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|08 - Hybridism||08-04 - Sterility not a special endowment, but incidental on other differences||10||
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.
As Gaertner 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 very different case from the difficulty of uniting two pure species, which have their reproductive organs perfect; yet these two distinct cases run to a certain 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 another 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 case of Hippeastrum, Lobelia, &c., which seeded much more freely when fertilised with the pollen of distinct species, than when self-fertilised with their own pollen.
We thus see, that although there is a clear and fundamental 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, chiefly in their reproductive systems.
These differences, in both cases, follow to a certain extent, as might have been expected, systematic affinity, by which every kind of resemblance and dissimilarity between organic beings is attempted to be expressed.
The facts by no means seem to me to indicate that the greater or lesser difficulty of either grafting or crossing together 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.
|08 - Hybridism||08-05 - Causes of the sterility of first crosses and of hybrids||10||
Causes of the Sterility of first Crosses and of Hybrids.
We may now look a little closer at the probable causes of the sterility of first crosses and of hybrids.
These two cases are fundamentally different, for, as just remarked, in the union of two pure species the male and female sexual elements are perfect, whereas in hybrids they are imperfect.
Even in first crosses, the greater or lesser difficulty in effecting a union apparently depends on several distinct causes.
There must sometimes be a physical impossibility in the male element reaching the ovule, as would be the case with a plant having a pistil too long for the pollen-tubes to reach the ovarium.
It has also been observed that when pollen of one species is placed on the stigma of a distantly allied species, though the pollen-tubes protrude, they do not penetrate the stigmatic surface.
Again, the male element may reach the female element, but be incapable of causing an embryo to be developed, as seems to have been the case with some of Thuret's experiments on Fuci.
No explanation can be given of these facts, any more than why certain trees cannot be grafted on others.
Lastly, an embryo may be developed, and then perish at an early period.
This latter alternative has not been sufficiently attended to; but I believe, from observations communicated to me by Mr. Hewitt, who has had great experience in hybridising gallinaceous birds, that the early death of the embryo is a very frequent cause of sterility in first crosses.
I was at first very unwilling to believe in this view; as hybrids, when once born, are generally healthy and long-lived, as we see in the case of the common mule.
Hybrids, however, are differently circumstanced before and after birth: when born and living in a country where their two parents can live, they are generally placed under suitable conditions of life.
But a hybrid partakes of only half of the nature and constitution of its mother, and therefore before birth, as long as it is nourished within its mother's womb or within the egg or seed produced by the mother, it may be exposed to conditions in some degree unsuitable, and consequently be liable to perish at an early period; more especially as all very young beings seem eminently sensitive to injurious or unnatural conditions of life.
In regard to the sterility of hybrids, in which the sexual elements are imperfectly developed, the case is very different.
I have more than once alluded to a large body of facts, which I have collected, showing that when animals and plants are removed from their natural conditions, they are extremely liable to have their reproductive systems seriously affected.
This, in fact, is the great bar to the domestication of animals.
Between the sterility thus superinduced and that of hybrids, there are many points of similarity.
In both cases the sterility is independent of general health, and is often accompanied by excess of size or great luxuriance.
In both cases, the sterility occurs in various degrees; in both, the male element is the most liable to be affected; but sometimes the female more than the male.
In both, the tendency goes to a certain extent with systematic affinity, or whole groups of animals and plants are rendered impotent by the same unnatural conditions; and whole groups of species tend to produce sterile hybrids.
On the other hand, one species in a group will sometimes resist great changes of conditions with unimpaired fertility; and certain species in a group will produce unusually fertile hybrids.
No one can tell, till he tries, whether any particular animal will breed under confinement or any plant seed freely under culture; nor can he tell, till he tries, whether any two species of a genus will produce more or less sterile hybrids.
Lastly, when organic beings are placed during several generations under conditions not natural to them, they are extremely liable to vary, which is due, as I believe, to their reproductive systems having been specially affected, though in a lesser degree than when sterility ensues.
So it is with hybrids, for hybrids in successive generations are eminently liable to vary, as every experimentalist has observed.
Thus we see that when organic beings are placed under new and unnatural conditions, and when hybrids are produced by the unnatural crossing of two species, the reproductive system, independently of the general state of health, is affected by sterility in a very similar manner.
In the one case, the conditions of life have been disturbed, though often in so slight a degree as to be inappreciable by us; in the other case, or that of hybrids,the external conditions have remained the same, but the organisation has been disturbed by two different structures and constitutions having been blended into one.
For it is scarcely possible that two organisations should be compounded into one, without some disturbance occurring in the development, or periodical action, or mutual relation of the different parts and organs one to another, or to the conditions of life.
When hybrids are able to breed inter se, they transmit to their offspring from generation to generation the same compounded organisation, and hence we need not be surprised that their sterility, though in some degree variable, rarely diminishes.
It must, however, be confessed that we cannot understand, excepting on vague hypotheses, several facts with respect to the sterility of hybrids; for instance, the unequal fertility of hybrids produced from reciprocal crosses; or the increased sterility in those hybrids which occasionally and exceptionally resemble closely either pure parent.
Nor do I pretend that the foregoing remarks go to the root of the matter: no explanation is offered why an organism, when placed under unnatural conditions, is rendered sterile.
All that I have attempted to show, is that in two cases, in some respects allied, sterility is the common result, in the one case from the conditions of life having been disturbed, in the other case from the organisation having been disturbed by two organisations having been compounded into one.
|08 - Hybridism||08-06 - Parallelism between the effects of changed conditions of life and crossing||10||
It may seem fanciful, but I suspect that a similar parallelism extends to an allied yet very different class of facts. It is an old and almost universal belief, founded, I think, on a considerable body of evidence, that slight changes in the conditions of life are beneficial to all living things.
We see this acted on by farmers and gardeners in their frequent exchanges of seed, tubers, &c., from one soil or climate to another, and back again.
During the convalescence of animals, we plainly see that great benefit is derived from almost any change in the habits of life.
Again, both with plants and animals, there is abundant evidence, that a cross between very distinct individuals of the same species, that is between members of different strains or sub-breeds, gives vigour and fertility to the offspring.
I believe, indeed, from the facts alluded to in our fourth chapter, that a certain amount of crossing is indispensable even with hermaphrodites; and that close interbreeding continued during several generations between the nearest relations, especially if these be kept under the same conditions of life, always induces weakness and sterility in the progeny.
Hence it seems that, on the one hand, slight changes in the conditions of life benefit all organic beings, and on the other hand, that slight crosses, that is crosses between the males and females of the same species which have varied and become slightly different, give vigour and fertility to the offspring.
But we have seen that greater changes, or changes of a particular nature, often render organic beings in some degree sterile; and that greater crosses, that is crosses between males and females which have become widely or specifically different, produce hybrids which are generally sterile in some degree.
I cannot persuade myself that this parallelism is an accident or an illusion.
Both series of facts seem to be connected together by some common but unknown bond, which is essentially related to the principle of life.
|08 - Hybridism||08-07 - Fertility of varieties when crossed and of their mongrel offspring not universal||10||
Fertility of Varieties when crossed, and of their Mongrel off-spring.
It may be urged, as a most forcible argument, that there must be some essential distinction between species and varieties, and that there must be some error in all the foregoing remarks, inasmuch as varieties, however much they may differ from each other in external appearance, cross with perfect facility, and yield perfectly fertile offspring.
I fully admit that this is almost invariably the case.
But if we look to varieties produced under nature, we are immediately involved in hopeless difficulties; for if two hitherto reputed varieties be found in any degree sterile together, they are at once ranked by most naturalists as species.
For instance, the blue and red pimpernel, the primrose and cowslip, which are considered by many of our best botanists as varieties, are said by Gaertner not to be quite fertile when crossed, and he consequently ranks them as undoubted species.
If we thus argue in a circle, the fertility of all varieties produced under nature will assuredly have to be granted.
If we turn to varieties, produced, or supposed to have been produced, under domestication, we are still involved in doubt.
For when it is stated, for instance, that the German Spitz dog unites more easily than other dogs with foxes, or that certain South American indigenous domestic dogs do not readily cross with European dogs, the explanation which will occur to everyone, and probably the true one, is that these dogs have descended from several aboriginally distinct species.
Nevertheless the perfect fertility of so many domestic varieties, differing widely from each other in appearance, for instance of the pigeon or of the cabbage, is a remarkable fact; more especially when we reflect how many species there are, which, though resembling each other most closely, are utterly sterile when intercrossed.
Several considerations, however, render the fertility of domestic varieties less remarkable than at first appears. It can, in the first place, be clearly shown that mere external dissimilarity between two species does not determine their greater or lesser degree of sterility when crossed; and we may apply the same rule to domestic varieties.
In the second place, some eminent naturalists believe that a long course of domestication tends to eliminate sterility in the successive generations of hybrids, which were at first only slightly sterile; and if this be so, we surely ought not to expect to find sterility both appearing and disappearing under nearly the same conditions of life.
Lastly, and this seems to me by far the most important consideration, new races of animals and plants are produced under domestication by man's methodical and unconscious power of selection, for his own use and pleasure: he neither wishes to select, nor could select, slight differences in the reproductive system, or other constitutional difference correlated with the reproductive system.
He supplies his several varieties with the same food; treats them in nearly the same manner, and does not wish to alter their general habits of life.
Nature acts uniformly and slowly during vast periods of time on the whole organization, in any way which may be for each creature's own good; and thus she may, either directly, or more probably indirectly, through correlation, modify the reproductive system in the several descendants from any one species.
Seeing this difference in the process of selection, as carried on by man and nature, we need not be surprised at some difference in the result.
I have as yet spoken as if the varieties of the same species were invariably fertile when intercrossed.
But it seems to me impossible to resist the evidence of the existence of a certain amount of sterility in the few following cases, which I will briefly abstract.
The evidence is at least as good as that from which we believe in the sterility of a multitude of species.
The evidence is, also, derived from hostile witnesses, who in all other cases consider fertility and sterility as safe criterions of specific distinction.
Gaertner kept during several years a dwarf kind of maize with yellow seeds, and a tall variety with red seeds, growing near each other in his garden; and although these plants have separated sexes, they never naturally crossed.
He then fertilized thirteen flowers of the one with the pollen of the other; but only a single head produced any seed, and this one head produced only five grains.
Manipulation in this case could not have been injurious, as the plants have separated sexes.
No one, I believe, has suspected that these varieties of maize are distinct species; and it is important to notice that the hybrid plants thus raised were themselves perfectly fertile; so that even Gaertner did not venture to consider the two varieties as specifically distinct.
Girou de Buzareingues crossed three varieties of gourd, which like the maize has separated sexes, and he asserts that their mutual fertilization is by so much the less easy as their differences are greater.
How far these experiments may be trusted, I know not; but the forms experimentised on, are ranked by Sagaret, who mainly founds his classification by the test of infertility, as varieties.
The following case is far more remarkable, and seems at first quite incredible; but it is the result of an astonishing number of experiments made during many years on nine species of Verbascum, by so good an observer and so hostile a witness, as Gaertner: namely, that yellow and white varieties of the same species of Verbascum when intercrossed produce less seed, than do either coloured varieties when fertilized with pollen from their own coloured flowers.
Moreover, he asserts that when yellow and white varieties of one species are crossed with yellow and white varieties of a distinct species, more seed is produced by the crosses between the same coloured flowers, than between those which are differently coloured.
Yet these varieties of Verbascum present no other difference besides the mere colour of the flower; and one variety can sometimes be raised from the seed of the other.
From observations which I have made on certain varieties of hollyhock, I am inclined to suspect that they present analogous facts.
Koelreuter, whose accuracy has been confirmed by every subsequent observer, has proved the remarkable fact, that one variety of the common tobacco is more fertile, when crossed with a widely distinct species, than are the other varieties.
He experimentised on five forms, which are commonly reputed to be varieties, and which he tested by the severest trial, namely, by reciprocal crosses, and he found their mongrel offspring perfectly fertile.
But one of these five varieties, when used either as father or mother, and crossed with the Nicotiana glutinosa, always yielded hybrids not so sterile as those which were produced from the four other varieties when crossed with N. glutinosa.
Hence the reproductive system of this one variety must have been in some manner and in some degree modified.
From these facts; from the great difficulty of ascertaining the infertility of varieties in a state of nature, for a supposed variety if infertile in any degree would generally be ranked as species; from man selecting only external characters in the production of the most distinct domestic varieties, and from not wishing or being able to produce recondite and functional differences in the reproductive system; from these several considerations and facts, I do not think that the very general fertility of varieties can be proved to be of universal occurrence, or to form a fundamental distinction between varieties and species.
The general fertility of varieties does not seem to me sufficient to overthrow the view which I have taken with respect to the very general, but not invariable, sterility of first crosses and of hybrids, namely, that it is not a special endowment, but is incidental on slowly acquired modifications, more especially in the reproductive systems of the forms which are crossed.