The Variation of Animals and Plants Under Domestication, Volume II (of 2). Darwin Charles

bears indirectly on two or three points connected with the modification of organic beings. As Buffon long ago remarked,242 domestic animals breed oftener in the year and produce more young at a birth than wild animals of the same species; they, also, sometimes breed at an earlier age. The case would hardly have deserved further notice, had not some authors lately attempted to show that fertility increases and decreases in an inverse ratio with the amount of food. This strange doctrine has apparently arisen from individual animals when supplied with an inordinate quantity of food, and from plants of many kinds when grown on excessively rich soil, as on a dunghill, becoming sterile; but to this latter point I shall have occasion presently to return. With hardly an exception, our domesticated animals, which have long been habituated to a regular and copious supply of food, without the labour of searching for it, are more fertile than the corresponding wild animals. It is notorious how frequently cats and dogs breed, and how many young they produce at a birth. The wild rabbit is said generally to breed four times yearly, and to produce from four to eight young; the tame rabbit breeds six or seven times yearly, and produces from four to eleven young. The ferret, though generally so closely confined, is more prolific than its supposed wild prototype. The wild sow is remarkably prolific, for she often breeds twice in the year, and produces from four to eight and sometimes even twelve young at a birth; but the domestic sow regularly breeds twice a year, and would breed oftener if permitted; and a sow that produces less than eight at a birth "is worth little, and the sooner she is fattened for the butcher the better." The amount of food affects the fertility even of the same individual: thus sheep, which on mountains never produce more than one lamb at a birth, when brought down to lowland pastures frequently bear twins. This difference apparently is not due to the cold of the higher land, for sheep and other domestic animals are said to be extremely prolific in Lapland. Hard living, also, retards the period at which animals conceive; for it has been found disadvantageous in the northern islands of Scotland to allow cows to bear calves before they are four years old.243

      Birds offer still better evidence of increased fertility from domestication: the hen of the wild Gallus bankiva lays from six to ten eggs, a number which would be thought nothing of with the domestic hen. The wild duck lays from five to ten eggs; the tame one in the course of the year from eighty to one hundred. The wild grey-lag goose lays from five to eight eggs; the tame from thirteen to eighteen, and she lays a second time; as Mr. Dixon has remarked, "high-feeding, care, and moderate warmth induce a habit of prolificacy which becomes in some measure hereditary." Whether the semi-domesticated dovecot pigeon is more fertile than the wild rock-pigeon C. livia, I know not; but the more thoroughly domesticated breeds are nearly twice as fertile as dovecots: the latter, however, when caged and highly fed, become equally fertile with house pigeons. The peahen alone of domesticated birds is rather more fertile, according to some accounts, when wild in its native Indian home, than when domesticated in Europe and exposed to our much colder climate.244

      With respect to plants, no one would expect wheat to tiller more, and each ear to produce more grain, in poor than in rich soil; or to get in poor soil a heavy crop of peas or beans. Seeds vary so much in number that it is difficult to estimate them; but on comparing beds of carrots saved for seed in a nursery garden with wild plants, the former seemed to produce about twice as much seed. Cultivated cabbages yielded thrice as many pods by measure as wild cabbages from the rocks of South Wales. The excess of berries produced by the cultivated Asparagus in comparison with the wild plant is enormous. No doubt many highly cultivated plants, such as pears, pineapples, bananas, sugar-cane, &c., are nearly or quite sterile; and I am inclined to attribute this sterility to excess of food and to other unnatural conditions; but to this subject I shall presently recur.

      In some cases, as with the pig, rabbit, &c., and with those plants which are valued for their seed, the direct selection of the more fertile individuals has probably much increased their fertility; and in all cases this may have occurred indirectly, from the better chance of the more numerous offspring produced by the more fertile individuals having survived. But with cats, ferrets, and dogs, and with plants like carrots, cabbages, and asparagus, which are not valued for their prolificacy, selection can have played only a subordinate part; and their increased fertility must be attributed to the more favourable conditions of life under which they have long existed.

       CHAPTER XVII.

      ON THE GOOD EFFECTS OF CROSSING, AND ON THE EVIL EFFECTS OF CLOSE INTERBREEDING

      DEFINITION OF CLOSE INTERBREEDING – AUGMENTATION OF MORBID TENDENCIES – GENERAL EVIDENCE ON THE GOOD EFFECTS DERIVED FROM CROSSING, AND ON THE EVIL EFFECTS FROM CLOSE INTERBREEDING – CATTLE, CLOSELY INTERBRED; HALF-WILD CATTLE LONG KEPT IN THE SAME PARKS – SHEEP – FALLOW-DEER – DOGS – RABBITS – PIGS – MAN, ORIGIN OF HIS ABHORRENCE OF INCESTUOUS MARRIAGES – FOWLS – PIGEONS – HIVE-BEES – PLANTS, GENERAL CONSIDERATIONS ON THE BENEFITS DERIVED FROM CROSSING – MELONS, FRUIT-TREES, PEAS, CABBAGES, WHEAT, AND FOREST-TREES – ON THE INCREASED SIZE OF HYBRID PLANTS, NOT EXCLUSIVELY DUE TO THEIR STERILITY – ON CERTAIN PLANTS WHICH EITHER NORMALLY OR ABNORMALLY ARE SELF-IMPOTENT, BUT ARE FERTILE, BOTH ON THE MALE AND FEMALE SIDE, WHEN CROSSED WITH DISTINCT INDIVIDUALS EITHER OF THE SAME OR ANOTHER SPECIES – CONCLUSION.

      The gain in constitutional vigour, derived from an occasional cross between individuals of the same variety, but belonging to distinct families, or between distinct varieties, has not been so largely or so frequently discussed, as have the evil effects of too close interbreeding. But the former point is the more important of the two, inasmuch as the evidence is more decisive. The evil results from close interbreeding are difficult to detect, for they accumulate slowly, and differ much in degree with different species; whilst the good effects which almost invariably follow a cross are from the first manifest. It should, however, be clearly understood that the advantage of close interbreeding, as far as the retention of character is concerned, is indisputable, and often outweighs the evil of a slight loss of constitutional vigour. In relation to the subject of domestication, the whole question is of some importance, as too close interbreeding interferes with the improvement of old races, and especially with the formation of new ones. It is important as indirectly bearing on Hybridism; and perhaps on the extinction of species, when any form has become so rare that only a few individuals remain within a confined area. It bears in an important manner on the influence of free intercrossing, in obliterating individual differences, and thus giving uniformity of character to the individuals of the same race or species; for if additional vigour and fertility be thus gained, the crossed offspring will multiply and prevail, and the ultimate result will be far greater than otherwise would have occurred. Lastly, the question is of high interest, as bearing on mankind. Hence I shall discuss this subject at full length. As the facts which prove the evil effects of close interbreeding are more copious, though less decisive, than those on the good effects of crossing, I shall, under each group of beings, begin with the former.

      There is no difficulty in defining what is meant by a cross; but this is by no means easy in regard to "breeding in and in" or "too close interbreeding," because, as we shall see, different species of animals are differently affected by the same degree of interbreeding. The pairing of a father and daughter, or mother and son, or brothers and sisters, if carried on during several generations, is the closest possible form of interbreeding. But some good judges, for instance Sir J. Sebright, believe that the pairing of a brother and sister is closer than that of parents and children; for when the father is matched with his daughter he crosses, as is said, with only half his own blood. The consequences of close interbreeding carried on for too long a time, are, as is generally believed, loss of size, constitutional vigour, and fertility, sometimes accompanied by a tendency to malformation. Manifest evil does not usually follow from pairing the nearest relations for two, three, or even four generations; but several causes interfere with our detecting the evil – such as the deterioration being very gradual, and the difficulty of distinguishing between such direct evil and the inevitable augmentation of any morbid tendencies which may be latent or apparent in the related parents. On the other hand, the benefit from a cross, even when there has not been any very close interbreeding, is almost invariably at once conspicuous. There is reason to believe, and this was the opinion of that most experienced observer Sir J. Sebright,

242

Quoted by Isid. Geoffroy St. Hilaire, 'Hist. Naturelle Générale,' tom. iii. p. 476. Since this MS. has been sent to press a full discussion on the present subject has appeared in Mr. Herbert Spencer's 'Principles of Biology,' vol. ii. 1867, p. 457 et seq.

243

For cats and dogs, &c., see Bellingeri, in 'Annal. des Sc. Nat.,' 2nd series, Zoolog., tom. xii. p. 155. For ferrets, Bechstein, 'Naturgeschichte Deutschlands,' Band i., 1801, s. 786, 795. For rabbits, ditto, s. 1123, 1131; and Bronn's 'Geschichte der Natur,' B. ii. s. 99. For mountain sheep, ditto, s. 102. For the fertility of the wild sow, see Bechstein's 'Naturgesch. Deutschlands,' B. i., 1801, s. 534; for the domestic pig, Sidney's edit. of Youatt on the Pig, 1860, p. 62. With respect to Lapland, see Acerbi's 'Travels to the North Cape,' Eng. translat., vol. ii. p. 222. About the Highland cows, see Hogg on Sheep, p. 263.

244

For the eggs of Gallus bankiva, see Blyth, in 'Annals and Mag. of Nat. Hist., 2nd series, vol. i., 1848, p. 456. For wild and tame ducks, Macgillivray, 'British Birds,' vol. v. p. 37; and 'Die Enten,' s. 87. For wild geese, L. Lloyd, 'Scandinavian Adventures,' vol. ii. 1854, p. 413; and for tame geese, 'Ornamental Poultry,' by Rev. E. S. Dixon, p. 139. On the breeding of pigeons, Pistor, 'Das Ganze der Taubenzucht,' 1831, s. 46; and Boitard and Corbié, 'Les Pigeons,' p. 158. With respect to peacocks, according to Temminck ('Hist. Nat. Gén. des Pigeons,' &c., 1813, tom. ii. p. 41), the hen lays in India even as many as twenty eggs; but according to Jerdon and another writer (quoted in Tegetmeier's 'Poultry Book,' 1866, pp. 280, 282), she there lays only from four to nine or ten eggs: in England she is said, in the 'Poultry Book,' to lay five or six, but another writer says from eight to twelve eggs.