On the Origin of Species by Means of Natural Selection
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Chapter VIII


Distinction between the sterility of first crosses and of hybrids -- Sterility various in degree, not universal, affected by close interbreeding, removed by domestication -- Laws governing the sterility of hybrids -- Sterility not a special endowment, but incidental on other differences -- Causes of the sterility of first crosses and of hybrids -- Parallelism between the effects of changed conditions of life and crossing -- Fertility of varieties when crossed and of their mongrel offspring not universal -- Hybrids and mongrels compared independently of their fertility -- Summary.

The view generally entertained by naturalists is that species, when intercrossed, have been specially endowed with the quality of sterility, in order to prevent the confusion of all organic forms. This view certainly seems at first probable, for species within the same country could hardly have kept distinct had they been capable of crossing freely. The importance of the fact that hybrids are very generally sterile, has, I think, been much underrated by some late writers. On the theory of natural selection the case is especially important, inasmuch as the sterility of hybrids could not possibly be of any advantage to them, and therefore could not have been acquired by the continued preservation of successive profitable degrees of sterility. I hope, however, to be able to show that sterility is not a specially acquired or endowed quality, but is incidental on other acquired differences.

In treating this subject, two classes of facts, to a large extent fundamentally different, have generally been confounded together; namely, the sterility of two species when first crossed, and the sterility of the hybrids produced from them.

Pure species have of course their organs of reproduction in a perfect condition, yet when intercrossed they produce either few or no offspring. Hybrids, on the other hand, have their reproductive organs functionally impotent, as may be clearly seen in the state of the male element in both plants and animals; though the organs themselves are perfect in structure, as far as the microscope reveals. In the first case the two sexual elements which go to form the embryo are perfect; in the second case they are either not at all developed, or are imperfectly developed. This distinction is important, when the cause of the sterility, which is common to the two cases, has to be considered. The distinction has probably been slurred over, owing to the sterility in both cases being looked on as a special endowment, beyond the province of our reasoning powers.

The fertility of varieties, that is of the forms known or believed to have descended from common parents, when intercrossed, and likewise the fertility of their mongrel offspring, is, on my theory, of equal importance with the sterility of species; for it seems to make a broad and clear distinction between varieties and species.

First, for the sterility of species when crossed and of their hybrid offspring. It is impossible to study the several memoirs and works of those two conscientious and admirable observers, Kolreuter and Gartner, who almost devoted their lives to this subject, without being deeply impressed with the high generality of some degree of sterility. Kolreuter makes the rule universal; but then he cuts the knot, for in ten cases in which he found two forms, considered by most authors as distinct species, quite fertile together, he unhesitatingly ranks them as varieties. Gartner, also, makes the rule equally universal; and he disputes the entire fertility of Kolreuter's ten cases. But in these and in many other cases, Gartner is obliged carefully to count the seeds, in order to show that there is any degree of sterility. He always compares the maximum number of seeds produced by two species when crossed and by their hybrid offspring, with the average number produced by both pure parent-species in a state of nature. But a serious cause of error seems to me to be here introduced: a plant to be hybridised must be castrated, and, what is often more important, must be secluded in order to prevent pollen being brought to it by insects from other plants. Nearly all the plants experimentised on by Gartner were potted, and apparently were kept in a chamber in his house. That these processes are often injurious to the fertility of a plant cannot be doubted; for Gartner gives in his table about a score of cases of plants which he castrated, and artificially fertilised with their own pollen, and (excluding all cases such as the Leguminosae, in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired. Moreover, as Gartner during several years repeatedly crossed the primrose and cowslip, which we have such good reason to believe to be varieties, and only once or twice succeeded in getting fertile seed; as he found the common red and blue pimpernels (Anagallis arvensis and coerulea), which the best botanists rank as varieties, absolutely sterile together; and as he came to the same conclusion in several other analogous cases; it seems to me that we may well be permitted to doubt whether many other species are really so sterile, when intercrossed, as Gartner believes.

It is certain, on the one hand, that the sterility of various species when crossed is so different in degree and graduates away so insensibly, and, on the other hand, that the fertility of pure species is so easily affected by various circumstances, that for all practical purposes it is most difficult to say where perfect fertility ends and sterility begins. I think no better evidence of this can be required than that the two most experienced observers who have ever lived, namely, Kolreuter and Gartner, should have arrived at diametrically opposite conclusions in regard to the very same species. It is also most instructive to compare--but I have not space here to enter on details--the evidence advanced by our best botanists on the question whether certain doubtful forms should be ranked as species or varieties, with the evidence from fertility adduced by different hybridisers, or by the same author, from experiments made during different years. It can thus be shown that neither sterility nor fertility affords any clear distinction between species and varieties; but that the evidence from this source graduates away, and is doubtful in the same degree as is the evidence derived from other constitutional and structural differences.

In regard to the sterility of hybrids in successive generations; though Gartner was enabled to rear some hybrids, carefully guarding them from a cross with either pure parent, for six or seven, and in one case for ten generations, yet he asserts positively that their fertility never increased, but generally greatly decreased. I do not doubt that this is usually the case, and that the fertility often suddenly decreases in the first few generations. Nevertheless I believe that in all these experiments the fertility has been diminished by an independent cause, namely, from close interbreeding. I have collected so large a body of facts, showing that close interbreeding lessens fertility, and, on the other hand, that an occasional cross with a distinct individual or variety increases fertility, that I cannot doubt the correctness of this almost universal belief amongst breeders. Hybrids are seldom raised by experimentalists in great numbers; and as the parent-species, or other allied hybrids, generally grow in the same garden, the visits of insects must be carefully prevented during the flowering season: hence hybrids will generally be fertilised during each generation by their own individual pollen; and I am convinced that this would be injurious to their fertility, already lessened by their hybrid origin. I am strengthened in this conviction by a remarkable statement repeatedly made by Gartner, namely, that if even the less fertile hybrids be artificially fertilised with hybrid pollen of the same kind, their fertility, notwithstanding the frequent ill effects of manipulation, sometimes decidedly increases, and goes on increasing. Now, in artificial fertilisation pollen is as often taken by chance (as I know from my own experience) from the anthers of another flower, as from the anthers of the flower itself which is to be fertilised; so that a cross between two flowers, though probably on the same plant, would be thus effected. Moreover, whenever complicated experiments are in progress, so careful an observer as Gartner would have castrated his hybrids, and this would have insured in each generation a cross with the pollen from a distinct flower, either from the same plant or from another plant of the same hybrid nature. And thus, the strange fact of the increase of fertility in the successive generations of artificially fertilised hybrids may, I believe, be accounted for by close interbreeding having been avoided.

Now let us turn to the results arrived at by the third most experienced hybridiser, namely, the Hon. and Rev. W. Herbert. He is as emphatic in his conclusion that some hybrids are perfectly fertile--as fertile as the pure parent-species--as are Kolreuter and Gartner that some degree of sterility between distinct species is a universal law of nature. He experimentised on some of the very same species as did Gartner. The difference in their results may, I think, be in part accounted for by Herbert's great horticultural skill, and by his having hothouses at his command. Of his many important statements I will here give only a single one as an example, namely, that 'every ovule in a pod of Crinum capense fertilised by C. revolutum produced a plant, which (he says) I never saw to occur in a case of its natural fecundation.' So that we here have perfect, or even more than commonly perfect, fertility in a first cross between two distinct species.

This case of the Crinum leads me to refer to a most singular fact, namely, that there are individual plants, as with certain species of Lobelia, and with all the species of the genus Hippeastrum, which can be far more easily fertilised by the pollen of another and distinct species, than by their own pollen. For these plants have been found to yield seed to the pollen of a distinct species, though quite sterile with their own pollen, notwithstanding that their own pollen was found to be perfectly good, for it fertilised distinct species. So that certain individual plants and all the individuals of certain species can actually be hybridised much more readily than they can be self-fertilised! For instance, a bulb of Hippeastrum aulicum produced four flowers; three were fertilised by Herbert with their own pollen, and the fourth was subsequently fertilised by the pollen of a compound hybrid descended from three other and distinct species: the result was that 'the ovaries of the three first flowers soon ceased to grow, and after a few days perished entirely, whereas the pod impregnated by the pollen of the hybrid made vigorous growth and rapid progress to maturity, and bore good seed, which vegetated freely.' In a letter to me, in 1839, Mr. Herbert told me that he had then tried the experiment during five years, and he continued to try it during several subsequent years, and always with the same result. This result has, also, been confirmed by other observers in the case of Hippeastrum with its sub-genera, and in the case of some other genera, as Lobelia, Passiflora and Verbascum. Although the plants in these experiments appeared perfectly healthy, and although both the ovules and pollen of the same flower were perfectly good with respect to other species, yet as they were functionally imperfect in their mutual self-action, we must infer that the plants were in an unnatural state. Nevertheless these facts show on what slight and mysterious causes the lesser or greater fertility of species when crossed, in comparison with the same species when self-fertilised, sometimes depends.

The practical experiments of horticulturists, though not made with scientific precision, deserve some notice. It is notorious in how complicated a manner the species of Pelargonium, Fuchsia, Calceolaria, Petunia, Rhododendron, &c., have been crossed, yet many of these hybrids seed freely. For instance, Herbert asserts that a hybrid from Calceolaria integrifolia and plantaginea, species most widely dissimilar in general habit, 'reproduced itself as perfectly as if it had been a natural species from the mountains of Chile.' I have taken some pains to ascertain the degree of fertility of some of the complex crosses of Rhododendrons, and I am assured that many of them are perfectly fertile. Mr. C. Noble, for instance, informs me that he raises stocks for grafting from a hybrid between Rhod. Ponticum and Catawbiense, and that this hybrid 'seeds as freely as it is possible to imagine.' Had hybrids, when fairly treated, gone on decreasing in fertility in each successive generation, as Gartner believes to be the case, the fact would have been notorious to nurserymen. Horticulturists raise large beds of the same hybrids, and such alone are fairly treated, for by insect agency the several individuals of the same hybrid variety are allowed to freely cross with each other, and the injurious influence of close interbreeding is thus prevented. Any one may readily convince himself of the efficiency of insect-agency by examining the flowers of the more sterile kinds of hybrid rhododendrons, which produce no pollen, for he will find on their stigmas plenty of pollen brought from other flowers.

In regard to animals, much fewer experiments have been carefully tried than with plants. If our systematic arrangements can be trusted, that is if the genera of animals are as distinct from each other, as are the genera of plants, then we may infer that animals more widely separated in the scale of nature can be more easily crossed than in the case of plants; but the hybrids themselves are, I think, more sterile. I doubt whether any case of a perfectly fertile hybrid animal can be considered as thoroughly well authenticated. It should, however, be borne in mind that, owing to few animals breeding freely under confinement, few experiments have been fairly tried: for instance, the canary-bird has been crossed with nine other finches, but as not one of these nine species breeds freely in confinement, we have no right to expect that the first crosses between them and the canary, or that their hybrids, should be perfectly fertile. Again, with respect to the fertility in successive generations of the more fertile hybrid animals, I hardly know of an instance in which two families of the same hybrid have been raised at the same time from different parents, so as to avoid the ill effects of close interbreeding. On the contrary, brothers and sisters have usually been crossed in each successive generation, in opposition to the constantly repeated admonition of every breeder. And in this case, it is not at all surprising that the inherent sterility in the hybrids should have gone on increasing. If we were to act thus, and pair brothers and sisters in the case of any pure animal, which from any cause had the least tendency to sterility, the breed would assuredly be lost in a very few generations.

Although I do not know of any thoroughly well-authenticated cases of perfectly fertile hybrid animals, I have some reason to believe that the hybrids from Cervulus vaginalis and Reevesii, and from Phasianus colchicus with P. torquatus and with P. versicolor are perfectly fertile. The hybrids from the common and Chinese geese (A. cygnoides), species which are so different that they are generally ranked in distinct genera, have often bred in this country with either pure parent, and in one single instance they have bred inter se. This was effected by Mr. Eyton, who raised two hybrids from the same parents but from different hatches; and from these two birds he raised no less than eight hybrids (grandchildren of the pure geese) from one nest. In India, however, these cross-bred geese must be far more fertile; for I am assured by two eminently capable judges, namely Mr. Blyth and Capt. Hutton, that whole flocks of these crossed geese are kept in various parts of the country; and as they are kept for profit, where neither pure parent-species exists, they must certainly be highly fertile.

A doctrine which originated with Pallas, has been largely accepted by modern naturalists; namely, that most of our domestic animals have descended from two or more aboriginal species, since commingled by intercrossing. On this view, the aboriginal species must either at first have produced quite fertile hybrids, or the hybrids must have become in subsequent generations quite fertile under domestication. This latter alternative seems to me the most probable, and I am inclined to believe in its truth, although it rests on no direct evidence. I believe, for instance, that our dogs have descended from several wild stocks; yet, with perhaps the exception of certain indigenous domestic dogs of South America, all are quite fertile together; and analogy makes me greatly doubt, whether the several aboriginal species would at first have freely bred together and have produced quite fertile hybrids. So again there is reason to believe that our European and the humped Indian cattle are quite fertile together; but from facts communicated to me by Mr. Blyth, I think they must be considered as distinct species. On this view of the origin of many of our domestic animals, we must either give up the belief of the almost universal sterility of distinct species of animals when crossed; or we must look at sterility, not as an indelible characteristic, but as one capable of being removed by domestication.

Finally, looking to all the ascertained facts on the intercrossing of plants and animals, it may be concluded that some degree of sterility, both in first crosses and in hybrids, is an extremely general result; but that it cannot, under our present state of knowledge, be considered as absolutely universal.

Laws governing the Sterility of first Crosses and of Hybrids. -- We will now consider a little more in detail the circumstances and rules governing the sterility of first crosses and of hybrids. Our chief object will be to see whether or not the rules indicate that species have specially been endowed with this quality, in order to prevent their crossing and blending together in utter confusion. The following rules and conclusions are chiefly drawn up from Gartner's admirable work on the hybridisation of plants. I have taken much pains to ascertain how far the rules apply to animals, and considering how scanty our knowledge is in regard to hybrid animals, I have been surprised to find how generally the same rules apply to both kingdoms.

It has been already remarked, that the degree of fertility, both of first crosses and of hybrids, graduates from zero to perfect fertility. It is surprising in how many curious ways this gradation can be shown to exist; but only the barest outline of the facts can here be given. When pollen from a plant of one family is placed on the stigma of a plant of a distinct family, it exerts no more influence than so much inorganic dust. From this absolute zero of fertility, the pollen of different species of the same genus applied to the stigma of some one species, yields a perfect gradation in the number of seeds produced, up to nearly complete or even quite complete fertility; and, as we have seen, in certain abnormal cases, even to an excess of fertility, beyond that which the plant's own pollen will produce. So in hybrids themselves, there are some which never have produced, and probably never would produce, even with the pollen of either pure parent, a single fertile seed: but in some of these cases a first trace of fertility may be detected, by the pollen of one of the pure parent-species causing the flower of the hybrid to wither earlier than it otherwise would have done; and the early withering of the flower is well known to be a sign of incipient fertilisation. From this extreme degree of sterility we have self-fertilised hybrids producing a greater and greater number of seeds up to perfect fertility.

Hybrids from two species which are very difficult to cross, and which rarely produce any offspring, are generally very sterile; but the parallelism between the difficulty of making a first cross, and the sterility of the hybrids thus produced--two classes of facts which are generally confounded together--is by no means strict. There are many cases, in which two pure species can be united with unusual facility, and produce numerous hybrid-offspring, yet these hybrids are remarkably sterile. On the other hand, there are species which can be crossed very rarely, or with extreme difficulty, but the hybrids, when at last produced, are very fertile. Even within the limits of the same genus, for instance in Dianthus, these two opposite cases occur.

The fertility, both of first crosses and of hybrids, is more easily affected by unfavourable conditions, than is the fertility of pure species. But the degree of fertility is likewise innately variable; for it is not always the same when the same two species are crossed under the same circumstances, but depends in part upon the constitution of the individuals which happen to have been chosen for the experiment. So it is with hybrids, for their degree of fertility is often found to differ greatly in the several individuals raised from seed out of the same capsule and exposed to exactly the same conditions.

By the term systematic affinity is meant, the resemblance between species in structure and in constitution, more especially in the structure of parts which are of high physiological importance and which differ little in the allied species. Now the fertility of first crosses between species, and of the hybrids produced from them, is largely governed by their systematic affinity. This is clearly shown by hybrids never having been raised between species ranked by systematists in distinct families; and on the other hand, by very closely allied species generally uniting with facility. But the correspondence between systematic affinity and the facility of crossing is by no means strict. A multitude of cases could be given of very closely allied species which will not unite, or only with extreme difficulty; and on the other hand of very distinct species which unite with the utmost facility. In the same family there may be a genus, as Dianthus, in which very many species can most readily be crossed; and another genus, as Silene, in which the most persevering efforts have failed to produce between extremely close species a single hybrid. Even within the limits of the same genus, we meet with this same difference; for instance, the many species of Nicotiana have been more largely crossed than the species of almost any other genus; but Gartner found that N. acuminata, which is not a particularly distinct species, obstinately failed to fertilise, or to be fertilised by, no less than eight other species of Nicotiana. Very many analogous facts could be given.

No one has been able to point out what kind, or what amount, of difference in any recognisable character is sufficient to prevent two species crossing. It can be shown that plants most widely different in habit and general appearance, and having strongly marked differences in every part of the flower, even in the pollen, in the fruit, and in the cotyledons, can be crossed. Annual and perennial plants, deciduous and evergreen trees, plants inhabiting different stations and fitted for extremely different climates, can often be crossed with ease.

By a reciprocal cross between two species, I mean the case, for instance, of a stallion-horse being first crossed with a female-ass, and then a male-ass with a mare: these two species may then be said to have been reciprocally crossed. There is often the widest possible difference in the facility of making reciprocal crosses. Such cases are highly important, for they prove that the capacity in any two species to cross is often completely independent of their systematic affinity, or of any recognisable difference in their whole organisation. On the other hand, these cases clearly show that the capacity for crossing is connected with constitutional differences imperceptible by us, and confined to the reproductive system. This difference in the result of reciprocal crosses between the same two species was long ago observed by Kolreuter. To give an instance: Mirabilis jalappa can easily be fertilised by the pollen of M. longiflora, and the hybrids thus produced are sufficiently fertile; but Kolreuter tried more than two hundred times, during eight following years, to fertilise reciprocally M. longiflora with the pollen of M. jalappa, and utterly failed. Several other equally striking cases could be given. Thuret has observed the same fact with certain sea-weeds or Fuci. Gartner, moreover, found that this difference of facility in making reciprocal crosses is extremely common in a lesser degree. He has observed it even between forms so closely related (as Matthiola annua and glabra) that many botanists rank them only as varieties. It is also a remarkable fact, that hybrids raised from reciprocal crosses, though of course compounded of the very same two species, the one species having first been used as the father and then as the mother, generally differ in fertility in a small, and occasionally in a high degree.

Several other singular rules could be given from Gartner: for instance, some species have a remarkable power of crossing with other species; other species of the same genus have a remarkable power of impressing their likeness on their hybrid offspring; but these two powers do not at all necessarily go together. There are certain hybrids which instead of having, as is usual, an intermediate character between their two parents, always closely resemble one of them; and such hybrids, though externally so like one of their pure parent-species, are with rare exceptions extremely sterile. So again amongst hybrids which are usually intermediate in structure between their parents, exceptional and abnormal individuals sometimes are born, which closely resemble one of their pure parents; and these hybrids are almost always utterly sterile, even when the other hybrids raised from seed from the same capsule have a considerable degree of fertility. These facts show how completely fertility in the hybrid is independent of its external resemblance to either pure parent.

Considering the several rules now given, which govern the fertility of first crosses and of hybrids, we see that when forms, which must be considered as good and distinct species, are united, their fertility graduates from zero to perfect fertility, or even to fertility under certain conditions in excess. That their fertility, besides being eminently susceptible to favourable and unfavourable conditions, is innately variable. That it is by no means always the same in degree in the first cross and in the hybrids produced from this cross. That the fertility of hybrids is not related to the degree in which they resemble in external appearance either parent. And lastly, that the facility of making a first cross between any two species is not always governed by their systematic affinity or degree of resemblance to each other. This latter statement is clearly proved by reciprocal crosses between the same two species, for according as the one species or the other is used as the father or the mother, there is generally some difference, and occasionally the widest possible difference, in the facility of effecting an union. The hybrids, moreover, produced from reciprocal crosses often differ in fertility.

Now do these complex and singular rules indicate that species have been endowed with sterility simply to prevent their becoming confounded in nature? I think not. For why should the sterility be so extremely different in degree, when various species are crossed, all of which we must suppose it would be equally important to keep from blending together? Why should the degree of sterility be innately variable in the individuals of the same species? Why should some species cross with facility, and yet produce very sterile hybrids; and other species cross with extreme difficulty, and yet produce fairly fertile hybrids? Why should there often be so great a difference in the result of a reciprocal cross between the same two species? Why, it may even be asked, has the production of hybrids been permitted? to grant to species the special power of producing hybrids, and then to stop their further propagation by different degrees of sterility, not strictly related to the facility of the first union between their parents, seems to be a strange arrangement.

The foregoing rules and facts, on the other hand, appear to me clearly to indicate that the sterility both of first crosses and of hybrids is simply incidental or dependent on unknown differences, chiefly in the reproductive systems, of the species which are crossed. The differences being of so peculiar and limited a nature, that, in reciprocal crosses between two species the male sexual element of the one will often freely act on the female sexual element of the other, but not in a reversed direction. It will be advisable to explain a little more fully by an example what I mean by sterility being incidental on other differences, and not a specially endowed quality. As the capacity of one plant to be grafted or budded on another is so entirely unimportant for its welfare in a state of nature, I presume that no one will suppose that this capacity is a specially endowed quality, but will admit that it is incidental on differences in the laws of growth of the two plants. We can sometimes see the reason why one tree will not take on another, from differences in their rate of growth, in the hardness of their wood, in the period of the flow or nature of their sap, &c.; but in a multitude of cases we can assign no reason whatever. Great diversity in the size of two plants, one being woody and the other herbaceous, one being evergreen and the other deciduous, and adaptation to widely different climates, does not always prevent the two grafting together. As in hybridisation, so with grafting, the capacity is limited by systematic affinity, for no one has been able to graft trees together belonging to quite distinct families; and, on the other hand, closely allied species, and varieties of the same species, can usually, but not invariably, be grafted with ease. But this capacity, as in hybridisation, is by no means absolutely governed by systematic affinity. Although many distinct genera within the same family have been grafted together, in other cases species of the same genus will not take on each other. The pear can be grafted far more readily on the quince, which is ranked as a distinct genus, than on the apple, which is a member of the same genus. Even different varieties of the pear take with different degrees of facility on the quince; so do different varieties of the apricot and peach on certain varieties of the plum.

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