With respect to the influence of the conditions of life on any two breeds which are allowed to cross freely, unless both are indigenous and have long been accustomed to the country where they live, they will, in all probability, be unequally affected by the conditions, and this will modify the result. Even with indigenous breeds, it will rarely or never occur that both are equally well adapted to the surrounding circumstances; more especially when permitted to roam freely, and not carefully tended, as will generally be the case with breeds allowed to cross. As a consequence of this, natural selection will to a certain extent come into action, and the best fitted will survive, and this will aid in determining the ultimate character of the commingled body.
How long a time it would require before such a crossed body of animals would assume within a limited area a uniform character no one can say; that they would ultimately become uniform from free intercrossing, and from the survival of the fittest, we may feel assured; but the character thus acquired would rarely or never, as we may infer from the several previous
considerations, be exactly intermediate between that of the two parent-breeds. With respect to the very slight differences by which the individuals of the same sub-variety, or even of allied varieties, are characterised, it is obvious that free crossing would soon obliterate such small distinctions. The formation of new varieties, independently of selection, would also thus be prevented; except when the same variation continually recurred from the action of some strongly predisposing cause. Hence we may conclude that free crossing has in all cases played an important part in giving to all the members of the same domestic race, and of the same natural species, uniformity of character, though largely modified by natural selection and by the direct action of the surrounding conditions.
On the possibility of all organic beings occasionally intercrossing.—But it may be asked, can free crossing occur with hermaphrodite animals and plants? All the higher animals, and the few insects which have been domesticated, have separated sexes, and must inevitably unite for each birth. With respect to the crossing of hermaphrodites, the subject is too large for the present volume, and will be more properly treated in a succeeding work. In my 'Origin of Species,' however, I have given a short abstract of the reasons which induce me to believe that all organic beings occasionally cross, though perhaps in some cases only at long intervals of time.[[192]] I will here just recall the fact that many plants, though hermaphrodite in structure, are unisexual in function;—such as those called by C. K. Sprengel dichogamous, in which the pollen and stigma of the same flower are matured at different periods; or those called by me reciprocally dimorphic, in which the flower's own pollen is not fitted to fertilise its own stigma; or again, the many kinds in which curious mechanical contrivances exist, effectually preventing self-fertilisation. There are, however, many hermaphrodite plants which are not in any way specially constructed to favour intercrossing, but which nevertheless commingle almost as freely as animals with separated sexes. This is the case with cabbages, radishes, and onions, as I know from
having experimented on them: even the peasants of Liguria say that cabbages must be prevented "from falling in love" with each other. In the orange tribe, Gallesio[[193]] remarks that the amelioration of the various kinds is checked by their continual and almost regular crossing. So it is with numerous other plants.
Nevertheless some cultivated plants can be named which rarely intercross, as the common pea, or which never intercross, as I have reason to believe is the case with the sweet-pea (Lathyrus odoratus); yet the structure of these flowers certainly favours an occasional cross. The varieties of the tomato and aubergine (Solanum) and pimenta (Pimenta vulgaris?) are said[[194]] never to cross, even when growing alongside each other. But it should be observed that these are all exotic plants, and we do not know how they would behave in their native country when visited by the proper insects.
It must also be admitted that some few natural species appear under our present state of knowledge to be perpetually self-fertilised, as in the case of the Bee Ophrys (O. apifera), though adapted in its structure to be occasionally crossed. The Leersia oryzoides produces minute enclosed flowers which cannot possibly be crossed, and these alone, to the exclusion of the ordinary flowers, have as yet been known to yield seed.[[195]] A few additional and analogous cases could be advanced. But these facts do not make me doubt that it is a general law of nature that the individuals of the same species occasionally intercross, and that some great advantage is derived from this act. It is well known (and I shall hereafter have to give instances) that some plants, both indigenous and naturalised, rarely or never produce flowers; or, if they flower, never produce seeds. But no one is thus led to doubt that it is a general law of nature that phanerogamic plants should produce flowers, and that these flowers should produce seed. When they fail, we believe that such plants would perform their proper functions under different conditions, or that they formerly did so and will do so again. On analogous grounds, I believe that the few flowers
which do not now intercross, either would do so under different conditions, or that they formerly fertilised each other at intervals—the means for effecting this being generally still retained—and they will do so again at some future period, unless indeed they become extinct. On this view alone, many points in the structure and action of the reproductive organs in hermaphrodite plants and animals are intelligible,—for instance, the male and female organs never being so completely enclosed as to render access from without impossible. Hence we may conclude that the most important of all the means for giving uniformity to the individuals of the same species, namely, the capacity of occasionally intercrossing, is present, or has been formerly present, with all organic beings.
On certain Characters not blending.—When two breeds are crossed their characters usually become intimately fused together; but some characters refuse to blend, and are transmitted in an unmodified state either from both parents or from one. When grey and white mice are paired, the young are not piebald nor of an intermediate tint, but are pure white or of the ordinary grey colour: so it is when white and common collared turtle-doves are paired. In breeding Game fowls, a great authority, Mr. J. Douglas, remarks, "I may here state a strange fact: if you cross a black with a white game, you get birds of both breeds of the clearest colour." Sir R. Heron crossed during many years white, black, brown, and fawn-coloured Angora rabbits, and never once got these colours mingled in the same animal, but often all four colours in the same litter.[[196]] Additional cases could be given, but this form of inheritance is very far from universal even with respect to the most distinct colours. When turnspit dogs and ancon sheep, both of which have dwarfed limbs, are crossed with common breeds, the offspring are not intermediate in structure, but take after either parent. When tailless or hornless animals are crossed with perfect animals, it frequently, but by no means invariably, happens that the offspring are either perfectly furnished with these organs or are quite destitute of them. According to Rengger, the hairless condition of the Paraguay dog is either perfectly or not at all transmitted to its mongrel offspring; but I have seen one partial exception in a dog of this parentage which had part of its skin hairy, and part naked; the parts being distinctly separated as in a piebald animal. When Dorking fowls with five toes are crossed with other breeds, the chickens often have five toes on one foot and four on the other. Some crossed pigs raised by Sir R. Heron between the solid-hoofed and common pig had not all four feet in an intermediate condition, but two feet were furnished with properly divided, and two with united hoofs.
Analogous facts have been observed with plants: Major Trevor Clarke crossed the little, glabrous-leaved, annual stock (Matthiola), with pollen of a large, red-flowered, rough-leaved, biennial stock, called cocardeau by the French, and the result was that half the seedlings had glabrous and the other half rough leaves, but none had leaves in an intermediate state. That the glabrous seedlings were the product of the rough-leaved variety, and not accidentally of the mother-plant's own pollen, was shown by their tall and strong habit of growth.[[197]] In the succeeding generations raised from the rough-leaved crossed seedlings, some glabrous plants appeared, showing that the glabrous character, though incapable of blending with and modifying the rough leaves, was all the time latent in this family of plants. The numerous plants formerly referred to, which I raised from reciprocal crosses between the peloric and common Antirrhinum, offer a nearly parallel case; for in the first generation all the plants resembled the common form, and in the next generation, out of one hundred and thirty-seven plants, two alone were in an intermediate condition, the others perfectly resembling either the peloric or common form. Major Trevor Clarke also fertilised the above-mentioned red-flowered stock with pollen from the purple Queen stock, and about half the seedlings scarcely differed in habit, and not at all in the red colour of the flower, from the mother-plant, the other half bearing blossoms of a rich purple, closely like those of the paternal plant. Gärtner crossed many white and yellow-flowered species and varieties of Verbascum; and these colours were never blended, but the offspring bore either pure white or pure yellow blossoms; the former in the larger proportion.[[198]] Dr. Herbert raised many seedlings, as he informed me, from Swedish turnips crossed by two other varieties, and these never produced flowers of an intermediate tint, but always like one of their parents. I fertilised the purple sweet-pea (Lathyrus odoratus), which has a dark reddish-purple standard-petal and violet-coloured wings and keel, with pollen of the painted-lady sweet-pea, which has a pale cherry-coloured standard, and almost white wings and keel; and from the same pod I twice raised plants perfectly resembling both sorts; the greater number resembling the father. So perfect was the resemblance, that I should have thought there had been some mistake, if the plants which were at first identical with the paternal variety, namely, the painted-lady, had not later in the season produced, as mentioned in a former chapter, flowers blotched and streaked with dark purple. I raised grandchildren and great-grandchildren from these crossed plants, and they continued to resemble the painted-lady, but during the later generations became rather more blotched with purple, yet none reverted completely to the original mother-plant, the purple sweet-pea. The following case is slightly different, but still shows the same principle: Naudin[[199]] raised numerous hybrids between the yellow Linaria vulgaris and the purple L. purpurea, and during three successive generations the colours kept distinct in different parts of the same flower.
From such cases as the foregoing, in which the offspring of the first generation perfectly resemble either parent, we come by a small step to those cases in which differently coloured flowers borne on the same root resemble both parents, and by another step to those in which the same flower or fruit is striped or blotched with the two parental colours, or bears a single stripe of the colour or other characteristic quality of one of the parent-forms. With hybrids and mongrels it frequently or even generally happens that one part of the body resembles more or less closely one parent and another part the other parent; and here again some resistance to fusion, or, what comes to the same thing, some mutual affinity between the organic atoms of the same nature, apparently comes into play, for otherwise all parts of the body would be equally intermediate in character. So again, when the offspring of hybrids or mongrels, which are themselves nearly intermediate in character, revert either wholly or by segments to their ancestors, the principle of the affinity of similar, or the repulsion of dissimilar atoms, must come into action. To this principle, which seems to be extremely general, we shall recur in the chapter on pangenesis.
It is remarkable, as has been strongly insisted upon by Isidore Geoffroy St. Hilaire in regard to animals, that the transmission of characters without fusion occurs most rarely when species are crossed; I know of one exception alone, namely, with the hybrids naturally produced between the common and hooded crow (Corvus corone and cornix), which, however, are closely allied species, differing in nothing except colour. Nor have I met with any well-ascertained cases of transmission of this kind, even when one form is strongly prepotent over another, when two races are crossed which have been slowly formed by man's selection, and therefore resemble to a certain extent natural species. Such cases as puppies in the same litter closely resembling two distinct breeds, are probably due to super-fœtation,—that is, to the influence of two fathers. All the characters above enumerated, which are transmitted in a perfect state to some of the offspring and not to others,—such as distinct colours, nakedness of skin, smoothness of leaves, absence of horns or tail, additional toes, pelorism, dwarfed structure, &c.,—have all been known to appear suddenly in individual animals and plants. From this fact, and from the several slight, aggregated differences which distinguish domestic races and species from each other, not being liable to this peculiar form of transmission, we may conclude that it is in some way connected with the sudden appearance of the characters in question.
On the Modification of old Races and the Formation of new Races by Crossing.—We have hitherto chiefly considered the effects of crossing in giving uniformity of character; we must now look to an opposite result. There can be no doubt that crossing, with the aid of rigorous selection during several generations, has been a potent means in modifying old races, and in forming new ones. Lord Orford crossed his famous stud of greyhounds once with the bulldog, which breed was chosen from being deficient in scenting powers, and from having what was wanted, courage and perseverance. In the course of six or seven generations all traces of the external form of the bulldog were eliminated, but courage and perseverance remained. Certain pointers have been crossed, as I hear from the Rev. W. D. Fox, with the foxhound, to give them dash and speed. Certain strains of Dorking fowls have had a slight infusion of Game blood; and I have known a great fancier who on a single occasion crossed his turbit-pigeons with barbs, for the sake of gaining greater breadth of beak.