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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.
08 - Hybridism 08-03 - Laws governing the sterility of hybrids 10 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 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.

horse
horse

ass
ass

mule
mule

hinny
hinny

Zebra Hinny
Zebra Hinny


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 Koelreuter.

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 Koelreuter 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.

Mirabilis Jalapa
Mirabilis Jalapa

Mirabilis Longiflora
Mirabilis Longiflora

Sea Weed
Sea Weed


Gaertner, 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.

matthiola
matthiola


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 Gaertner: 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.
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.

sap
sap


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.

graft
graft


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.

pear
pear

quince
quince

apple
apple


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.

apricot
apricot

peach
peach

plum
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.

gooseberry
gooseberry

Black Currant
Black Currant

Red Currant
Red Currant


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.

robinia
robinia


On the other hand, certain species of Sorbus, when grafted on other species, yielded twice as much fruit as when on their own roots.

sorbus
sorbus


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.

hippeastrum
hippeastrum

lobelia
lobelia


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.

pistil
pistil


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.

mule
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.