but without being able to assign any cause, that, when a new character appears, it is occasionally from the first well fixed, or fluctuates much, or wholly fails to be transmitted. So it is with the aggregate of slight differences which characterise a new variety, for some propagate their kind from the first much truer than others. Even with plants multiplied by bulbs, layers, &c., which may in one sense be said to form parts of the same individual, it is well known that certain varieties retain and transmit through successive bud-generations their newly-acquired characters more truly than others. In none of these, nor in the following cases, does there appear to be any relation between the force with which a character is transmissible and the length of time during which it has already been transmitted. Some varieties, such as white and yellow hyacinths and white sweet-peas, transmit their colours more faithfully than do the varieties which have retained their natural colour. In the Irish family, mentioned in the twelfth chapter, the peculiar tortoiseshell-like colouring of the eyes was transmitted far more faithfully than any ordinary colour. Ancon and Mauchamp sheep and niata cattle, which are all comparatively modern breeds, exhibit remarkably strong powers of inheritance. Many similar cases could be adduced.
As all domesticated animals and cultivated plants have varied, and yet are descended from aboriginally wild forms, which no doubt had retained the same character from an immensely remote epoch, we see that scarcely any degree of antiquity ensures a character being transmitted perfectly true. In this case, however, it may be said that changed conditions of life induce certain modifications, and not that the power of inheritance fails; but in every case of failure, some cause, either internal or external, must interfere. It will generally be found that the parts in our domesticated productions which have varied, or which still continue to vary,—that is, which fail to retain their primordial state,—are the same with the parts which differ in the natural species of the same genus. As, on the theory of descent with modification, the species of the same genus have been modified since they branched off from a common progenitor, it follows that the characters by which they differ from each other have varied whilst other parts of the organisation have remained unchanged; and it might be argued that
these same characters now vary under domestication, or fail to be inherited, owing to their lesser antiquity. But we must believe structures, which have already varied, would be more liable to go on varying, rather than structures which during an immense lapse of time have remained unaltered; and this variation is probably the result of certain relations between the conditions of life and the organisation, quite independently of the greater or less antiquity of each particular character.
Fixedness of character, or the strength of inheritance, has often been judged of by the preponderance of certain characters in the crossed offspring between distinct races; but prepotency of transmission here comes into play, and this, as we shall immediately see, is a very different consideration from the strength or weakness of inheritance. It has often been observed[[136]] that breeds of animals inhabiting wild and mountainous countries cannot be permanently modified by our improved breeds; and as these latter are of modern origin, it has been thought that the greater antiquity of the wilder breeds has been the cause of their resistance to improvement by crossing; but it is more probably due to their structure and constitution being better adapted to the surrounding conditions. When plants are first subjected to culture, it has been found that, during several generations, they transmit their characters truly, that is, do not vary, and this has been attributed to ancient characters being strongly inherited; but it may with equal or greater probability be consequent on changed conditions of life requiring a long time for their accumulative action. Notwithstanding these considerations, it would perhaps be rash to deny that characters become more strongly fixed the longer they are transmitted; but I believe that the proposition resolves itself into this,—that all characters of all kinds, whether new or old, tend to be inherited, and that those which have already withstood all counteracting 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 believed[[139]] to have had a prepotent influence on the shorthorn race. It has also been observed[[140]] 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 Buzareingues[[142]] 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 said[[144]] 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 Corbié[[145]] 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 states[[147]] 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 shown[[150]] 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 upright figure on these hybrids.
I am aware that such cases as the foregoing have been ascribed by various authors, not to one species, race, or individual being prepotent over the other in impressing it character on its crossed offspring, but to such rules as that the father influences the external characters and the mother the internal or vital organs. But the great diversity of the rules given by various authors almost proves their falseness. Dr. Prosper Lucas has fully discussed this point, and has shown[[153]] that none of the rules (and I could add others to those quoted by him) apply to all animals. Similar rules have been enounced for plants, and have been proved by Gärtner[[154]] to be all erroneous. If we confine our view to the domesticated races of a single species, or perhaps even to the species of the same genus, some such rules may hold good; for instance, it seems that in reciprocally crossing various breeds of fowls the male generally gives colour;[[155]] but conspicuous exceptions have passed under my own eyes. In sheep it seems that the ram usually gives its peculiar horns and fleece to its crossed offspring, and the bull the presence or absence of horns.
In the following chapter on Crossing I shall have occasion to show that certain characters are rarely or never blended by crossing, but are transmitted in an unmodified state from either parent-form; I refer to this fact here because it is sometimes accompanied on the one side by prepotency, which thus acquires the false appearance of unusual strength. In the same chapter I shall show that the rate at which a species or breed absorbs and obliterates another by repeated crosses, depends in chief part on prepotency in transmission.
In conclusion, some of the cases above given,—for instance, that of the trumpeter pigeon,—prove that there is a wide difference between mere inheritance and prepotency. This latter power seems to us, in our ignorance, to act in most cases quite capriciously. The very same character, even though it be an abnormal or monstrous one, such as silky feathers, may be transmitted by different species, when crossed, either with prepotent force or singular feebleness. It is obvious, that a purely-bred form of either sex, in all cases in which prepotency does not run more strongly in one sex than the other, will transmit its character with prepotent force over a mongrelized and already variable form.[[156]] From several of the above-given cases we may conclude that mere antiquity of character does not by any means necessarily make it prepotent. In some cases prepotency apparently depends on the same character being present and visible in one of the two breeds which are crossed, and latent or invisible in the other breed; and in this case it is natural that the character which is potentially present in both should be prepotent. Thus, we have reason to believe that there is a latent tendency in all horses to be dun-coloured and striped; and when a horse of this kind is crossed with one of any other colour, it is said that the offspring are almost sure to be striped. Sheep have a similar latent tendency to become dark-coloured, and we have seen with what prepotent force a ram with a few black spots, when crossed with sheep of various breeds, coloured its offspring. All pigeons have a latent tendency to become slaty-blue, with certain characteristic marks, and it is known that, when a bird thus coloured is crossed with one of any other colour, it is most difficult afterwards to eradicate the blue tint. A nearly parallel case is offered by those black bantams which, as they grow