Origin of Species by Charles Darwin
Chapter IX. Hybridism
Laws governing the sterility of first crosses and of hybrids
We will now consider a little more in detail the laws governing the sterility of first crosses and of hybrids. Our chief object will be to see whether or not these laws indicate that species have been specially endowed with this quality, in order to prevent their crossing and blending together in utter confusion. The following conclusions are drawn up chiefly from Gartner's admirable work on the hybridisation of plants. I have taken much pains to ascertain how far they 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; 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 applied to the stigma of some one species of the same genus, 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 produces. So in hybrids themselves, there are some which never have produced, and probably never would produce, even with the pollen of the pure parents, 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.
The hybrids raised 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, as in the genus Verbascum, 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 that of pure species. But the fertility of first crosses is likewise innately variable; for it is not always the same in degree when the same two species are crossed under the same circumstances; it 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 the same conditions.
By the term systematic affinity is meant, the general resemblance between species in structure and constitution. Now the fertility of first crosses, 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. 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 female-ass being first crossed by a stallion, and then a mare by a male-ass: 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, that is of any difference in their structure or constitution, excepting in their reproductive systems. The diversity of the result in reciprocal crosses between the same two species was long ago observed by Kolreuter. To give an instance: Mirabilis jalapa 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. jalapa, 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 closely related forms (as Matthiola annua and glabra) which many botanists rank 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, though they rarely differ in external characters, yet 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 among 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 the fertility of a hybrid may be 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 the difference in the result of 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 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 in their reproductive systems; the differences being of so peculiar and limited a nature, that, in reciprocal crosses between the same 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 unimportant for their 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, etc.; 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 together trees 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.
As Gartner found that there was sometimes an innate difference in different individuals of the same two species in crossing; so Sagaret believes this to be the case with different individuals of the same two species in being grafted together. As in reciprocal crosses, the facility of effecting an union is often very far from equal, so it sometimes is in grafting. The common gooseberry, for instance, cannot be grafted on the currant, whereas the currant will take, though with difficulty, on the gooseberry.
We have seen that the sterility of hybrids which have their reproductive organs in an imperfect condition, is a different case from the difficulty of uniting two pure species, which have their reproductive organs perfect; yet these two distinct classes of cases run to a large extent parallel. Something analogous occurs in grafting; for Thouin found that three species of Robinia, which seeded freely on their own roots, and which could be grafted with no great difficulty on a fourth species, when thus grafted were rendered barren. On the other hand, certain species of Sorbus, when grafted on other species, yielded twice as much fruit as when on their own roots. We are reminded by this latter fact of the extraordinary cases of Hippeastrum, Passiflora, etc., which seed much more freely when fertilised with the pollen of a distinct species than when fertilised with pollen from the same plant.
We thus see that, although there is a clear and great difference between the mere adhesion of grafted stocks and the union of the male and female elements in the act of reproduction, yet that there is a rude degree of parallelism in the results of grafting and of crossing distinct species. And as we must look at the curious and complex laws governing the facility with which trees can be grafted on each other as incidental on unknown differences in their vegetative systems, so I believe that the still more complex laws governing the facility of first crosses are incidental on unknown differences in their reproductive systems. These differences in both cases follow, to a certain extent, as might have been expected, systematic affinity, by which term every kind of resemblance and dissimilarity between organic beings is attempted to be expressed. The facts by no means seem to indicate that the greater or lesser difficulty of either grafting or crossing various species has been a special endowment; although in the case of crossing, the difficulty is as important for the endurance and stability of specific forms as in the case of grafting it is unimportant for their welfare.