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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.
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.

tuber
tuber


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.

hermaphrodite
hermaphrodite

snail
snail


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.