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08 - Hybridism 08-02 - Sterility various in degree, not universal, affected by close interbreeding, removed by domestication 10 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, Koelreuter and Gaertner, 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 Gaertner 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 Gaertner, 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 Gaertner 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 Koelreuter and Gaertner 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 Gaertner.

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.

hothouse
hothouse


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.'

crinum
crinum


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.

lobelia
lobelia

hippeastrum
hippeastrum


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.

passiflora
passiflora

verbascum
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.

ovule
ovule

pollen
pollen


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.

pelargonium
pelargonium

fuchsia
fuchsia

calceolaria
calceolaria

petunia
petunia

rhododendron
rhododendron


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.'

integrifolia
integrifolia

plantaginea
plantaginea

Chile
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.

rhododendrons
rhododendrons


Ponticum and Catawbiense, and that this hybrid 'seeds as freely as it is possible to imagine.'

rhododendron ponticum
rhododendron ponticum

rhododendron catawbiense
rhododendron catawbiense


Had hybrids, when fairly treated, gone on decreasing in fertility in each successive generation, as Gaertner 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.

canary
canary

finch
finch


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.

reevesii
reevesii

Phasianus Colchicus
Phasianus Colchicus

Phasianus Torquatus
Phasianus Torquatus


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.

goose
goose

Chinese Goose
Chinese Goose


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.

India
India


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 its 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.

dog
dog

South America
South America


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 Gaertner'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.

pollen
pollen


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.

stigma
stigma


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.

dianthus
dianthus

silene
silene


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 Gaertner 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.

nicotiana
nicotiana


Very many analogous facts could be given.