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

in a greater or lesser degree, to one or both of their progenitors. Several authors have maintained that hybrids and mongrels include all the characters of both parents, not fused together, but merely mingled in different proportions in different parts of the body; or, as Naudin113 has expressed it, a hybrid is a living mosaic-work, in which the eye cannot distinguish the discordant elements, so completely are they intermingled. We can hardly doubt that, in a certain sense, this is true, as when we behold in a hybrid the elements of both species segregating themselves into segments in the same flower or fruit, by a process of self-attraction or self-affinity; this segregation taking place either by seminal or by bud-propagation. Naudin further believes that the segregation of the two specific elements or essences is eminently liable to occur in the male and female reproductive matter; and he thus explains the almost universal tendency to reversion in successive hybrid generations. For this would be the natural result of the union of pollen and ovules, in both of which the elements of the same species had been segregated by self-affinity. If, on the other hand, pollen which included the elements of one species happened to unite with ovules including the elements of the other species, the intermediate or hybrid state would still be retained, and there would be no reversion. But it would, as I suspect, be more correct to say that the elements of both parent-species exist in every hybrid in a double state, namely, blended together and completely separate. How this is possible, and what the term specific essence or element may be supposed to express, I shall attempt to show in the hypothetical chapter on pangenesis.

      But Naudin's view, as propounded by him, is not applicable to the reappearance of characters lost long ago by variation; and it is hardly applicable to races or species which, after having been crossed at some former period with a distinct form, and having since lost all traces of the cross, nevertheless occasionally yield an individual which reverts (as in the case of the great-great-grandchild of the pointer Sappho) to the crossing form. The most simple case of reversion, namely, of a hybrid or mongrel to its grandparents, is connected by an almost perfect series with the extreme case of a purely-bred race recovering characters which had been lost during many ages; and we are thus led to infer that all the cases must be related by some common bond.

      Gärtner believed that only those hybrid plants which are highly sterile exhibit any tendency to reversion to their parent-forms. It is rash to doubt so good an observer, but this conclusion must I think be an error; and it may perhaps be accounted for by the nature of the genera observed by him, for he admits that the tendency differs in different genera. The statement is also directly contradicted by Naudin's observations, and by the notorious fact that perfectly fertile mongrels exhibit the tendency in a high degree, – even in a higher degree, according to Gärtner himself, than hybrids.114

      Gärtner further states that reversions rarely occur with hybrid plants raised from species which have not been cultivated, whilst, with those which have been long cultivated, they are of frequent occurrence. This conclusion explains a curious discrepancy: Max Wichura,115 who worked exclusively on willows, which had not been subjected to culture, never saw an instance of reversion; and he goes so far as to suspect that the careful Gärtner had not sufficiently protected his hybrids from the pollen of the parent-species: Naudin, on the other hand, who chiefly experimented on cucurbitaceous and other cultivated plants, insists more strenuously than any other author on the tendency to reversion in all hybrids. The conclusion that the condition of the parent-species, as affected by culture, is one of the proximate causes leading to reversion, agrees fairly well with the converse case of domesticated animals and cultivated plants being liable to reversion when they become feral; for in both cases the organisation or constitution must be disturbed, though in a very different way.

      Finally, we have seen that characters often reappear in purely-bred races without our being able to assign any proximate cause; but when they become feral this is either indirectly or directly induced by the change in their conditions of life. With crossed breeds, the act of crossing in itself certainly leads to the recovery of long-lost characters, as well as of those derived from either parent-form. Changed conditions, consequent on cultivation, and the relative position of buds, flowers, and seeds on the plant, all apparently aid in giving this same tendency. Reversion may occur either through seminal or bud generation, generally at birth, but sometimes only with an advance of age. Segments or portions of the individual may alone be thus affected. That a being should be born resembling in certain characters an ancestor removed by two or three, and in some cases by hundreds or even thousands of generations, is assuredly a wonderful fact. In these cases the child is commonly said to inherit such characters directly from its grandparents or more remote ancestors. But this view is hardly conceivable. If, however, we suppose that every character is derived exclusively from the father or mother, but that many characters lie latent in both parents during a long succession of generations, the foregoing facts are intelligible. In what manner characters may be conceived to lie latent, will be considered in a future chapter to which I have lately alluded.

      Latent Characters.– But I must explain what is meant by characters lying latent. The most obvious illustration is afforded by secondary sexual characters. In every female all the secondary male characters, and in every male all the secondary female characters, apparently exist in a latent state, ready to be evolved under certain conditions. It is well known that a large number of female birds, such as fowls, various pheasants, partridges, peahens, ducks, &c., when old or diseased, or when operated on, partly assume the secondary male characters of their species. In the case of the hen-pheasant this has been observed to occur far more frequently during certain seasons than during others.116 A duck ten years old has been known to assume both the perfect winter and summer plumage of the drake.117 Waterton118 gives a curious case of a hen which had ceased laying, and had assumed the plumage, voice, spurs, and warlike disposition of the cock; when opposed to an enemy she would erect her hackles and show fight. Thus every character, even to the instinct and manner of fighting, must have lain dormant in this hen as long as her ovaria continued to act. The females of two kinds of deer, when old, have been known to acquire horns; and, as Hunter has remarked, we see something of an analogous nature in the human species.

      On the other hand, with male animals, it is notorious that the secondary sexual characters are more or less completely lost when they are subjected to castration. Thus, if the operation be performed on a young cock, he never, as Yarrell states, crows again; the comb, wattles, and spurs do not grow to their full size, and the hackles assume an intermediate appearance between true hackles and the feathers of the hen. Cases are recorded of confinement alone causing analogous results. But characters properly confined to the female are likewise acquired; the capon takes to sitting on eggs, and will bring up chickens; and what is more curious, the utterly sterile male hybrids from the pheasant and the fowl act in the same manner, "their delight being to watch when the hens leave their nests, and to take on themselves the office of a sitter."119 That admirable observer Réaumur120 asserts that a cock, by being long confined in solitude and darkness, can be taught to take charge of young chickens; he then utters a peculiar cry, and retains during his whole life this newly acquired maternal instinct. The many well-ascertained cases of various male mammals giving milk, show that their rudimentary mammary glands retain this capacity in a latent condition.

      We thus see that in many, probably in all cases, the secondary characters of each sex lie dormant or latent in the opposite sex, ready to be evolved under peculiar circumstances. We can thus understand how, for instance, it is possible for a good milking cow to transmit her good qualities through her male offspring to future generations; for we may confidently believe that these qualities are present, though latent, in the males of each generation. So it is with the game-cock, who can transmit his superiority in courage and vigour through his female to his male offspring; and with man it is known 121 that diseases, such as hydrocele, necessarily confined to the male sex, can be transmitted through the female to the grandson. Such cases as these offer, as was remarked at the commencement of this chapter, the simplest possible examples

113

'Nouvelles Archives du Muséum,' tom. i. p. 151.

114

'Bastarderzeugung,' s. 582, 438, &c.

115

'Die Bastardbefruchtung … der Weiden,' 1865, s. 23. For Gärtner's remarks on this head, see 'Bastarderzeugung,' s. 474, 582.

116

Yarrell, 'Phil. Transact.,' 1827, p. 268; Dr. Hamilton, in 'Proc. Zoolog. Soc.,' 1862, p. 23.

117

'Archiv. Skand. Beiträge zur Naturgesch.,' viii. s. 397-413.

118

In his 'Essays on Nat. Hist.,' 1838. Mr. Hewitt gives analogous cases with hen-pheasants in 'Journal of Horticulture,' July 12, 1864, p. 37. Isidore Geoffroy Saint Hilaire, in his 'Essais de Zoolog. Gén.' (suites à Buffon, 1842, pp. 496-513), has collected such cases in ten different kinds of birds. It appears that Aristotle was well aware of the change in mental disposition in old hens. The case of the female deer acquiring horns is given at p. 513.

119

'Cottage Gardener,' 1860, p. 379.

120

'Art de faire Eclorre,' &c., 1749, tom. ii. p. 8.

121

Sir H. Holland, 'Medical Notes and Reflections,' 3rd edit., 1855, p. 31.