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

influences and been truly transmitted, will, as a general rule, continue to withstand them, and consequently be faithfully inherited.

Prepotency in the Transmission of Character

      When individuals distinct enough to be recognised, but of the same family, or when two well-marked races, or two species, are crossed, the usual result, as stated in the previous chapter, is, that the offspring in the first generation are intermediate between their parents, or resemble one parent in one part and the other parent in another part. But this is by no means the invariable rule; for in many cases it is found that certain individuals, races, and species are prepotent in transmitting their likeness. This subject has been ably discussed by Prosper Lucas,137 but is rendered extremely complicated by the prepotency sometimes running equally in both sexes, and sometimes more strongly in one sex than in the other; it is likewise complicated by the presence of secondary sexual characters, which render the comparison of mongrels with their parent-breeds difficult.

      It would appear that in certain families some one ancestor, and after him others in the same family, must have had great power in transmitting their likeness through the male line; for we cannot otherwise understand how the same features should so often be transmitted after marriages with various females, as has been the case with the Austrian Emperors, and as, according to Niebuhr, formerly occurred in certain Roman families with their mental qualities.138 The famous bull Favourite is believed139 to have had a prepotent influence on the shorthorn race. It has also been observed140 with English race-horses that certain mares have generally transmitted their own character, whilst other mares of equally pure blood have allowed the character of the sire to prevail.

      The truth of the principle of prepotency comes out more clearly when certain races are crossed. The improved Shorthorns, notwithstanding that the breed is comparatively modern, are generally acknowledged to possess great power in impressing their likeness on all other breeds; and it is chiefly in consequence of this power that they are so highly valued for exportation.141 Godine has given a curious case of a ram of a goat-like breed of sheep from the Cape of Good Hope, which produced offspring hardly to be distinguished from himself, when crossed with ewes of twelve other breeds. But two of these half-bred ewes, when put to a merino ram, produced lambs closely resembling the merino breed. Girou de Buzareingues142 found that of two races of French sheep the ewes of one, when crossed during successive generations with merino rams, yielded up their character far sooner than the ewes of the other race. Sturm and Girou have given analogous cases with other breeds of sheep and with cattle, the prepotency running in these cases through the male side; but I was assured on good authority in South America, that when niata cattle are crossed with common cattle, though the niata breed is prepotent whether males or females are used, yet that the prepotency is strongest through the female line. The Manx cat is tailless and has long hind legs; Dr. Wilson crossed a male Manx with common cats, and, out of twenty-three kittens, seventeen were destitute of tails; but when the female Manx was crossed by common male cats all the kittens had tails, though they were generally short and imperfect.143

      In making reciprocal crosses between pouter and fantail pigeons, the pouter-race seemed to be prepotent through both sexes over the fantail. But this is probably due to weak power in the fantail rather than to any unusually strong power in the pouter, for I have observed that barbs also preponderated over fantails. This weakness of transmission in the fantail, though the breed is an ancient one, is said144 to be general; but I have observed one exception to the rule, namely, in a cross between a fantail and laugher. The most curious instance known to me of weak power in both sexes is in the trumpeter pigeon. This breed has been well known for at least 130 years: it breeds perfectly true, as I have been assured by those who have long kept many birds: it is characterised by a peculiar tuft of feathers over the beak, by a crest on the head, by a most peculiar coo quite unlike that of any other breed, and by much-feathered feet. I have crossed both sexes with turbits of two sub-breeds, with almond tumblers, spots, and runts, and reared many mongrels and recrossed them; and though the crest on the head and feathered feet were inherited (as is generally the case with most breeds), I have never seen a vestige of the tuft over the beak or heard the peculiar coo. Boitard and Corbié145 assert that this is the invariable result of crossing trumpeters with any other breed: Neumeister,146 however, states that in Germany mongrels have been obtained, though very rarely, which were furnished with the tuft and would trumpet: but a pair of these mongrels with a tuft, which I imported, never trumpeted. Mr. Brent states147 that the crossed offspring of a trumpeter were crossed with trumpeters for three generations, by which time the mongrels had 7-8ths of this blood in their veins, yet the tuft over the beak did not appear. At the fourth generation the tuft appeared, but the birds, though now having 15-16ths trumpeter's blood, still did not trumpet. This case well shows the wide difference between inheritance and prepotency; for here we have a well-established old race which transmits it characters faithfully, but which, when crossed with any other race, has the feeblest power of transmitting its two chief characteristic qualities.

      I will give one other instance with fowls and pigeons of weakness and strength in the transmission of the same character to their crossed offspring. The Silk-fowl breeds true, and there is reason to believe is a very ancient race; but when I reared a large number of mongrels from a Silk-hen by a Spanish cock, not one exhibited even a trace of the so-called silkiness. Mr. Hewitt also asserts that in no instance are the silky feathers transmitted by this breed when crossed with any other variety. But three birds out of many raised by Mr. Orton from a cross between a silk-cock and a bantam-hen, had silky feathers.148 So that it is certain that this breed very seldom has the power of transmitting its peculiar plumage to its crossed progeny. On the other hand, there is a silk sub-variety of the fantail pigeon, which has its feathers in nearly the same state as in the Silk-fowl: now we have already seen that fantails, when crossed, possess singularly weak power in transmitting their general qualities; but the silk sub-variety when crossed with any other small-sized race invariably transmits its silky feathers!149

      The law of prepotency comes into action when species are crossed, as with races and individuals. Gärtner has unequivocally shown150 that this is the case with plants. To give one instance: when Nicotiana paniculata and vincæflora are crossed, the character of N. paniculata is almost completely lost in the hybrid; but if N. quadrivalvis be crossed with N. vincæflora, this later species, which was before so prepotent, now in its turn almost disappears under the power of N. quadrivalvis. It is remarkable that the prepotency of one species over another in transmission is quite independent, as shown by Gärtner, of the greater or less facility with which the one fertilises the other.

      With animals, the jackal is prepotent over the dog, as is stated by Flourens who made many crosses between these animals; and this was likewise the case with a hybrid which I once saw between a jackal and terrier. I cannot doubt, from the observations of Colin and others, that the ass is prepotent over the horse; the prepotency in this instance running more strongly through the male than through the female ass; so that the mule resembles the ass more closely than does the hinny.151 The male pheasant, judging from Mr. Hewitt's descriptions,152 and from the hybrids which I have seen, preponderates over the domestic fowl; but the latter, as far as colour is concerned, has considerable power of transmission, for hybrids raised from five differently coloured hens differed greatly in plumage. I formerly examined some curious hybrids in the Zoological Gardens, between the Penguin variety of the common duck and the Egyptian goose (Tadorna Ægyptiaca); and although I will not assert that the domesticated variety preponderated over the natural species, yet it had strongly impressed its unnatural

137

'Héréd. Nat.,' tom. ii. pp. 112-120.

138

Sir H. Holland, 'Chapters on Mental Physiology,' 1852, p. 234.

139

'Gardener's Chronicle,' 1860, p. 270.

140

Mr. N. H. Smith, Observations on Breeding, quoted in 'Encyclop. of Rural Sports,' p. 278.

141

Quoted by Bronn, 'Geschichte der Natur,' b. ii. s. 170. See Sturm, 'Ueber Racen,' 1825, s. 104-107. For the niata cattle, see my 'Journal of Researches,' 1845, p. 146.

142

Lucas, 'l'Hérédité Nat.,' tom. ii. p. 112.

143

Mr. Orton, 'Physiology of Breeding,' 1855, p. 9.

144

Boitard and Corbié, 'Les Pigeons,' 1824, p. 224.

145

'Les Pigeons, pp. 168, 198.

146

'Das Ganze,' &c., 1837, s. 39.

147

'The Pigeon Book,' p. 46.

148

'Physiology of Breeding,' p.22; Mr. Hewitt, in 'The Poultry Book,' by Tegetmeier, 1866, p. 224.

149

Boitard and Corbié, 'Les Pigeons,' 1824, p. 226.

150

'Bastarderzeugung,' s. 256, 290, &c. Naudin ('Nouvelles Archives du Muséum,' tom. i. p. 149) gives a striking instance of prepotency in Datura stramonium when crossed with two other species.

151

Flourens, 'Longévité Humaine,' p. 144, on crossed jackals. With respect to the difference between the mule and the hinny, I am aware that this has generally been attributed to the sire and dam transmitting their characters differently; but Colin, who has given in his 'Traité Phys. Comp.,' tom. ii. pp. 537-539, the fullest description which I have met with of these reciprocal hybrids, is strongly of opinion that the ass preponderates in both crosses, but in an unequal degree. This is likewise the conclusion of Flourens, and of Bechstein in his 'Naturgeschichte Deutschlands,' b. i. s. 294. The tail of the hinny is much more like that of the horse than is the tail of the mule, and this is generally accounted for by the males of both species transmitting with greater power this part of their structure; but a compound hybrid which I saw in the Zoological Gardens, from a mare by a hybrid ass-zebra, closely resembled its mother in its tail.

152

Mr. Hewitt, who has had such great experience in raising these hybrids, says ('Poultry Book,' by Mr. Tegetmeier, 1866, pp. 165-167) that in all, the head was destitute of wattles, comb, and ear-lappets; and all closely resembled the pheasant in the shape of the tail and general contour of the body. These hybrids were raised from hens of several breeds by a cock-pheasant; but another hybrid, described by Mr. Hewitt, was raised from a hen-pheasant by a silver-laced Bantam cock, and this possessed a rudimental comb and wattles.