Hybrid (biology)

“ And now we see that this same conclusion is independently and strongly fortified by the consideration of the illegitimate unions of trimorphic and dimorphic heterostyled plants. In so complex and obscure a subject as hybridism it is no slight gain to arrive at a definite conclusion, namely, that we must look exclusively to functional differences in the sexual elements, as the cause of the sterility of species when first crossed and of their hybrid offspring. ”

“ It is remarkable how closely and in how many points illegitimate unions between the two or three forms of the same heterostyled species, together with their illegitimate offspring, resemble hybrid unions between distinct species together with their hybrid offspring. In both cases we meet with every degree of sterility, from very slightly lessened fertility to absolute barrenness, when not even a single seed-capsule is produced. ”

“ Perhaps on close inspection you may find in some cases a slight difference, some character being not as fully developed in the hybrid as in the species. But when such is not the case, or where the opportunity for such a closer examination is wanting, a hybrid may easily be taken for a specimen of the pure race. Now take the seeds of your plant and sow them. If you had not supposed it to be hybrid you will be astonished at finding among its progeny some of a wholly different type. You will be led to conclude that you are observing a sudden change in structure such as is usually called a sport. ”

“ Darwin himself drew attention to this.
Let us next call to mind some of the peculiarities connected with hybridisation. We know that hybrids are often characterized by their large size, rapidity of growth, earlier production of flowers, wealth of flower-production and a longer life; hybrids, if crossed with one of the two parent forms, are usually more fertile than when they are crossed together or with another hybrid. But the characters which hybrids exhibit on self-fertilisation are rather variable. ”

“ 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. ”

“ What interests us here is the fact that hybrids of varieties do not remain constant in the second generation but usually split as it is said, remaining hybrid only in part of their offspring, the other portion returning to the parental types. ”

“ Hybridity.—The subject of hybridity has been fully discussed by many, and by no one more ably than by Darwin himself. It has generally been assumed that the hybrid of any two species is sterile, and, in fact, hybridity has been looked upon as one of the best tests of specific value next to genetic incapacity. The assumption finds its greatest support in genesis among the higher animals, and the most thoroughly differentiated species ”

“ Secondly, a large number, no doubt the greater number of novelties, are of hybrid origin. Here we may discern two cases. Hybrids may be produced by the crossing of old types, either of two old cultivated forms or newly introduced species, or ordinarily between an old and an introduced variety. Such novelties are excluded from our present discussion. Secondly, hybrids may be produced between a true, new mutation and some of the already existing varieties of the same species. ”

“ Until becoming acquainted with these facts, I was unwilling to believe in the frequent early death of hybrid embryos; for 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 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 ”

“ The recognition of symbasis, or the necessity of a broad foundation to sustain the organic structure, permits the inference that some hybrids are sterile and variable for the same reason that closely inbred plants and animals decline in fertility and produce mutations or deviations from the normal type. A hybrid is a mixture or cross between individuals which would not be expected to mix in nature. ”

“ Hybrids are very common among plants, and quite sufficiently so among animals to show that the boundaries of these so-called immutable species are not so well defined as has been supposed. Often, indeed, there is no offspring between the individuals of what are called distinct species, especially when they are widely different, and again, the offspring when produced is generally sterile; but when there is less difference between the parents, both the difficulty of breeding the hybrid, and its sterility when produced, are found to disappear. ”

“ But, in addition to their structural distinctions, the species of animals and plants, or at least a great number of them, exhibit physiological characters—what are known as distinct species, structurally, being for the most part either altogether incompetent to breed one with another; or if they breed, the resulting mule, or hybrid, is unable to perpetuate its race with another hybrid of the same kind.
A true physical cause is, however, admitted to be such only on one condition—that it shall account for all the phenomena which come within the range of its operation. ”

“ We here see that the formative elements included within the male element or pollen of one variety can affect and hybridise, not the part which they are properly adapted to affect, namely, the ovules, but the partially-developed tissues of a distinct variety or species. We are thus brought half-way towards a graft-hybrid, in which the formative elements included within the tissues of one individual combine with those included in the tissues of a distinct variety or species, thus giving rise to a new and intermediate form, independently of the male or female sexual organs. ”

“ The very important and interesting phenomenon of hybridism also belongs to this law of mixed or amphigonous211 transmission. It alone, when rightly estimated, is quite sufficient to refute the prevailing dogma of the constancy of species. Plants, as well as animals, belonging to quite different species, may sexually mingle with one another and produce descendants which in many cases can again propagate themselves, and that indeed either (more frequently) by mingling with one of the two parental species, or (more rarely) by pure in-breeding, hybrid mixing with hybrid. ”

“ But recalling our former consideration of this subject the supposition seems allowable that the tendency to revert is not connected with the type of the hybrid, but is apt to occur in some rare individuals of every type. But since most of the sterile hybrids are only known to us in a single individual and its vegetative offspring, this surmise offers an explanation of the rare occurrence of sports. ”

“ The sterility of hybrids is entirely due to the reproductive organs being affected; in the case of plants they continue to propagate freely by buds. The hybrid cells throw off hybrid gemmules which collect in the buds but cannot do so in the reproductive organs. ”

“ In such cases it is evident that the heritages of the parents remain uninfluenced by each other during the whole life of the hybrid, working side by side, but the active element always prevails over its latent opponent which is ready to break free whenever an opportunity is offered. ”

“ Sterile Hybrids.—The original notion of a hybrid, or at least the most popular meaning of the term, is that of a cross between organic types so widely diverse that the progeny are in some way abnormal or defective, especially with reference to reproduction. Among animals sterile hybrids can not be propagated, but in plants they can be grown from cuttings or buds, and are thus preserved as horticultural * varieties. ”

“ 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. ”

“ I well know the large amount of natural as well as artificial hybridisation that goes on. But, on the other hand, there are so many species which either will not cross at all, or produce sterile hybrids, that, taking a general view of all species together, mutual sterility does become by far the most generally distributed single peculiarity—i.e. is the one peculiarity which, more than any other that can be named, is common to numberless species. ”

“ In the case of hybrids, it apparently depends on their whole organisation having been disturbed by being compounded from two distinct forms; the sterility being closely allied to that which so frequently affects pure species, when exposed to new and unnatural conditions of life. He who will explain these latter cases will be able to explain the sterility of hybrids. This view is strongly supported by a parallelism of another kind: namely, that, firstly, slight changes in the conditions of life add to the vigour and fertility of all organic beings ”

“ The occurrence of hybrids in nature explains the difficulty experienced by botanists in deciding on what is a species, and the widely different limitations of the term adopted by different observers in the case of willows, roses, brambles, &c. The artificial process is practically the same in hybridization as in cross-fertilization, but usually requires more care. ”

“ He is as emphatic in his conclusion that some hybrids are perfectly fertile — as fertile as the pure parent-species — as are Kölreuter and Gärtner that some degree of sterility between distinct species is a universal law of nature. ”

“ It is by no means enough to point to natural and fertile hybrids as cases opposed to physiological selection unless it has been shown by experiment through a generation or two that these hybrids are fully fertile—i.e. as fertile as their parent species. Now, experiments of this kind have rarely been carried through. ”

“ For, you see, if you have not done that you have not strictly fulfilled all the conditions of the problem; you have not shown that you can produce, by the cause assumed, all the phenomena which you have in nature. Here are the phenomena of Hybridism staring you in the face, and you cannot say, 'I can, by selective modification, produce these same results.' Now, it is admitted on all hands that, at present, so far as experiments have gone, it has not been found possible to produce this complete physiological divergence by selective breeding. ”

“ But it is not as yet proved that a race ever exhibits, when crossed with another race of the same species, those phenomena of hybridization which are exhibited by many species when crossed with other species. On the other hand, not only is it not proved that all species give rise to hybrids infertile 'inter se', but there is much reason to believe that, in crossing, species exhibit every gradation from perfect sterility to perfect fertility. ”

“ Fifthly, the degree of sterility of first crosses and of hybrids runs, to a certain extent, parallel with the general or systematic affinity of the forms which are united. For species belonging to distinct genera can rarely, and those belonging to distinct families can never, be crossed. The parallelism, however, is far from complete; for a multitude of closely allied species will not unite, or unite with extreme difficulty, whilst other species, widely different from one another, can be crossed with perfect facility. ”

“ 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 relations 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, does not diminish ”

“ 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. ”