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 longcivilised 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." 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
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, 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." 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, "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 longcontinued 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
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 In North America plum-trees are liable to a disease which Downing 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. 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. 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. 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. White verbenas are especially liable to mildew. 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. 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 lightcoloured animals suffer most: in Thuringia 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, was peculiarly sensitive to the bites of insects. In the West Indies 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; 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. 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.
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. Prof. G. Jaeger 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 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." 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 yellowberried 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 birdsmuch 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. 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.
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. 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. 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. 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 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 flowerbuds, 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 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 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. 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";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, 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. 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, 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.
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, 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." 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: 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 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." At Kew, as Mr. Beaton remarks, where many seedlings of common plants are raised, "you see new forms of Laburnums, Spiraeas, and other shrubs." So with animals: Marshall, 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."
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, 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 Wagnerremarks, 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 200l. 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 cattleshows, a day being given for their exhibition.
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. The common mignonette (Reseda odorata), from bearing inconspicuous flowers, valued solely for their fragrance, "remains in the same unimproved condition as when first introduced." 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, 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