they occur far more frequently on plants which have been highly cultivated during a length of time, than on other and less highly cultivated plants; and very few well-marked instances have been observed with plants growing under strictly natural conditions. I have given one instance of an ash-tree growing in a gentleman’s pleasure-grounds; and occasionally there may be seen, on beech and other trees, twigs leafing at a different period from the other branches. But our forest trees in England can hardly be considered as living under strictly natural conditions; the seedlings are raised and protected in nursery-grounds, and must often be transplanted into places where wild trees of the kind would not naturally grow. It would be esteemed a prodigy if a dog-rose growing in a hedge produced by bud-variation a moss-rose, or a wild bullace or wild cherry-tree yielded a branch bearing fruit of a different shape and colour from the ordinary fruit. The prodigy would be enhanced if these varying branches were found capable of propagation, not only by grafts, but sometimes by seed; yet analogous cases have occurred with many of our highly cultivated trees and herbs.

These several considerations alone render it probable that variability of every kind is directly or indirectly caused by changed conditions of life. Or, to put the case under another point of view, if it were possible to expose all the individuals of a species during many generations to absolutely uniform conditions of life, there would be no variability.

On the Nature of the Changes in the Conditions of Life which induce Variability.

From a remote period to the present day, under climates and circumstances as different as it is possible to conceive, organic beings of all kinds, when domesticated or cultivated, have varied. We see this with the many domestic races of quadrupeds and birds belonging to different orders, with goldfish and silkworms, with plants of many kinds, raised in various quarters of the world. In the deserts of northern Africa the date-palm has yielded thirty-eight varieties; in the fertile plains of India it is notorious how many varieties of rice and of a host of other plants exist; in a single Polynesian island, twenty-four varieties of the bread-fruit, the same number of the banana, and twenty-two varieties of the arum, are cultivated by the natives; the mulberry-tree in India and Europe has yielded many varieties serving as food for the silkworm; and in China sixty-three varieties of the bamboo are used for various domestic purposes.[5] These facts, and innumerable others which could be added, indicate that a change of almost any kind in the conditions of life suffices to cause variability—different changes acting on different organisms.

Andrew Knight[6] attributed the variation of both animals and plants to a more abundant supply of nourishment, or to a more favourable climate, than that natural to the species. A more genial climate, however, is far from necessary; the kidney-bean, which is often injured by our spring frosts, and peaches, which require the protection of a wall, have varied much in England, as has the orange-tree in northern Italy, where it is barely able to exist.[7] Nor can we overlook the fact, though not immediately connected with our present subject, that the plants and shells of the Arctic regions are eminently variable.[8] Moreover, it does not appear that a change of climate, whether more or less genial, is one of the most potent causes of variability; for in regard to plants Alph. De Candolle, in his ‘Géographie Botanique’ repeatedly shows that the native country of a plant, where in most cases it has been longest cultivated, is that where it has yielded the greatest number of varieties.

It is doubtful whether a change in the nature of the food is a potent cause of variability. Scarcely any domesticated animal has varied more than the pigeon or the fowl, but their food, especially that of highly-bred pigeons, is generally the same. Nor can our cattle and sheep have been subjected to any great change in this respect. But in all these cases the food probably is much less varied in kind than that which was consumed by the species in its natural state.[9]

Of all the causes which induce variability, excess of food, whether or not changed in nature, is probably the most powerful. This view was held with regard to plants by Andrew Knight, and is now held by Schleiden, more especially in reference to the inorganic elements of the food.[10] In order to give a plant more food it suffices in most cases to grow it separately, and thus prevent other plants robbing its roots. It is surprising, as I have often seen, how vigorously our common wild species flourish when planted by themselves, though not in highly manured land; separate growth is, in fact, the first step in cultivation. We see the converse of the belief that excess of food induces variability in the following statement by a great raiser of seeds of all kinds:[11] “It is a rule invariably with us, when we desire to keep a true stock of any one kind of seed, to grow it on poor land without dung; but when we grow for quantity, we act contrary, and sometimes have dearly to repent of it.” According also to Carrière, who has had great experience with flower-garden seeds, “On remarque en général les plantes de vigeur moyenne sont celles qui conservent le mieux leurs caractères.”

In the case of animals the want of a proper amount of exercise, as Bechstein remarked, has perhaps played, independently of the direct effects of the disuse of any particular organ, an important part in causing variability. We can see in a vague manner that, when the organised and nutrient fluids of the body are not used during growth, or by the wear and tear of the tissues, they will be in excess; and as growth, nutrition, and reproduction are intimately allied processes, this superfluity might disturb the due and proper action of the reproductive organs, and consequently affect the character of the future offspring. But it may be argued that neither an excess of food nor a superfluity in the organised fluids of the body necessarily induces variability. The goose and the turkey have been well fed for many generations, yet have varied very little. Our fruit-trees and culinary plants, which are so variable, have been cultivated from an ancient period, and, though they probably still receive more nutriment than in their natural state, yet they must have received during many generations nearly the same amount; and it might be thought that they would have become habituated to the excess. Nevertheless, on the whole, Knight’s view, that excess of food is one of the most potent causes of variability, appears, as far as I can judge, probable.

Whether or not our various cultivated plants have received nutriment in excess, all have been exposed to changes of various kinds. Fruit-trees are grafted on different stocks, and grown in various soils. The seeds of culinary and agricultural plants are carried from place to place; and during the last century the rotation of our crops and the manures used have been greatly changed.

Slight changes of treatment often suffice to induce variability. The simple fact of almost all our cultivated plants and domesticated animals having varied in all places and at all times, leads to this conclusion. Seeds taken from common English forest-trees, grown under their native climate, not highly manured or otherwise artificially treated, yield seedlings which vary much, as may be seen in every extensive seed-bed. I have shown in a former chapter what a number of well-marked and singular varieties the thorn (Cratægus oxycantha) has produced: yet this tree has been subjected to hardly any cultivation. In Staffordshire I carefully examined a large number of two British plants, namely Geranium phæum and pyrenaicum, which have never been highly cultivated. These plants had spread spontaneously by seed from a common garden into an open plantation; and the seedlings varied in almost every single character, both in their flower and foliage, to a degree which I have never seen exceeded; yet they could not have been exposed to any great change in their conditions.

With respect to animals, Azara has remarked with much surprise[12] that, whilst the feral horses on the Pampas are always of one of three colours, and the cattle always of a uniform colour, yet these animals, when bred on the unenclosed estancias, though kept in a state which can hardly be called domesticated, and apparently exposed to almost identically the same conditions as when they are feral, nevertheless display a great diversity of colour. So again in India several species of fresh-water fish are only so far treated artificially, that they are reared in great tanks; but this small change is sufficient to induce much variability.[13]

Some facts on the effects of grafting, in regard to the variability of trees, deserve attention. Cabanis asserts that when certain pears are grafted on the quince, their seeds yield a greater number of varieties than do the seeds of the same variety of pear when grafted on the wild pear.[14] But as the pear and quince are distinct species, though so closely related that the one can be readily grafted and succeeds admirably on the other, the fact of variability being thus caused is not surprising; as we are here enabled to see the cause, namely, the very different nature of the stock and graft. Several North American varieties of the plum and peach are well known to reproduce themselves truly by seed; but Downing asserts,[15] “that when a graft is taken from one of these trees and placed upon another stock, this grafted tree is found to lose its singular property of producing the same variety by seed, and becomes like all other worked trees;”—that is, its seedlings become highly variable. Another case is worth giving: the Lalande variety of the walnut-tree leafs between April 20th and May 15th, and its seedlings invariably inherit the same habit; whilst several other varieties of the walnut leaf in June. Now, if seedlings are raised from the May-leafing Lalande variety, grafted on another May-leafing variety, though both stock and graft have the same early habit of leafing, yet the seedlings leaf at various times, even as late as the 5th of June.[16] Such facts as these are well fitted to show on what obscure and slight causes variability depends.

I may here just allude to the appearance of new and valuable varieties of fruit-trees and of wheat in woods and waste places, which at first sight seems a most anomalous circumstance. In France a considerable number of the best pears have been discovered in woods; and this has occurred so frequently, that Poiteau asserts that “improved varieties of our cultivated fruits rarely originate with nurserymen.”[17] In England, on the other hand, no instance of a good pear having been found wild has been recorded; and Mr. Rivers informs me that he knows of only one instance with apples, namely, the Bess Poole, which was discovered in a wood in Nottinghamshire. This difference between the two countries may be in part accounted for by the more favourable climate of France, but chiefly from the great number of seedlings which spring up there in the woods. I infer that this is the case from a remark made by a French gardener,[18] who regards it as a national calamity that such a number of pear-trees are periodically cut down for firewood, before they have borne fruit. The new varieties which thus spring up in the woods, though they cannot have received any excess of nutriment, will have been exposed to abruptly changed conditions, but whether this is the cause of their production is very doubtful. These varieties, however, are probably all descended[19] from old cultivated kinds growing in adjoining orchards— a circumstance which will account for their variability; and out of a vast number of varying trees there will always be a good chance of the appearance of a valuable kind. In North America, where fruit-trees frequently spring up in waste places, the Washington pear was found in a hedge, and the Emperor peach in a wood.[20]

With respect to wheat, some writers have spoken[21] as if it were an ordinary event for new varieties to be found in waste places; the Fenton wheat was certainly discovered growing on a pile of basaltic detritus in a quarry, but in such a situation the plant would probably receive a sufficient amount of nutriment. The Chidham wheat was raised from an ear found on a hedge; and Hunter’s wheat was discovered by the roadside in Scotland, but it is not said that this latter variety grew where it was found.[22]

Whether our domestic productions would ever become so completely habituated to the conditions under which they now live, as to cease varying, we have no sufficient means for judging. But, in fact, our domestic productions are never exposed for a great length of time to uniform conditions, and it is certain that our most anciently cultivated plants, as well as animals, still go on varying, for all have recently undergone marked improvement. In some few cases, however, plants have become habituated to new conditions. Thus, Metzger, who cultivated in Germany during many years numerous varieties of wheat, brought from different countries,[23] states that some kinds were at first extremely variable, but gradually, in one instance after an interval of twenty-five years, became constant; and it does not appear that this resulted from the selection of the more constant forms.

On the Accumulative Action of changed Conditions of Life.—We have good grounds for believing that the influence of changed conditions accumulates, so that no effect is produced on a species until it has been exposed during several generations to continued cultivation or domestication. Universal experience shows us that when new flowers are first introduced into our gardens they do not vary; but ultimately all, with the rarest exceptions, vary to a greater or less extent. In a few cases the requisite number of generations, as well as the successive steps in the progress of variation, have been recorded, as in the often quoted instance of the Dahlia.[24] After several years’ culture the Zinnia has only lately (1860) begun to vary in any great degree. “In the first seven or eight years of high cultivation, the Swan River daisy (Brachycome iberidifolia) kept to its original colour; it then varied into lilac and purple and other minor shades.”[25] Analogous facts have been recorded with the Scotch rose. In discussing the variability of plants several experienced horticulturists have spoken to the same general effect. Mr. Salter[26] remarks, “Every one knows that the chief difficulty is in breaking through the original form and colour of the species, and every one will be on the look-out for any natural sport, either from seed or branch; that being once obtained, however trifling the change may be, the result depends upon himself.” M. de Jonghe, who has had so much success in raising new varieties of pears and strawberries,[27] remarks with respect to the former, “There is another principle, namely, that the more a type has entered into a state of variation, the greater is its tendency to continue doing so; and the more it has varied from the original type, the more it is disposed to vary still farther.” We have, indeed, already discussed this latter point when treating of the power which man possesses, through selection, of continually augmenting in the same direction each modification; for this power depends on continued variability of the same general kind. The most celebrated horticulturist in France, namely, Vilmorin,[28] even maintains that, when any particular variation is desired, the first step is to get the plant to vary in any manner whatever, and to go on selecting the most variable individuals, even though they vary in the wrong direction; for the fixed character of the species being once broken, the desired variation will sooner or later appear.

As nearly all our animals were domesticated at an extremely remote epoch, we cannot, of course, say whether they varied quickly or slowly when first subjected to new conditions. But Dr. Bachman[29] states that he has seen turkeys raised from the eggs of the wild species lose their metallic tints and become spotted with white in the third generation. Mr. Yarrell many years ago informed me that the wild ducks bred on the ponds in St. James’s Park, which had never been crossed, as it is believed, with domestic ducks, lost their true plumage after a few generations. An excellent observer,[30] who has often reared ducks from the eggs of the wild bird, and who took precautions that there should be no crossing with domestic breeds, has given, as previously stated, full details on the changes which they gradually undergo. He found that he could not breed these wild ducks true for more than five or six generations, “as they then proved so much less beautiful. The white collar round the neck of the mallard became much broader and more irregular, and white feathers appeared in the ducklings’ wings.” They increased also in size of body; their legs became less fine, and they lost their elegant carriage. Fresh eggs were then procured from wild birds; but again the same result followed. In these cases of the duck and turkey we see that animals, like plants, do not depart from their primitive type until they have been subjected during several generations to domestication. On the other hand, Mr. Yarrell informed me that the Australian dingos, bred in the Zoological Gardens, almost invariably produced in the first generation puppies marked with white and other colours; but, these introduced dingos had probably been procured from the natives, who keep them in a semi-domesticated state. It is certainly a remarkable fact that changed conditions should at first produce, as far as we can see, absolutely no effect; but that they should subsequently cause the character of the species to change. In the chapter on pangenesis I shall attempt to throw a little light on this fact.

Returning now to the causes which are supposed to induce variability. Some authors[31] believe that close interbreeding gives this tendency, and leads to the production of monstrosities. In the seventeenth chapter some few facts were advanced, showing that monstrosities are, as it appears, occasionally thus induced; and there can be no doubt that close interbreeding causes lessened fertility and a weakened constitution; hence it may lead to variability: but I have not sufficient evidence on this head. On the other hand, close interbreeding, if not carried to an injurious extreme, far from causing variability, tends to fix the character of each breed.

It was formerly a common belief, still held by some persons, that the imagination of the mother affects the child in the womb.[32] This view is evidently not applicable to the lower animals, which lay unimpregnated eggs, or to plants. Dr. William Hunter, in the last century, told my father that during many years every woman in a large London Lying-in Hospital was asked before her confinement whether anything had specially affected her mind, and the answer was written down; and it so happened that in no one instance could a coincidence be detected between the woman’s answer and any abnormal structure; but when she knew the nature of the structure, she frequently suggested some fresh cause. The belief in the power of the mother’s imagination may perhaps have arisen from the children of a second marriage resembling the previous father, as certainly sometimes occurs, in accordance with the facts given in the eleventh chapter.

Crossing as a Cause of Variability.—In an early part of this chapter it was stated that Pallas[33] and a few other naturalists maintain that variability is wholly due to crossing. If this means that new characters never spontaneously appear in our domestic races, but that they are all directly derived from certain aboriginal species, the doctrine is little less than absurd; for it implies that animals like Italian greyhounds, pug-dogs, bull-dogs, pouter and fantail pigeons, etc., were able to exist in a state of nature. But the doctrine may mean something widely different, namely, that the crossing of distinct species is the sole cause of the first appearance of new characters, and that without this aid man could not have formed his various breeds. As, however, new characters have appeared in certain cases by bud-variation, we may conclude with certainty that crossing is not necessary for variability. It is, moreover, certain that the breeds of various animals, such as of the rabbit, pigeon, duck, etc., and the varieties of several plants, are the modified descendants of a single wild species. Nevertheless, it is probable that the crossing of two forms, when one or both have long been domesticated or cultivated, adds to the variability of the offspring, independently of the commingling of the characters derived from the two parent-forms; and this implies that new characters actually arise. But we must not forget the facts advanced in the thirteenth chapter, which clearly prove that the act of crossing often leads to the reappearance or reversion of long-lost characters; and in most cases it would be impossible to distinguish between the reappearance of ancient characters and the first appearance of absolutely new characters. Practically, whether new or old, they would be new to the breed in which they reappeared.

Gärtner declares,[34] and his experience is of the highest value on such a point, that, when he crossed native plants which had not been cultivated, he never once saw in the offspring any new character; but that from the odd manner in which the characters derived from the parents were combined, they sometimes appeared as if new. When, on the other hand, he crossed cultivated plants, he admits that new characters occasionally appeared, but he is strongly inclined to attribute their appearance to ordinary variability, not in any way to the cross. An opposite conclusion, however, appears to me the more probable. According to Kölreuter, hybrids in the genus Mirabilis vary almost infinitely, and he describes new and singular characters in the form of the seeds, in the colour of the anthers, in the cotyledons being of immense size, in new and highly peculiar odours, in the flowers expanding early in the season, and in their closing at night. With respect to one lot of these hybrids, he remarks that they presented characters exactly the reverse of what might have been expected from their parentage.[35]

Prof. Lecoq[36] speaks strongly to the same effect in regard to this same genus, and asserts that many of the hybrids from Mirabilis jalapa and multiflora might easily be mistaken for distinct species, and adds that they differed in a greater degree than the other species of the genus, from M. jalapa. Herbert, also, has described[37] certain hybrid Rhododendrons as being “as unlike all others in foliage, as if they had been a separate species.” The common experience of floriculturists proves that the crossing and recrossing of distinct but allied plants, such as the species of Petunia, Calceolaria, Fuchsia, Verbena, etc., induces excessive variability; hence the appearance of quite new characters is probable. M. Carrière[38] has lately discussed this subject: he states that Erythrina cristagalli had been multiplied by seed for many years, but had not yielded any varieties: it was then crossed with the allied E. herbacea, and “the resistance was now overcome, and varieties were produced with flowers of extremely different size, form, and colour.”

From the general and apparently well-founded belief that the crossing of distinct species, besides commingling their characters, adds greatly to their variability, it has probably arisen that some botanists have gone so far as to maintain[39] that, when a genus includes only a single species, this when cultivated never varies. The proposition made so broadly cannot be admitted; but it is probably true that the variability of monotypic genera when cultivated is generally less than that of genera including numerous species, and this quite independently of the effects of crossing. I have shown in my ‘Origin of Species’ that the species belonging to small genera generally yield a less number of varieties in a state of nature than those belonging to large genera. Hence the species of small genera would, it is probable, produce fewer varieties under cultivation than the already variable species of larger genera.

Although we have not at present sufficient evidence that the crossing of species, which have never been cultivated, leads to the appearance of new characters, this apparently does occur with species which have been already rendered in some degree variable through cultivation. Hence crossing, like any other change in the conditions of life, seems to be an element, probably a potent one, in causing variability. But we seldom have the means of distinguishing, as previously remarked, between the appearance of really new characters and the reappearance of long-lost characters, evoked through the act of crossing. I will give an instance of the difficulty in distinguishing such cases. The species of Datura may be divided into two sections, those having white flowers with green stems, and those having purple flowers with brown stems: now Naudin[40] crossed Datura lævis and ferox, both of which belong to the white section, and raised from them 205 hybrids. Of these hybrids, every one had brown stems and bore purple flowers; so that they resembled the species of the other section of the genus, and not their own two parents. Naudin was so much astonished at this fact, that he was led carefully to observe both parent-species, and he discovered that the pure seedlings of D. ferox, immediately after germination, had dark purple stems, extending from the young roots up to the cotyledons, and that this tint remained ever afterwards as a ring round the base of the stem of the plant when old. Now I have shown in the thirteenth chapter that the retention or exaggeration of an early character is so intimately related to reversion, that it evidently comes under the same principle. Hence probably we ought to look at the purple flowers and brown stems of these hybrids, not as new characters due to variability, but as a return to the former state of some ancient progenitor.

Independently of the appearance of new characters from crossing, a few words may be added to what has been said in former chapters on the unequal combination and transmission of the characters proper to the two parent-forms. When two species or races are crossed, the offspring of the first generation are generally uniform, but those subsequently produced display an almost infinite diversity of character. He who wishes, says Kölreuter,[41] to obtain an endless number of varieties from hybrids should cross and recross them. There is also much variability when hybrids or mongrels are reduced or absorbed by repeated crosses with either pure parent-form: and a still higher degree of variability when three distinct species, and most of all when four species, are blended together by successive crosses. Beyond this point Gärtner,[42] on whose authority the foregoing statements are made, never succeeded in effecting a union; but Max Wichura[43] united six distinct species of willows into a single hybrid. The sex of the parent species affects in an inexplicable manner the degree of variability of hybrids; for Gärtner[44] repeatedly found that when a hybrid was used as a father and either one of the pure parent-species, or a third species, was used as the mother, the offspring were more variable than when the same hybrid was used as the mother, and either pure parent or the same third species as the father: thus seedlings from Dianthus barbatus crossed by the hybrid D. chinensi-barbatus were more variable than those raised from this latter hybrid fertilised by the pure D. barbatus. Max Wichura[45] insists strongly on an analogous result with his hybrid willows. Again Gärtner[46] asserts that the degree of variability sometimes differs in hybrids raised from reciprocal crosses between the same two species; and here the sole difference is, that the one species is first used as the father and then as the mother. On the whole we see that, independently of the appearance of new characters, the variability of successive crossed generations is extremely complex, partly from the offspring partaking unequally of the characters of the two parent-forms, and more especially from their unequal tendency to revert to such characters or to those of more ancient progenitors.

On the Manner and on the Period of Action of the Causes which induce Variability.—This is an extremely obscure subject, and we need here only consider, whether inherited variations are due to certain parts being acted on after they have been formed, or through the reproductive system being affected before their formation; and in the former case at what period of growth or development the effect is produced. We shall see in the two following chapters that various agencies, such as an abundant supply of food, exposure to a different climate, increased use or disuse of parts, etc., prolonged during several generations, certainly modify either the whole organisation or certain organs; and it is clear at least in the case of bud-variation that the action cannot have been through the reproductive system.

With respect to the part which the reproductive system takes in causing variability, we have seen in the eighteenth chapter that even slight changes in the conditions of life have a remarkable power in causing a greater or less degree of sterility. Hence it seems not improbable that beings generated through a system so easily affected should themselves be affected, or should fail to inherit, or inherit in excess, characters proper to their parents. We know that certain groups of organic beings, but with exceptions in each group, have their reproductive systems much more easily affected by changed conditions than other groups; for instance, carnivorous birds, more readily than carnivorous mammals, and parrots more readily than pigeons; and this fact harmonises with the apparently capricious manner and degree in which various groups of animals and plants vary under domestication.

Kölreuter[47] was struck with the parallelism between the excessive variability of hybrids when crossed and recrossed in various ways,—these hybrids having their reproductive powers more or less affected,—and the variability of anciently cultivated plants. Max Wichura[48] has gone one step farther, and shows that with many of our highly cultivated plants, such as the hyacinth, tulip, auricula, snapdragon, potato, cabbage, etc., which there is no reason to believe have been hybridised, the anthers contain many irregular pollen-grains in the same state as in hybrids. He finds also in certain wild forms, the same coincidence between the state of the pollen and a high degree of variability, as in many species of Rubus; but in R. caesius and idaeus, which are not highly variable species, the pollen is sound. It is also notorious that many cultivated plants, such as the banana, pineapple, bread-fruit, and others previously mentioned, have their reproductive organs so seriously affected as to be generally quite sterile; and when they do yield seed, the seedlings, judging from the large number of cultivated races which exist, must be variable in an extreme degree. These facts indicate that there is some relation between the state of the reproductive organs and a tendency to variability; but we must not conclude that the relation is strict. Although many of our highly cultivated plants may have their pollen in a deteriorated condition, yet, as we have previously seen, they yield more seeds, and our anciently domesticated animals are more prolific, than the corresponding species in a state of nature. The peacock is almost the only bird which is believed to be less fertile under domestication than in its native state, and it has varied in a remarkably small degree. From these considerations it would seem that changes in the conditions of life lead either to sterility or to variability, or to both; and not that sterility induces variability. On the whole it is probable that any cause affecting the organs of reproduction would likewise affect their product,—that is, the offspring thus generated.

The period of life at which the causes that induce variability act, is likewise an obscure subject, which has been discussed by various authors.[49] In some of the cases, to be given in the following chapter, of modifications from the direct action of changed conditions, which are inherited, there can be no doubt that the causes have acted on the mature or nearly mature animal. On the other hand, monstrosities, which cannot be distinctly separated from lesser variations, are often caused by the embryo being injured whilst in the mother’s womb or in the egg. Thus I. Geoffroy Saint-Hilaire[50] asserts that poor women who work hard during their pregnancy, and the mothers of illegitimate children troubled in their minds and forced to conceal their state, are far more liable to give birth to monsters than women in easy circumstances. The eggs of the fowl when placed upright or otherwise treated unnaturally frequently produce monstrous chickens. It would, however, appear that complex monstrosities are induced more frequently during a rather late than during a very early period of embryonic life; but this may partly result from some one part, which has been injured during an early period, affecting by its abnormal growth other parts subsequently developed; and this would be less likely to occur with parts injured at a later period.[51] When any part or organ becomes monstrous through abortion, a rudiment is generally left, and this likewise indicates that its development had already commenced.

Insects sometimes have their antennae or legs in a monstrous condition, the larvae of which do not possess either antennae or legs; and in these cases, as Quatrefages[52] believes, we are enabled to see the precise period at which the normal progress of development was troubled. But the nature of the food given to a caterpillar sometimes affects the colours of the moth, without the caterpillar itself being affected; therefore it seems possible that other characters in the mature insect might be indirectly modified through the larvae. There is no reason to suppose that organs which have been rendered monstrous have always been acted on during their development; the cause may have acted on the organisation at a much earlier stage. It is even probable that either the male or female sexual elements, or both, before their union, may be affected in such a manner as to lead to modifications in organs developed at a late period of life; in nearly the same manner as a child may inherit from his father a disease which does not appear until old age.

In accordance with the facts above given, which prove that in many cases a close relation exists between variability and the sterility following from changed conditions, we may conclude that the exciting cause often acts at the earliest possible period, namely, on the sexual elements, before impregnation has taken place. That an affection of the female sexual element may induce variability we may likewise infer as probable from the occurrence of bud-variations; for a bud seems to be the analogue of an ovule. But the male element is apparently much oftener affected by changed conditions, at least in a visible manner, than the female element or ovule and we know from Gärtner’s and Wichura’s statements that a hybrid used as the father and crossed with a pure species gives a greater degree of variability to the offspring, than does the same hybrid when used as the mother. Lastly, it is certain that variability may be transmitted through either sexual element, whether or not originally excited in them, for Kölreuter and Gärtner[53] found that when two species were crossed, if either one was variable, the offspring were rendered variable.

Summary.—From the facts given in this chapter, we may conclude that the variability of organic beings under domestication, although so general, is not an inevitable contingent on life, but results from the conditions to which the parents have been exposed. Changes of any kind in the conditions of life, even extremely slight changes, often suffice to cause variability. Excess of nutriment is perhaps the most efficient single exciting cause. Animals and plants continue to be variable for an immense period after their first domestication; but the conditions to which they are exposed never long remain quite constant. In the course of time they can be habituated to certain changes, so as to become less variable; and it is possible that when first domesticated they may have been even more variable than at present. There is good evidence that the power of changed conditions accumulates; so that two, three, or more generations must be exposed to new conditions before any effect is visible. The crossing of distinct forms, which have already become variable, increases in the offspring the tendency to further variability, by the unequal commingling of the characters of the two parents, by the reappearance of long-lost characters, and by the appearance of absolutely new characters. Some variations are induced by the direct action of the surrounding conditions on the whole organisation, or on certain parts alone; other variations appear to be induced indirectly through the reproductive system being affected, as we know is often the case with various beings, which when removed from their natural conditions become sterile. The causes which induce variability act on the mature organism, on the embryo, and, probably, on the sexual elements before impregnation has been effected.

REFERENCES

[1] ‘Des Jacinthes,’ etc., Amsterdam, 1768, p. 43; Verlot, ‘Des Variétés,’ etc., p. 86. On the reindeer see Linnæus, ‘Tour in Lapland,’ translated by Sir J. E. Smith, vol. i. p. 314. The statement in regard to German shepherds is given on the authority of Dr. Weinland.Return to passage

[2] Müller’s ‘Physiology,’ Eng. translation, vol. ii. p. 1662. With respect to the similarity of twins in constitution, Dr. William Ogle has given me the following extract from Professor Trousseau’s Lectures (‘Clinique Médicale,’ tom. i.1 p. 523), in which a curious case is recorded:—“J’ai donné mes soins à deux frères jumeaux, tous deux si extraordinairement ressemblants qu’il m’était impossible de les reconnaitre, à moin de les voir l’un à côté de l’autre. Cette ressemblance physique s’étendait plus loin: ils avaient, permettez-moi l’expression, une similitude pathologique plus remarquable encore. Ainsi l’un d’eux que je voyais aux néothermes à Paris malade d’une ophthalmie rhumatismale me disait, ‘En ce moment mon frere doit avoir une ophthalmie comme la mienne;’ et comme je m’etais recrie, il me montrait quelques jours apres une lettre qu’il venait de recevoir de ce frère alors à Vienne, et qui lui écrivait en effet—‘J’ai mon ophthalmie, tu dois avoir la tienne.’ Quelque singulier que ceci puisse paraître, le fait n’en est pas moins exact: on ne me l’a pas raconté, je l’ai vu, et j’en ai vu d’autres analogues dans ma pratique. Ces deux jumeaux étaient aussi tous deux asthmatiques, et asthmatiques a un effroyable degré. Originaires de Marseille, ils n’ont jamais pu demeurer dans cette ville, ou leurs intérêts les appelaient souvent, sans etre pris de leurs acces; jamais ils n’en eprouvaient a Paris. Bien mieux, il leur suffisait de gagner Toulon pour être guéris de leurs attaques de Marseille. Voyageant sans cesse et dans tous pays pour leurs affaires, ils avaient remarque que certaines localités leur étaient funestes, que dans d’autres ils etaient exempts de tout phénomène d’oppression.”Return to passage

[3] Isid. Geoffroy St.-Hilaire, ‘Hist. des Anomalies,’ tom. iii. p. 352; Moquin-Tandon, ‘Tératologie Végétale,’ 1841, p. 115.Return to passage

[4] Metzger, ‘Die Getreidarten,’ 1841, s. 39.Return to passage

[5] On the date-palm see Vogel, ‘Annals and Mag. of Nat. Hist.,’ 1854, p. 460. On Indian varieties, Dr. F. Hamilton, ‘Transact. Linn. Soc.,’ vol. xiv. p. 296. On the varieties cultivated in Tahiti, see Dr. Bennett, in Loudon’s ‘Mag. of N. Hist.,’ vol. v. 1832, p. 484. Also Ellis, ‘Polynesian Researches,’ vol. i. pp. 370, 375. On twenty varieties of the Pandanus and other trees in the Marianne Island, see ‘Hooker’s Miscellany,’ vol. i. p. 308. On the bamboo in China, see Huc’s ‘Chinese Empire,’ vol. ii. p. 307.Return to passage

[6] ‘Treatise on the Culture of the Apple,’ etc., p. 3.Return to passage

[7] Gallesio, ‘Teoria della Riproduzione Veg.,’ p. 125.Return to passage

[8] See Dr. Hooker’s Memoir on Arctic Plants in ‘Linn. Transact.,’ vol. xxiii. part ii. Mr. Woodward, and a higher authority cannot be quoted, speaks of the Arctic mollusca (in his ‘Rudimentary Treatise,’ 1856, p. 355) as remarkably subject to variation.Return to passage

[9] Bechstein, in his ‘Naturgeschichte der Stubenvögel,’ 1840, s. 238, has some good remarks on this subject. He states that his canary-birds varied in colour, though kept on uniform food.Return to passage

[10] ‘The Plant,’ by Schleiden, translated by Henfrey, 1848, p. 169. See also Alex. Braun, in ‘Bot. Memoirs,’ Ray Soc., 1853, p. 313.Return to passage

[11] Messrs. Hardy and Son, of Maldon, in ‘Gardener’s Chronicle,’ 1856, p. 458. Carrière, ‘Production et Fixation des Variétés,’ 1865, p. 31.Return to passage

[12] ‘Quadrupedes du Paraguay,’ 1801, tom. ii. p. 319.Return to passage

[13] M’Clelland on Indian Cyprinidæ, ‘Asiatic Researches,’ vol. xix. part ii., 1839, pp. 266, 268, 313.Return to passage

[14] Quoted by Sageret, ‘Pom. Phys.,’ 1830, p. 43. This statement, however, is not believed by Decaisne.Return to passage

[15] ‘The Fruits of America,’ 1845, p. 5.Return to passage

[16] M. Cardan, in ‘Comptes Rendus,’ Dec. 1848, quoted in ‘Gardener’s Chronicle,’ 1849, p. 101.Return to passage

[17] M. Alexis Jordan mentions four excellent pears found in woods in France, and alludes to others (‘Mém. Acad. de Lyon,’ tom. ii. 1852, p. 159). Poiteau’s remark is quoted in ‘Gardener’s Mag.,’ vol. iv., 1828, p. 385. See ‘Gardener’s Chronicle,’ 1862, p. 335, for another case of a new variety of the pear found in a hedge in France. Also for another case, see Loudon’s ‘Encyclop. of Gardening,’ p. 901. Mr. Rivers has given me similar information.Return to passage

[18] Duval, ‘Hist. du Poirier,’ 1849, p. 2.Return to passage

[19] I infer that this is the fact from Van Mons’ statement (‘Arbres Fruitiers,’ 1835, tom. i. p. 446) that he finds in the woods seedlings resembling all the chief cultivated races of both the pear and apple. Van Mons, however, looked at these wild varieties as aboriginal species.Return to passage

[20] Downing, ‘Fruit-trees of North America,’ p. 422; Foley, in ‘Transact. Hort. Soc.,’ vol. vi. p. 412.Return to passage

[21] ‘Gardener’s Chronicle,’ 1847, p. 244.Return to passage

[22] ‘Gardener’s Chronicle,’ 1841, p. 383; 1850, p. 700; 1854, p. 650.Return to passage

[23] ‘Die Getreidearten,’ 1843, s. 66, 116, 117.Return to passage

[24] Sabine, in ‘Hort. Transact.,’ vol. iii. p. 225; Bronn, ‘Geschichte der Natur,’ b. ii. s. 119.Return to passage

[25] ‘Journal of Horticulture,’ 1861, p. 112; on Zinnia, ‘Gardener’s Chronicle,’ 1860, p. 852.Return to passage

[26] ‘The Chrysanthemum, its History, etc.,’ 1865, p. 3.Return to passage

[27] ‘Gardener’s Chronicle,’ 1855, p. 54; ‘Journal of Horticulture,’ May 9, 1865, p. 363.Return to passage

[28] Quoted by Verlot, ‘Des Variétés,’ etc., 1865, p. 28.Return to passage

[29] ‘Examination of the Characteristics of Genera and Species,’ Charleston, 1855, p. 14.Return to passage

[30] Mr. Hewitt, ‘Journal of Hort.,’ 1863, p. 39.Return to passage

[31] Devay, ‘Mariages Consanguins,’ pp. 97, 125. In conversation I have found two or three naturalists of the same opinion.Return to passage

[32] Müller has conclusively argued against this belief, ‘Elements of Phys.,’ Eng. translat., vol. ii. 1842, p. 1405.Return to passage

[33] ‘Act. Acad. St. Petersburg,’ 1780, part ii. p. 84, etc.Return to passage

[34] ‘Bastarderzeugung,’ s. 249, 255, 295.Return to passage

[35] ‘Nova Acta, St. Petersburg,’ 1794, p. 378; 1795, pp. 307, 313, 316; 1787, p. 407.Return to passage

[36] ‘De la Fécondation,’ 1862, p. 311.Return to passage

[37] ‘Amaryllidaceæ,’ 1837, p. 362.Return to passage

[38] Abstracted in ‘Gardener’s Chronicle,’ 1860, p. 1081.Return to passage

[39] This was the opinion of the elder De Candolle, as quoted in ‘Dic. Class. d’Hist. Nat.,’ tom. viii. p. 405. Puvis, in his work, ‘De la Dégénération,’ 1837, p. 37, has discussed this same point.Return to passage

[40] ‘Comptes Rendus,’ Novembre 21, 1864, p. 838.Return to passage

[41] ‘Nova Acta, St. Petersburg,’ 1794, p. 391.Return to passage

[42] ‘Bastarderzeugung,’ s. 507, 516, 572.Return to passage

[43] ‘Die Bastardbefruchtung,’ etc., 1865, s. 24.Return to passage

[44] ‘Bastarderzeugung,’ s. 452, 507.Return to passage

[45] ‘Die Bastardbefruchtung,’ s. 56.Return to passage

[46] ‘Bastarderzeugung,’ s. 423.Return to passage

[47] ‘Dritte Fortsetzung,’ etc., 1766, s. 85.Return to passage

[48] ‘Die Bastardbefruchtung,’ etc., 1865, s. 92: see also the Rev. M. J. Berkeley on the same subject, in ‘Journal of Royal Hort. Soc.,’ 1866, p. 80.Return to passage

[49] Dr. P. Lucas has given a history of opinion on this subject: ‘Héréd. Nat.,’ 1847, tom. i. p. 175.Return to passage

[50] ‘Hist. des Anomalies,’ tom. iii. p. 499.Return to passage

[51] Ibid., tom. iii. pp. 392, 502. The several memoirs by M. Dareste hereafter referred to are of special value on this whole subject.Return to passage

[52] See his interesting work, ‘Métamorphoses de l’Homme,’ etc., 1862, p. 129.Return to passage

[53] ‘Dritte Fortsetzung,’ etc., s. 123; ‘Bastarderzeugung’ s. 249.Return to passage