DEFINITION OF CLOSE INTERBREEDING — AUGMENTATION OF MORBID TENDENCIES — GENERAL EVIDENCE OF THE GOOD EFFECTS DERIVED FROM CROSSING, AND ON THE EVIL EFFECTS FROM CLOSE INTERBREEDING — CATTLE, CLOSELY INTERBRED; HALF-WILD CATTLE LONG KEPT IN THE SAME PARKS — SHEEP — FALLOW-DEER — DOGS, RABBITS, PIGS — MAN, ORIGIN OF HIS ABHORRENCE OF INCESTUOUS MARRIAGES — FOWLS — PIGEONS — HIVE-BEES — PLANTS, GENERAL CONSIDERATIONS ON THE BENEFITS DERIVED FROM CROSSING — MELONS, FRUIT-TREES, PEAS, CABBAGES, WHEAT, AND FOREST-TREES — ON THE INCREASED SIZE OF HYBRID PLANTS, NOT EXCLUSIVELY DUE TO THEIR STERILITY — ON CERTAIN PLANTS WHICH EITHER NORMALLY OR ABNORMALLY ARE SELF-IMPOTENT, BUT ARE FERTILE, BOTH ON THE MALE AND FEMALE SIDE, WHEN CROSSED WITH DISTINCT INDIVIDUALS EITHER OF THE SAME OR ANOTHER SPECIES — CONCLUSION.
The gain in constitutional vigour, derived from an occasional cross between individuals of the same variety, but belonging to distinct families, or between distinct varieties, has not been so largely or so frequently discussed, as have the evil effects of too close interbreeding. But the former point is the more important of the two, inasmuch as the evidence is more decisive. The evil results from close interbreeding are difficult to detect, for they accumulate slowly, and differ much in degree with different species; whilst the good effects which almost invariably follow a cross are from the first manifest. It should, however, be clearly understood that the advantage of close interbreeding, as far as the retention of character is concerned, is indisputable, and often outweighs the evil of a slight loss of constitutional vigour. In relation to the subject of domestication, the whole question is of some importance, as too close interbreeding interferes with the improvement of old races. It is important as indirectly bearing on Hybridism; and possibly on the extinction of species, when any form has become so rare that only a few individuals remain within a confined area. It bears in an important manner on the influence of free intercrossing, in obliterating individual differences, and thus giving uniformity of character to the individuals of the same race or species; for if additional vigour and fertility be thus gained, the crossed offspring will multiply and prevail, and the ultimate result will be far greater than otherwise would have occurred. Lastly, the question is of high interest, as bearing on mankind. I shall therefore discuss this subject at full length. As the facts which prove the evil effects of close interbreeding are more copious, though less decisive, than those on the good effects of crossing, I shall, under each group of beings, begin with the former.
There is no difficulty in defining what is meant by a cross; but this is by no means easy in regard to "breeding in and in" or "too close interbreeding," because, as we shall see, different species of animals are differently affected by the same degree of interbreeding. The pairing of a father and daughter, or mother and son, or brothers and sisters, if carried on during several generations, is the closest possible form of interbreeding. But some good judges, for instance Sir J. Sebright, believe that the pairing of a brother and sister is much closer than that of parents and children; for when the father is matched with his daughter he crosses, as is said, with only half his own blood. The consequences of close interbreeding carried on for too long a time, are, as is generally believed, loss of size, constitutional vigour, and fertility, sometimes accompanied by a tendency to malformation. Manifest evil does not usually follow from pairing the nearest relations for two, three, or even four generations; but several causes interfere with our detecting the evil—such as the deterioration being very gradual, and the difficulty of distinguishing between such direct evil and the inevitable augmentation of any morbid tendencies which may be latent or apparent in the related parents. On the other hand, the benefit from a cross, even when there has not been any very close interbreeding, is almost invariably at once conspicuous. There is good reason to believe, and this was the opinion of that most experienced observer Sir J. Sebright, that the evil effects of close interbreeding may be checked or quite prevented by the related individuals being separated for a few generations and exposed to different conditions of life. This conclusion is now held by many breeders; for instance Mr. Carr remarks, it is a well-known "fact that a change of soil and climate effects perhaps almost as great a change in the constitution as would result from an infusion of fresh blood." I hope to show in a future work that consanguinity by itself counts for nothing, but acts solely from related organisms generally having a similar constitution, and having been exposed in most cases to similar conditions.
That any evil directly follows from the closest interbreeding has been denied by many persons; but rarely by any practical breeder; and never, as far as I know, by one who has largely bred animals which propagate their kind quickly. Many physiologists attribute the evil exclusively to the combination and consequent increase of morbid tendencies common to both parents; and that this is an active source of mischief there can be no doubt. It is unfortunately too notorious that men and various domestic animals endowed with a wretched constitution, and with a strong hereditary disposition to disease, if not actually ill, are fully capable of procreating their kind. Close interbreeding, on the other hand, often induces sterility; and this indicates something quite distinct from the augmentation of morbid tendencies common to both parents. The evidence immediately to be given convinces me that it is a great law of nature, that all organic beings profit from an occasional cross with individuals not closely related to them in blood; and that, on the other hand, long-continued close interbreeding is injurious.
Various general considerations have had much influence in leading me to this conclusion; but the reader will probably rely more on special facts and opinions. The authority of experienced observers, even when they do not advance the grounds of their belief, is of some little value. Now almost all men who have bred many kinds of animals and have written on the subject, such as Sir J. Sebright, Andrew Knight, etc., have expressed the strongest conviction on the impossibility of long-continued close interbreeding. Those who have compiled works on agriculture, and have associated much with breeders, such as the sagacious Youatt, Low, etc., have strongly declared their opinion to the same effect. Prosper Lucas, trusting largely to French authorities, has come to a similar conclusion. The distinguished German agriculturist Hermann von Nathusius, who has written the most able treatise on this subject which I have met with, concurs; and as I shall have to quote from this treatise, I may state that Nathusius is not only intimately acquainted with works on agriculture in all languages, and knows the pedigrees of our British breeds better than most Englishmen, but has imported many of our improved animals, and is himself an experienced breeder.
Evidence of the evil effects of close interbreeding can most readily be acquired in the case of animals, such as fowls, pigeons, etc., which propagate quickly, and, from being kept in the same place, are exposed to the same conditions. Now I have inquired of very many breeders of these birds, and I have hitherto not met with a single man who was not thoroughly convinced that an occasional cross with another strain of the same sub-variety was absolutely necessary. Most breeders of highly improved or fancy birds value their own strain, and are most unwilling, at the risk, in their opinion, of deterioration, to make a cross. The purchase of a first-rate bird of another strain is expensive, and exchanges are troublesome; yet all breeders, as far as I can hear, excepting those who keep large stocks at different places for the sake of crossing, are driven after a time to take this step.
Another general consideration which has had great influence on my mind is, that with all hermaphrodite animals and plants, which it might have been thought would have perpetually fertilised themselves and been thus subjected for long ages to the closest interbreeding, there is not a single species, as far as I can discover, in which the structure ensures self-fertilisation. On the contrary, there are in a multitude of cases, as briefly stated in the fifteenth chapter, manifest adaptations which favour or inevitably lead to an occasional cross between one hermaphrodite and another of the same species; and these adaptive structures are utterly purposeless, as far as we can see, for any other end.
With Cattle there can be no doubt that extremely close interbreeding may be long carried on advantageously with respect to external characters, and with no manifest evil as far as constitution is concerned. The case of Bakewell's Longhorns, which were closely interbred for a long period, has often been quoted; yet Youatt says the breed "had acquired a delicacy of constitution inconsistent with common management," and "the propagation of the species was not always certain." But the Shorthorns offer the most striking case of close interbreeding; for instance, the famous bull Favourite (who was himself the offspring of a half-brother and sister from Foljambe) was matched with his own daughter, granddaughter, offspring of a half-brother and sister from Foljambe) was matched with his own daughter, granddaughter, 16ths, or 93·75 per cent of the blood of Favourite in her veins. This cow was matched with the bull Wellington, having 62·5 per cent of Favourite blood in his veins, and produced Clarissa; Clarissa was matched with the bull Lancaster, having 68·75 of the same blood, and she yielded valuable offspring. Nevertheless Collings, who reared these animals, and was a strong advocate for close breeding, once crossed his stock with a Galloway, and the cows from this cross realised the highest prices. Bates's herd was esteemed the most celebrated in the world. For thirteen years he bred most closely in and in; but during the next seventeen years, though he had the most exalted notion of the value of his own stock, he thrice infused fresh blood into his herd: it is said that he did this, not to improve the form of his animals, but on account of their lessened fertility. Mr. Bates's own view, as given by a celebrated was, that "to breed inand-in from a bad stock was ruin and devastation; yet that the practice may be safely followed within certain limits when the parents so related are descended from first-rate animals." We thus see that there has been much close interbreeding with Shorthorns; but Nathusius, after the most careful study of their pedigrees, says that he can find no instance of a breeder who has strictly followed this practice during his whole life. From this study and his own experience, he concludes that close interbreeding is necessary to ennoble the stock; but that in effecting this the greatest care is necessary, on account of the tendency to infertility and weakness. It may be added, that another high authority asserts that many more calves are born cripples from Shorthorns than from other and less closely interbred races of cattle.
Although by carefully selecting the best animals (as Nature effectually does by the law of battle) close interbreeding may be long carried on with cattle, yet the good effects of a cross between almost any twobreeds is at once shown by the greater size and vigour of the offspring; as Mr. Spooner writes to me, "crossing distinct breeds certainly improves cattle for the butcher." Such crossed animals are of course of no value to the breeder; but they have been raised during many years in several parts of England to be slaughtered; and their merit is now so fully recognised, that at fat-cattle shows a separate class has been formed for their reception. The best fat ox at the great show at Islington in 1862 was a crossed animal.
The half-wild cattle, which have been kept in British parks probably for 400 or 500 years, or even for a longer period, have been advanced by Culley and others as a case of long-continued interbreeding within the limits of the same herd without any consequent injury. With respect to the cattle at Chillingham, the late Lord Tankerville owned that they were bad breeders The agent, Mr. Hardy, estimates (in a letter to me, dated May, 1861) that in the herd of about fifty the average number annually slaughtered, killed by fighting, and dying, is about ten, or one in five. As the herd is kept up to nearly the same average number, the annual rate of increase must be likewise about one in five. The bulls, I may add, engage in furious battles, of which battles the present Lord Tankerville has given me a graphic description, so that there will always be rigorous selection of the most vigorous males. I procured in 1855 from Mr. D. Gardner, agent to the Duke of Hamilton, the following account of the wild cattle kept in the Duke's park in Lanarkshire, which is about 200 acres in extent. The number of cattle varies from sixty-five to eighty; and the number annually killed (I presume by all causes) is from eight to ten; so that the annual rate of increase can hardly be more than one in six. Now in South America, where the herds are half-wild, and therefore offer a nearly fair standard of comparison, according to Azara the natural increase of the cattle on an estancia is from onethird to one-fourth of the total number, or one in between three and four and this, no doubt, applies exclusively to adult animals fit for consumption. Hence the half-wild British cattle which have long interbred within the limits of the same herd are relatively far less fertile. Although in an unenclosed country like Paraguay there must be some crossing between the different herds, yet even there the inhabitants believe that the occasional introduction of animals from distant localities is necessary to prevent "degeneration in size and diminution of fertility." The decrease in size from ancient times in the Chillingham and Hamilton cattle must have been prodigious, for Professor Rütimeyer has shown that they are almost certainly the descendants of the gigantic Bos primigenius. No doubt this decrease in size may be largely attributed to less favourable conditions of life; yet animals roaming over large parks, and fed during severe winters, can hardly be considered as placed under very unfavourable conditions.
With Sheep there has often been long-continued interbreeding within the limits of the same flock; but whether the nearest relations have been matched so frequently as in the case of Shorthorn cattle, I do not know. The Messrs. Brown during fifty years have never infused fresh blood into their excellent flock of Leicesters. Since 1810 Mr. Barford has acted on the same principle with the Foscote flock. He asserts that half a century of experience has convinced him that when two nearly related animals are quite sound in constitution, in-and-in breeding does not induce degeneracy; but he adds that he "does not pride himself on breeding from the nearest affinities." In France the Naz flock has been bred for sixty years without the introduction of a single strange ram. Nevertheless, most great breeders of sheep have protested against close interbreeding prolonged for too great a length of time. The most celebrated of recent breeders, Jonas Webb, kept five separate families to work on, thus "retaining the requisite distance of relationship between the sexes"; and what is probably of greater importance, the separate flocks will have been exposed to somewhat different conditions.
Although by the aid of careful selection the near interbreeding of sheep may be long continued without any manifest evil, yet it has often been the practice with farmers to cross distinct breeds to obtain animals for the butcher, which plainly shows that good of some kind is derived from this practice. We have excellent evidence on this head from Mr. S. Druce, who gives in detail the comparative numbers of four pure breeds and of a cross-breed which can be supported on the same ground, and he gives their produce in fleece and carcase. A high authority, Mr. Pusey, sums up the result in money value during an equal length of time, namely (neglecting shillings), for Cotswolds 248l., for Leicesters 223l., for Southdowns 204l., for Hampshire Downs 264l., and for the crossbred 293l. A former celebrated breeder, Lord Somerville, states that his half-breeds from Ryelands and Spanish sheep were larger animals than either the pure Ryelands or pure Spanish sheep. Mr. Spooner concludes his excellent Essay on Crossing by asserting that there is a pecuniary advantage in judicious cross-breeding, especially when the male is larger than the female
As some of our British parks are ancient, it occurred to me that there must have been long-continued close interbreeding with the fallow-deer (Cervus dama) kept in them; but on inquiry I find that it is a common practice to infuse new blood by procuring bucks from other parks. Mr. Shirley, who has carefully studied the management of deer, admits that in some parks there has been no admixture of foreign blood from a time beyond the memory of man. But he concludes "that in the end the constant breeding in-and-in is sure to tell to the disadvantage of the whole herd, though it may take a very long time to prove it; moreover, when we find, as is very constantly the case, that the introduction of fresh blood has been of the very greatest use to deer, both by improving their size and appearance, and particularly by being of service in removing the taint of 'rickback,' if not of other diseases, to which deer are sometimes subject when the blood has not been changed, there can, I think, be no doubt but that a judicious cross with a good stock is of the greatest consequence, and is indeed essential, sooner or later, to the prosperity of every well-ordered park."
Mr. Meynell's famous foxhounds have been adduced, as showing that no ill effects follow from close interbreeding; and Sir J. Sebright ascertained from him that he frequently bred from father and daughter, mother and son, and sometimes even from brothers and sisters. With greyhounds also there has been much close interbreeding, but the best breeders agree that it may be carried too far. But Sir J. Sebright declares, that by breeding in-and-in, by which he means matching brothers and sisters, he has actually seen the offspring of strong spaniels degenerate into weak and diminutive lapdogs. The Rev. W. D. Fox has communicated to me the case of a small lot of bloodhounds, long kept in the same family, which had become very bad breeders, and nearly all had a bony enlargement in the tail. A single cross with a distinct strain of bloodhounds restored their fertility, and drove away the tendency to malformation in the tail. I have heard the particulars of another case with bloodhounds, in which the female had to be held to the male. Considering how rapid is the natural increase of the dog, it is difficult to understand the large price of all highly improved breeds, which almost implies long-continued close interbreeding, except on the belief that this process lessens fertility and increases liability to distemper and other diseases. A high authority, Mr. Scrope, attributes the rarity and deterioration in size of the Scotch deerhound (the few individuals formerly existing throughout the country being all related) in large part to close interbreeding.
With all highly-bred animals there is more or less difficulty in getting them to procreate quickly, and all suffer much from delicacy of constitution. A great judge of rabbits says, "the long-eared does are often too highly bred or forced in their youth to be of much value as breeders, often turning out barren or bad mothers." They often desert their young, so that it is necessary to have nurse-rabbits, but I do not pretend to attribute all these evil results to close interbreeding.
With respect to Pigs there is more unanimity amongst breeders on the evil effects of close interbreeding than, perhaps, with any other large animal. Mr. Druce, a great and successful breeder of the Improved Oxfordshires (a crossed race), writes, "without a change of boars of a different tribe, but of the same breed, constitution cannot be preserved." Mr. Fisher Hobbs, the raiser of the celebrated Improved Essex breed, divided his stock into three separate families, by which means he maintained the breed for more than twenty years, "by judicious selection from the three distinct families." Lord Western was the first importer of a Neapolitan boar and sow. "From this pair he bred in-and-in, until the breed was in danger of becoming extinct, a sure result (as Mr. Sidney remarks) of in-and-in breeding." Lord Western then crossed his Neapolitan pigs with the old Essex, and made the first great step towards the Improved Essex breed. Here is a more interesting case. Mr. J. Wright, well known as a breeder, crossed the same boar with the daughter, granddaughter, and great-granddaughter, and so on for seven generations. The result was, that in many instances the offspring failed to breed; in others they produced few that lived; and of the latter many were idiotic, without sense, even to suck, and when attempting to move could not walk straight. Now it deserves especial notice, that the two last sows produced by this long course of interbreeding were sent to other boars, and they bore several litters of healthy pigs. The best sow in external appearance produced during the whole seven generations was one in the last stage of descent; but the litter consisted of this one sow. She would not breed to her sire, yet bred at the first trial to a stranger in blood. So that, in Mr. Wright's case, long-continued and extremely close interbreeding did not affect the external form or merit of the young; but with many of them the general constitution and mental powers, and especially the reproductive functions, were seriously affected.
Nathusius gives an analogous and even more striking case: he imported from England a pregnant sow of the large Yorkshire breed, and bred the product closely in-and-in for three generations: the result was unfavourable, as the young were weak in constitution, with impaired fertility. One of the latest sows, which he esteemed a good animal, produced, when paired with her own uncle (who was known to be productive with sows of other breeds), a litter of six, and a second time a litter of only five weak young pigs. He then paired this sow with a boar of a small black breed, which he had likewise imported from England; this boar, when matched with sows of his own breed, produced from seven to nine young. Now, the sow of the large breed, which was so unproductive when paired with her own uncle, yielded to the small black boar, in the first litter twenty-one, and in the second litter eighteen young pigs; so that in one year she produced thirtynine fine young animals!
As in the case of several other animals already mentioned, even when no injury is perceptible from moderately close interbreeding, yet, to quote the words of Mr. Coate (who five times won the annual gold medal of the Smithfield Club Show for the best pen of pigs), "Crosses answer well for profit to the farmer, as you get more constitution and quicker growth; but for me, who sell a great number of pigs for breeding purposes, I find it will not do, as it requires many years to get anything like purity of blood again."
Almost all the animals as yet mentioned are gregarious, and the males must frequently pair with their own daughters, for they expel the young males as well as all intruders, until forced by old age and loss of strength to yield to some stronger male. It is therefore not improbable that gregarious animals may have been rendered less susceptible than non-social species to the evil consequences of close interbreeding, so that they may be enabled to live in herds without injury to their offspring. Unfortunately we do not know whether an animal like the cat, which is not gregarious, would suffer from close interbreeding in a greater degree than our other domesticated animals. But the pig is not, as far as I can discover, strictly gregarious, and we have seen that it appears eminently liable to the evil effects of close interbreeding. Mr. Huth, in the case of the pig, attributes (Chapter XXIV) these effects to their having been "cultivated most for their fat," or to the selected individuals having had a weak constitution; but we must remember that it is great breeders who have brought forward the above cases, and who are far more familiar than ordinary men can be, with the causes which are likely to interfere with the fertility of their animals.
The effects of close interbreeding in the case of man is a difficult subject, on which I will say but little. It has been discussed by various authors under many points of view Mr. Tylor has shown that with widely different races in the most distant quarters of the world, marriages between relations—even between distant relations—have been strictly prohibited. There are, however, many exceptions to the rule, which are fully given by Mr. Huth. It is a curious problem how these prohibitions arose during early and barbarous times. Mr. Tylor is inclined to attribute them to the evil effects of consanguineous marriages having been observed; and he ingeniously attempts to explain some apparent anomalies in the prohibition not extending equally to the relations on the male and female side. He admits, however, that other causes, such as the extension of friendly alliances, may have come into play. Mr. W. Adam, on the other hand, concludes that related marriages are prohibited and viewed with repugnance, from the confusion which would thus arise in the descent of property, and from other still more recondite reasons. But I cannot accept these views, seeing that incest is held in abhorrence by savages such as those of Australia and South America, who have no property to bequeath, or fine moral feelings to confuse, and who are not likely to reflect on distant evils to their progeny. According to Mr. Huth the feeling is the indirect result of exogamy, inasmuch as when this practice ceased in any tribe and it became endogamous, so that marriages were strictly confined to the same tribe, it is not unlikely that a vestige of the former practice would still be retained, so that closely-related marriages would be prohibited. With respect to exogamy itself Mr. MacLennan believes that it arose from a scarcity of women, owing to female infanticide, aided perhaps by other causes.
It has been clearly shown by Mr. Huth that there is no instinctive feeling in man against incest any more than in gregarious animals. We know also how readily any prejudice or feeling may rise to abhorrence, as shown by Hindus in regard to objects causing defilement. Although there seems to be no strong inherited feeling in mankind against incest, it seems possible that men during primeval times may have been more excited by strange females than by those with whom they habitually lived; in the same manner as according to Mr. Cupples, male deerhounds are inclined towards strange females, while the females prefer dogs with whom they have associated. If any such feeling formerly existed in man, this would have led to a preference for marriages beyond the nearest kin, and might have been strengthened by the offspring of such marriages surviving in greater numbers, as analogy would lead us to believe would have occurred.
Whether consanguineous marriages, such as are permitted in civilised nations, and which would not be considered as close interbreeding in the case of our domesticated animals, cause any injury will never be known with certainty until a census is taken with this object in view. My son, George Darwin, has done what is possible at present by a statistical investigation and he has come to the conclusion, from his own researches and those of Dr. Mitchell, that the evidence as to any evil thus caused is conflicting, but on the whole points to the evil being very small.
Birds. —In the case of the Fowl a whole array of authorities could be given against too close interbreeding. Sir J. Sebright positively asserts that he made many trials, and that his fowls, when thus treated, became long in the legs, small in the body, and bad breeders. He produced the famous Sebright Bantams by complicated crosses, and by breeding in-and-in; and since his time there has been much close interbreeding with these animals; and they are now notoriously bad breeders. I have seen Silver Bantams, directly descended from his stock, which had become almost as barren as hybrids; for not a single chicken had been that year hatched from two full nests of eggs. Mr. Hewitt says that with these Bantams the sterility of the male stands, with rare exceptions, in the closest relation with their loss of certain secondary male characters: he adds, "I have noticed, as a general rule, that even the slightest deviation from feminine character in the tail of the male Sebright—say the elongation by only half an inch of the two principal tail feathers—brings with it improved probability of increased fertility."
An experienced writer remarks that the same amateur, as is well known, seldom long maintains the superiority of his birds; and this, he adds, undoubtedly is due to all his stock "being of the same blood;" hence it is indispensable that he should occasionally procure a bird of another strain. But this is not necessary with those who keep a stock of fowls at different stations. Thus, Mr. Ballance, who has bred Malays for thirty years, and has won more prizes with these birds than any other fancier in England, says that breeding in-and-in does not necessarily cause deterioration; "but all depends upon how this is managed. My plan has been to keep about five or six distinct runs, and to rear about two hundred or three hundred chickens each year, and select the best birds from each run for crossing. I thus secure sufficient crossing to prevent deterioration."
Mr. Wright states33 that Mr. Clark, "whose fighting-cocks were so notorious, continued to breed from his own kind till they lost their disposition to fight, but stood to be cut up without making any resistance, and were so reduced in size as to be under those weights required for the best prizes; but on obtaining a cross from Mr. Leighton, they again resumed their former courage and weight." It should be borne in mind that game-cocks before they fought were always weighed, so that nothing was left to the imagination about any reduction or increase of weight. Mr. Clark does not seem to have bred from brothers and sisters, which is the most injurious kind of union; and he found, after repeated trials, that there was a greater reduction in weight in the young from a father paired with his daughter, than from a mother with her son. I may add that Mr. Eyton of Eyton, th