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28
HYBRIDISM


dealing with birds, concluded that no hybrids were fertile with one another beyond the second generation, but thought that they were fertile with members of the parent races. Wallace, on the other hand, cites from Quatrefages the case of hybrids between the moths Bombyx cynthia and B. arrindia, which were stated to be fertile inter se for eight generations. He also states that hybrids between the sheep and goat have a limited fertility inter se. Charles Darwin, however, had evidence that some hybrid pheasants were completely fertile, and he himself interbred the progeny of crosses between the common and Chinese geese, whilst there appears to be no doubt as to the complete fertility of the crosses between many species of ducks, J. L. Bonhote having interbred in various crosses for several generations the mallard (Anas boschas), the Indian spot-bill duck (A. poecilorhyncha), the New Zealand grey duck (A. superciliosa) and the pin-tail (Dafila acuta). Podmore’s pigeon hybrids were fertile inter se, a specimen having been exhibited at the London Zoological Gardens. The hybrids between the brown and polar bears bred at Halle proved to be fertile, both with one of the parent species and with one another.

Cornevin and Lesbre state that in 1873 an Arab mule was fertilized in Africa by a stallion, and gave birth to female offspring which she suckled. All three were brought to the Jardin d’Acclimatation in Paris, and there the mule had a second female colt to the same father, and subsequently two male colts in succession to an ass and to a stallion. The female progeny were fertilized, but their offspring were feeble and died at birth. Cossar Ewart gives an account of a recent Indian case in which a female mule gave birth to a male colt. He points out, however, that many mistakes have been made about the breeding of hybrids, and is not altogether inclined to accept this supposed case. Very little has been published with regard to the most important question, as to the actual condition of the sexual organs and cells in hybrids. There does not appear to be gross anatomical defect to account for the infertility of hybrids, but microscopical examination in a large number of cases is wanted. Cossar Ewart, to whom indeed much of the most interesting recent work on hybrids is due, states that in male zebra-hybrids the sexual cells were immature, the tails of the spermatozoa being much shorter than those of the similar cells in stallions and zebras. He adds, however, that the male hybrids he examined were young, and might not have been sexually mature. He examined microscopically the ovary of a female zebra-hybrid and found one large and several small Graafian follicles, in all respects similar to those in a normal mare or female zebra. A careful study of the sexual organs in animal and plant hybrids is very much to be desired, but it may be said that so far as our present knowledge goes there is not to be expected any obvious microscopical cause of the relative infertility of hybrids.

The relative variability of hybrids has received considerable attention from many writers. Horticulturists, as Bateson has written, are “aware of the great and striking variations which occur in so many orders of plants when hybridization is effected.” The phrase has been used “breaking the constitution of a plant” to indicate the effect produced in the offspring of a hybrid union, and the device is frequently used by those who are seeking for novelties to introduce on the market. It may be said generally that hybrids are variable, and that the products of hybrids are still more variable. J. L. Bonhote found extreme variations amongst his hybrid ducks. Y. Delage states that in reciprocal crosses there is always a marked tendency for the offspring to resemble the male parents; he quotes from Huxley that the mule, whose male parent is an ass, is more like the ass, and that the hinny, whose male parent is a horse, is more like the horse. Standfuss found among Lepidoptera that males were produced much more often than females, and that these males paired readily. The freshly hatched larvae closely resembled the larvae of the female parent, but in the course of growth the resemblance to the male increased, the extent of the final approximation to the male depending on the relative phylogenetic age of the two parents, the parent of the older species being prepotent. In reciprocal pairing, he found that the male was able to transmit the characters of the parents in a higher degree. Cossar Ewart, in relation to zebra hybrids, has discussed the matter of resemblance to parents in very great detail, and fuller information must be sought in his writings. He shows that the wild parent is not necessarily prepotent, although many writers have urged that view. He described three hybrids bred out of a zebra mare by different horses, and found in all cases that the resemblance to the male or horse parent was more profound. Similarly, zebra-donkey hybrids out of zebra mares bred in France and in Australia were in characters and disposition far more like the donkey parents. The results which he obtained in the hybrids which he bred from a zebra stallion and different mothers were more variable, but there was rather a balance in favour of zebra disposition and against zebra shape and marking.

“Of the nine zebra-horse hybrids I have bred,” he says, “only two in their make and disposition take decidedly after the wild parent. As explained fully below, all the hybrids differ profoundly in the plan of their markings from the zebra, while in their ground colour they take after their respective dams or the ancestors of their dams far more than after the zebra—the hybrid out of the yellow and white Iceland pony, e.g. instead of being light in colour, as I anticipated, is for the most part of a dark dun colour, with but indistinct stripes. The hoofs, mane and tail of the hybrids are at the most intermediate, but this is perhaps partly owing to reversion towards the ancestors of these respective dams. In their disposition and habits they all undoubtedly agree more with the wild sire.”

Ewart’s experiments and his discussion of them also throw important light on the general relation of hybrids to their parents. He found that the coloration and pattern of his zebra hybrids resembled far more those of the Somali or Grévy’s zebra than those of their sire—a Burchell’s zebra. In a general discussion of the stripings of horses, asses and zebras, he came to the conclusion that the Somali zebra represented the older type, and that therefore his zebra hybrids furnished important evidence of the effect of crossing in producing reversion to ancestral type. The same subject has of course been discussed at length by Darwin, in relation to the cross-breeding of varieties of pigeons; but the modern experimentalists who are following the work of Mendel interpret reversion differently (see Mendelism).

Graft-Hybridism.—It is well known that, when two varieties or allied species are grafted together, each retains its distinctive characters. But to this general, if not universal, rule there are on record several alleged exceptions, in which either the scion is said to have partaken of the qualities of the stock, the stock of the scion, or each to have affected the other. Supposing any of these influences to have been exerted, the resulting product would deserve to be called a graft-hybrid. It is clearly a matter of great interest to ascertain whether such formation of hybrids by grafting is really possible; for, if even one instance of such formation could be unequivocally proved, it would show that sexual and asexual reproduction are essentially identical.

The cases of alleged graft-hybridism are exceedingly few, considering the enormous number of grafts that are made every year by horticulturists, and have been so made for centuries. Of these cases the most celebrated are those of Adam’s laburnum (Cytisus Adami) and the bizzarria orange. Adam’s laburnum is now flourishing in numerous places throughout Europe, all the trees having been raised as cuttings from the original graft, which was made by inserting a bud of the purple laburnum into a stock of the yellow. M. Adam, who made the graft, has left on record that from it there sprang the existing hybrid. There can be no question as to the truly hybrid character of the latter—all the peculiarities of both parent species being often blended in the same raceme, flower or even petal; but until the experiment shall have been successfully repeated there must always remain a strong suspicion that, notwithstanding the assertion and doubtless the belief of M. Adam, the hybrid arose as a cross in the ordinary way of seminal reproduction. Similarly, the bizzarria orange, which is unquestionably a hybrid between the bitter orange and the citron—since it presents the remarkable spectacle of these two different fruits blended into one—is stated by the gardener who first succeeded in producing it to have arisen as a graft-hybrid; but here again a similar doubt, similarly due to the need of corroboration, attaches to the statement. And the same remark applies to the still more wonderful case of the so-called trifacial orange, which blends three distinct kinds of fruit in one, and which is said to have been produced by artificially splitting and uniting the seeds taken from the three distinct species, the fruits of which now occur blended in the triple hybrid.

The other instances of alleged graft-hybridism are too numerous to be here noticed in detail; they refer to jessamine, ash, hazel, vine, hyacinth, potato, beet and rose. Of these the cases of the vine, beet and rose are the strongest as evidence of graft-hybridization, from the fact that some of them were produced