When an individual having some recognisable peculiarity unites with another of the same sub-variety, not having the peculiarity in question, it often reappears in the descendants after an interval of several generations. Every one must have noticed, or heard from old people of children closely resembling in appearance or mental disposition, or in so small and complex a character as expression, one of their grandparents, or some more distant collateral relation. Very many anomalies of structure and diseases (13/16. Mr. Sedgwick gives many instances in the 'British and Foreign Med. — Chirurg. Review' April and July 1863 pages 448, 188.) of which instances have been given in the last chapter, have come into a family from one parent, and have reappeared in the progeny after passing over two or three generations. The following case has been communicated to me on good authority, and may, I believe, be fully trusted: a pointer-bitch produced seven puppies; four were marked with blue and white, which is so unusual a colour with pointers that she was thought to have played false with one of the greyhounds, and the whole litter was condemned; but the gamekeeper was permitted to save one as a curiosity. Two years afterwards a friend of the owner saw the young dog, and declared that he was the image of his old pointer-bitch Sappho, the only blue and white pointer of pure descent which he had ever seen. This led to close inquiry, and it was proved that he was the great-great-grandson of Sappho; so that, according to the common expression, he had only 1/16th of her blood in his veins. I may give one other instance, on the authority of Mr. R. Walker, a large cattle- breeder in Kincardineshire. He bought a black bull, the son of a black cow with white legs, white belly and part of the tail white; and in 1870 a calf the gr. — gr. — gr. — gr. — grandchild of this cow was born coloured in the same very peculiar manner; all the intermediate offspring having been black. In these cases there can hardly be a doubt that a character derived from a cross with an individual of the same variety reappeared after passing over three generations in the one case, and five in the other.
When two distinct races are crossed, it is notorious that the tendency in the offspring to revert to one or both parent-forms is strong, and endures for many generations. I have myself seen the clearest evidence of this in crossed pigeons and with various plants. Mr. Sidney (13/17. In his edition of 'Youatt on the Pig' 1860 page 27.) states that, in a litter of Essex pigs, two young ones appeared which were the image of the Berkshire boar that had been used twenty-eight years before in giving size and constitution to the breed. I observed in the farmyard at Betley Hall some fowls showing a strong likeness to the Malay breed, and was told by Mr. Tollet that he had forty years before crossed his birds with Malays; and that, though he had at first attempted to get rid of this strain, he had subsequently given up the attempt in despair, as the Malay character would reappear.
This strong tendency in crossed breeds to revert has given rise to endless discussions in how many generations after a single cross, either with a distinct breed or merely with an inferior animal, the breed may be considered as pure, and free from all danger of reversion. No one supposes that less than three generations suffices, and most breeders think that six, seven, or eight are necessary, and some go to still greater lengths. (13/18. Dr. P. Lucas, 'Hered. Nat.' tome 2 pages 314, 892: see a good practical article on the subject in 'Gardener's Chronicle' 1856 page 620. I could add a vast number of references, but they would be superfluous.) But neither in the case of a breed which has been contaminated by a single cross, nor when, in the attempt to form an intermediate breed, half-bred animals have been matched together during many generations, can any rule be laid down how soon the tendency to reversion will be obliterated. It depends on the difference in the strength or prepotency of transmission in the two parent-forms, on their actual amount of difference, and on the nature of the conditions of life to which the crossed offspring are exposed. But we must be careful not to confound these cases of reversion to characters which were gained by a cross, with those under the first class, in which characters originally common to BOTH parents, but lost at some former period, reappear; for such characters may recur after an almost indefinite number of generations.
The law of reversion is as powerful with hybrids, when they are sufficiently fertile to breed together, or when they are repeatedly crossed with either pure parent-form, as in the case of mongrels. It is not necessary to give instances. With plants almost every one who has worked on this subject, from the time of Kolreuter to the present day, has insisted on this tendency. Gartner has recorded some good instances; but no one has given more striking ones than Naudin. (13/19. Kolreuter gives curious cases in his 'Dritte Fortsetzung' 1766 ss. 53, 59; and in his well-known 'Memoirs on Lavatera and Jalapa.' Gartner 'Bastarderzeugung' ss. 437, 441, etc. Naudin in his "Recherches sur l'Hybridite" 'Nouvelles Archives du Museum' tome 1 page 25.) The tendency differs in degree or strength in different groups, and partly depends, as we shall presently see, on whether the parent-plants have been long cultivated. Although the tendency to reversion is extremely general with nearly all mongrels and hybrids, it cannot be considered as invariably characteristic of them; it may also be mastered by long-continued selection; but these subjects will more properly be discussed in a future chapter on Crossing. From what we see of the power and scope of reversion, both in pure races, and when varieties or species are crossed, we may infer that characters of almost every kind are capable of reappearing after having been lost for a great length of time. But it does not follow from this that in each particular case certain characters will reappear; for instance, this will not occur when a race is crossed with another endowed with prepotency of transmission. Sometimes the power of reversion wholly fails, without our being able to assign any cause for the failure: thus it has been stated that in a French family in which 85 out of above 600 members, during six generations, had been subject to night-blindness, "there has not been a single example of this affection in the children of parents who were themselves free from it." (13/20. Quoted by Mr. Sedgwick in 'Med. — Chirurg. Review' April 1861 page 485. Dr. H. Dobell in 'Med. — Chirurg. Transactions' volume 46 gives an analogous case in which, in a large family, fingers with thickened joints were transmitted to several members during five generations; but when the blemish once disappeared it never reappeared.)
REVERSION THROUGH BUD-PROPAGATION — PARTIAL REVERSION, BY SEGMENTS IN THE SAME FLOWER OR FRUIT, OR IN DIFFERENT PARTS OF THE BODY IN THE SAME INDIVIDUAL ANIMAL.
In the eleventh chapter many cases of reversion by buds, independently of seminal generation, were given — as when a leaf-bud on a variegated, a curled, or laciniated variety suddenly reassumes its proper character; or as when a Provence-rose appears on a moss-rose, or a peach on a nectarine-tree. In some of these cases only half the flower or fruit, or a smaller segment, or mere stripes, reassume their former character; and here we have reversion by segments. Vilmorin (13/21. Verlot 'Des Varietes' 1865 page 63.) has also recorded several cases with plants derived from seed, of flowers reverting by stripes or blotches to their primitive colours: he states that in all such cases a white or pale-coloured variety must first be formed, and, when this is propagated for a length of time by seed, striped seedlings occasionally make their appearance; and these can afterwards by care be multiplied by seed.
The stripes and segments just referred to are not due, as far as is known, to reversion to characters derived from a cross, but to characters lost by variation. These cases, however, as Naudin (13/22. 'Nouvelles Archives du Museum' tome 1 page 25. Alex. Braun (in his 'Rejuvenescence' Ray Soc. 1853 page 315) apparently holds a similar opinion.) insists in his discussion on disjunction of character, are closely analogous with those given in the eleventh chapter, in which crossed plants have been known to produce half-and- half or striped flowers and fruit, or distinct kinds of flowers on the same root resembling the two parent-forms. Many piebald animals probably come under this same head. Such cases, as we shall see in the chapter on Crossing, apparently result from certain characters not readily blending together, and, as a consequence of this incapacity for fusion, the offspring either perfectly resemble one of their two parents, or resemble one parent in one part, and the other parent in another part; or whilst young are intermediate in character, but with advancing age revert wholly or by segments to either parent-form, or to both. Thus, young trees of the Cytisus adami are intermediate in foliage and flowers between the two parent-forms; but when older the buds continually revert either partially or wholly to both forms. The cases given in the eleventh chapter on the changes which occurred during growth in crossed plants of Tropaeolum, Cereus, Datura, and Lathyrus are all analogous. As, however, these plants are hybrids of the first generation, and as their buds after a time come to resemble their parents and not their grandparents, these cases do not at first appear to come under the law of reversion in the ordinary sense of the word; nevertheless, as the change is effected through a succession of bud-generations on the same plant, they may be thus included.
Analogous facts have been observed in the animal kingdom, and are more remarkable, as they occur in the same individual in the strictest sense, and not as with plants through a succession of bud-generations. With animals the act of reversion, if it can be so designated, does not pass over a true generation, but merely over the early stages of growth in the same individual. For instance, I crossed several white hens with a black cock, and many of the chickens were, during the first year, perfectly white, but acquired during the second year black feathers; on the other hand, some of the chickens which were at first black, became during the second year piebald with white. A great breeder (13/23. Mr. Teebay in 'The Poultry Book' by Mr. Tegetmeier 1866 page 72.) says, that a Pencilled Brahma hen which has any of the blood of the Light Brahma in her, will "occasionally produce a pullet well pencilled during the first year, but she will most likely moult brown on the shoulders and become quite unlike her original colours in the second year." The same thing occurs with light Brahmas if of impure blood. I have observed exactly similar cases with the crossed offspring from differently coloured pigeons. But here is a more remarkable fact: I crossed a turbit, which has a frill formed by the feathers being reversed on its breast, with a trumpeter; and one of the young pigeons thus raised at first showed not a trace of the frill, but, after moulting thrice, a small yet unmistakably distinct frill appeared on its breast. According to Girou (13/24. Quoted by Hofacker 'Ueber die Eigenschaften' etc. s. 98.) calves produced from a red cow by a black bull, or from a black cow by a red bull, are not rarely born red, and subsequently become black. I possess a dog, the daughter of a white terrier by a fox- coloured bulldog; as a puppy she was quite white, but when about six months old a black spot appeared on her nose, and brown spots on her ears. When a little older she was badly wounded on the back, and the hair which grew on the cicatrix was of a brown colour, apparently derived from her father. This is the more remarkable, as with most animals having coloured hair, that which grows on a wounded surface is white.
In the foregoing cases, the characters which with advancing age reappeared, were present in the immediately preceding generations; but characters sometimes reappear in the same manner after a much longer interval of time. Thus the calves of a hornless race of cattle which originated in Corrientes, though at first quite hornless, as they become adult sometimes acquire small, crooked, and loose horns; and these in succeeding years occasionally become attached to the skull. (13/25. Azara 'Essais Hist. Nat. de Paraguay' tome 2 1801 page 372.) White and black Bantams, both of which generally breed true, sometimes assume as they grow old a saffron or red plumage. For instance, a first-rate black bantam has been described, which during three seasons was perfectly black, but then annually became more and more red; and it deserves notice that this tendency to change, whenever it occurs in a bantam, "is almost certain to prove hereditary." (13/26. These facts are given on the high authority of Mr. Hewitt in 'The Poultry Book' by Mr. Tegetmeier 1866 page 248.) The cuckoo or blue-mottled Dorking cock, when old, is liable to acquire yellow or orange hackles in place of his proper bluish-grey hackles. (13/27. 'The Poultry Book' by Tegetmeier 1866 page 97.) Now as Gallus bankiva is coloured red and orange, and as Dorking fowls and bantams are descended from this species, we can hardly doubt that the change which occasionally occurs in the plumage of these birds as their age advances, results from a tendency in the individual to revert to the primitive type.
CROSSING AS A DIRECT CAUSE OF REVERSION.
It has long been notorious that hybrids and mongrels often revert to both or to one of their parent-forms, after an interval of from two to seven or eight, or, according to some authorities, even a greater number of generations. But that the act of crossing in itself gives an impulse towards reversion, as shown by the reappearance of long-lost characters, has never, I believe, been hitherto proved. The proof lies in certain peculiarities, which do not characterise the immediate parents, and therefore cannot have been derived from them, frequently appearing in the offspring of two breeds when crossed, which peculiarities never appear, or appear with extreme rarity, in these same breeds, as long as they are precluded from crossing. As this conclusion seems to me highly curious and novel, I will give the evidence in detail.
[My attention was first called to this subject, and I was led to make numerous experiments, by MM. Boitard and Corbie having stated that, when they crossed certain breeds of pigeons, birds coloured like the wild C. livia, or the common dovecote — namely, slaty-blue, with double black wing-bars, sometimes chequered with black, white loins, the tail barred with black, with the outer feathers edged with white, — were almost invariably produced. The breeds which I crossed, and the remarkable results attained, have been fully described in the sixth chapter. I selected pigeons belonging to true and ancient breeds, which had not a trace of blue or any of the above specified marks; but when crossed, and their mongrels recrossed, young birds were often produced, more or less plainly coloured slaty-blue, with some or all of the proper characteristic marks. I may recall to the reader's memory one case, namely, that of a pigeon, hardly distinguishable from the wild Shetland species, the grandchild of a red-spot, white fantail, and two black barbs, from any of which, when purely-bred, the production of a pigeon coloured like the wild C. livia would have been almost a prodigy.
I was thus led to make the experiments, recorded in the seventh chapter, on fowls. I selected long-established pure breeds, in which there was not a trace of red, yet in several of the mongrels feathers of this colour appeared; and one magnificent bird, the offspring of a black Spanish cock and white Silk hen, was coloured almost exactly like the wild Gallus bankiva. All who know anything of the breeding of poultry will admit that tens of thousands of pure Spanish and of pure white Silk fowls might have been reared without the appearance of a red feather. The fact, given on the authority of Mr. Tegetmeier, of the frequent appearance, in mongrel fowls, of pencilled or transversely-barred feathers, like those common to many gallinaceous birds, is likewise apparently a case of reversion to a character formerly possessed by some ancient progenitor of the family. I owe to the kindness of this excellent observer the opportunity of inspecting some neck-hackles and tail-feathers from a hybrid between the common fowl and a very distinct species, the Gallus varius; and these feathers are transversely striped in a conspicuous manner with dark metallic blue and grey, a character which could not have been derived from either immediate parent.
I have been informed by Mr. B.P. Brent, that he crossed a white Aylesbury drake and a black so-called Labrador duck, both of which are true breeds, and he obtained a young drake closely like the mallard (A. boschas). Of the musk- duck (Cairina moschata, Linn.) there are two sub-breeds, namely, white and slate-coloured; and these I am informed breed true, or nearly true. But the Rev. W.D. Fox tells me that, by putting a white drake to a slate-coloured duck, black birds, pied with white, like the wild musk-duck, were always produced. I hear from Mr. Blyth that hybrids from the canary and gold-finch almost always have streaked feathers on their backs; and this streaking must be derived from the original wild canary.
We have seen in the fourth chapter, that the so-called Himalayan rabbit, with its snow-white body, black ears, nose, tail, and feet, breeds perfectly true. This race is known to have been formed by the union of two varieties of silver-grey rabbits. Now, when a Himalayan doe was crossed by a sandy-coloured buck, a silver-grey rabbit was produced; and this is evidently a case of reversion to one of the parent varieties. The young of the Himalayan rabbit are born snow-white, and the dark marks do not appear until some time subsequently; but occasionally young Himalayan rabbits are born of a light silver-grey, which colour soon disappears; so that here we have a trace of reversion, during an early period of life, to the parent varieties, independently of any recent cross.
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