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 Kölreuter. 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 Kölreuter 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. Gärtner, 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 Gärtner: 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.