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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.
06 - Difficutiles in Theory 06-03 - Absence or Rarity of Transitional Varieties 10 But, as by this theory innumerable transitional forms must have existed, why do we not find them embedded in countless numbers in the crust of the earth?

It will be more convenient to discuss this question in the chapter on the Imperfection of the Geological Record; and I will here only state that I believe the answer mainly lies in the record being incomparably less perfect than is generally supposed.


The crust of the earth is a vast museum; but the natural connections have been imperfectly made, and only at long intervals of time.

fossil
fossil
03 - Struggle for Existence 03-04 - Rapid Increase of naturalised Animals and Plants 10 But we have better evidence on this subject than mere theoretical calculations, namely, the numerous recorded cases of the astonishingly rapid increase of various animals in a state of nature, when circumstances have been favourable to them during two or three following seasons.

Still more striking is the evidence from our domestic animals of many kinds which have run wild in several parts of the world; if the statements of the rate of increase of slow-breeding cattle and horses in South America, and latterly in Australia, had not been well authenticated, they would have been incredible.

cattle
cattle

horses
horses


So it is with plants; cases could be given of introduced plants which have become common throughout whole islands in a period of less than ten years.

Several of the plants, such as the cardoon and a tall thistle, which are now the commonest over the whole plains of La Plata, clothing square leagues of surface almost to the exclusion of every other plant, have been introduced from Europe; and there are plants which now range in India, as I hear from Dr. Falconer, from Cape Comorin to the Himalaya, which have been imported from America since its discovery.

cardoon
cardoon

thistle
thistle

La Plata
La Plata

Europe
Europe

India
India

Cape Comorin
Cape Comorin

Himalaya
Himalaya

America
America


In such cases, and endless others could be given, no one supposes that the fertility of the animals or plants has been suddenly and temporarily increased in any sensible degree.

The obvious explanation is that the conditions of life have been highly favourable, and that there has consequently been less destruction of the old and young, and that nearly all the young have been enabled to breed.

Their geometrical ratio of increase, the result of which never fails to be surprising, simply explains their extraordinarily rapid increase and wide diffusion in their new homes.
09 - On the Imperfection of the Geological Record 09-05 - On the intermittence of geological formations 10 But the imperfection in the geological record mainly results from another and more important cause than any of the foregoing; namely, from the several formations being separated from each other by wide intervals of time.

When we see the formations tabulated in written works, or when we follow them in nature, it is difficult to avoid believing that they are closely consecutive.

But we know, for instance, from Sir R. Murchison's great work on Russia, what wide gaps there are in that country between the superimposed formations; so it is in North America, and in many other parts of the world.

North America
North America


The most skilful geologist, if his attention had been exclusively confined to these large territories, would never have suspected that during the periods which were blank and barren in his own country, great piles of sediment, charged with new and peculiar forms of life, had elsewhere been accumulated.

And if in each separate territory, hardly any idea can be formed of the length of time which has elapsed between the consecutive formations, we may infer that this could nowhere be ascertained.

The frequent and great changes in the mineralogical composition of consecutive formations, generally implying great changes in the geography of the surrounding lands, whence the sediment has been derived, accords with the belief of vast intervals of time having elapsed between each formation.

But we can, I think, see why the geological formations of each region are almost invariably intermittent; that is, have not followed each other in close sequence.

Scarcely any fact struck me more when examining many hundred miles of the South American coasts, which have been upraised several hundred feet within the recent period, than the absence of any recent deposits sufficiently extensive to last for even a short geological period.

South America
South America


Along the whole west coast, which is inhabited by a peculiar marine fauna, tertiary beds are so scantily developed, that no record of several successive and peculiar marine faunas will probably be preserved to a distant age.

A little reflection will explain why along the rising coast of the western side of South America, no extensive formations with recent or tertiary remains can anywhere be found, though the supply of sediment must for ages have been great, from the enormous degradation of the coast-rocks and from muddy streams entering the sea.

The explanation, no doubt, is, that the littoral and sub-littoral deposits are continually worn away, as soon as they are brought up by the slow and gradual rising of the land within the grinding action of the coast-waves.

We may, I think, safely conclude that sediment must be accumulated in extremely thick, solid, or extensive masses, in order to withstand the incessant action of the waves, when first upraised and during subsequent oscillations of level.

Such thick and extensive accumulations of sediment may be formed in two ways; either, in profound depths of the sea, in which case, judging from the researches of E. Forbes, we may conclude that the bottom will be inhabited by extremely few animals, and the mass when upraised will give a most imperfect record of the forms of life which then existed; or, sediment may be accumulated to any thickness and extent over a shallow bottom, if it continue slowly to subside.

In this latter case, as long as the rate of subsidence and supply of sediment nearly balance each other, the sea will remain shallow and favourable for life, and thus a fossiliferous formation thick enough, when upraised, to resist any amount of degradation, may be formed.

fossil
fossil


I am convinced that all our ancient formations, which are rich in fossils, have thus been formed during subsidence.

Since publishing my views on this subject in 1845, I have watched the progress of Geology, and have been surprised to note how author after author, in treating of this or that great formation, has come to the conclusion that it was accumulated during subsidence.

I may add, that the only ancient tertiary formation on the west coast of South America, which has been bulky enough to resist such degradation as it has as yet suffered, but which will hardly last to a distant geological age, was certainly deposited during a downward oscillation of level, and thus gained considerable thickness.

South America
South America


All geological facts tell us plainly that each area has undergone numerous slow oscillations of level, and apparently these oscillations have affected wide spaces.

Consequently formations rich in fossils and sufficiently thick and extensive to resist subsequent degradation, may have been formed over wide spaces during periods of subsidence, but only where the supply of sediment was sufficient to keep the sea shallow and to embed and preserve the remains before they had time to decay.

On the other hand, as long as the bed of the sea remained stationary, thick deposits could not have been accumulated in the shallow parts, which are the most favourable to life.

Still less could this have happened during the alternate periods of elevation; or, to speak more accurately, the beds which were then accumulated will have been destroyed by being upraised and brought within the limits of the coast-action.

Thus the geological record will almost necessarily be rendered intermittent.

I feel much confidence in the truth of these views, for they are in strict accordance with the general principles inculcated by Sir C. Lyell; and E. Forbes independently arrived at a similar conclusion.

Sir Charles Lyell
Sir Charles Lyell


One remark is here worth a passing notice.

During periods of elevation the area of the land and of the adjoining shoal parts of the sea will be increased, and new stations will often be formed; all circumstances most favourable, as previously explained, for the formation of new varieties and species; but during such periods there will generally be a blank in the geological record.

On the other hand, during subsidence, the inhabited area and number of inhabitants will decrease (excepting the productions on the shores of a continent when first broken up into an archipelago), and consequently during subsidence, though there will be much extinction, fewer new varieties or species will be formed; and it is during these very periods of subsidence, that our great deposits rich in fossils have been accumulated.

Nature may almost be said to have guarded against the frequent discovery of her transitional or linking forms.

From the foregoing considerations it cannot be doubted that the geological record, viewed as a whole, is extremely imperfect; but if we confine our attention to any one formation, it becomes more difficult to understand, why we do not therein find closely graduated varieties between the allied species which lived at its commencement and at its close.

Some cases are on record of the same species presenting distinct varieties in the upper and lower parts of the same formation, but, as they are rare, they may be here passed over.

Although each formation has indisputably required a vast number of years for its deposition, I can see several reasons why each should not include a graduated series of links between the species which then lived; but I can by no means pretend to assign due proportional weight to the following considerations.

Although each formation may mark a very long lapse of years, each perhaps is short compared with the period requisite to change one species into another.

I am aware that two palaeontologists, whose opinions are worthy of much deference, namely Bronn and Woodward, have concluded that the average duration of each formation is twice or thrice as long as the average duration of specific forms.

Heinrich Georg Bronn
Heinrich Georg Bronn


But insuperable difficulties, as it seems to me, prevent us coming to any just conclusion on this head.

When we see a species first appearing in the middle of any formation, it would be rash in the extreme to infer that it had not elsewhere previously existed.

So again when we find a species disappearing before the uppermost layers have been deposited, it would be equally rash to suppose that it then became wholly extinct.

We forget how small the area of Europe is compared with the rest of the world; nor have the several stages of the same formation throughout Europe been correlated with perfect accuracy.

europe
europe


With marine animals of all kinds, we may safely infer a large amount of migration during climatal and other changes; and when we see a species first appearing in any formation, the probability is that it only then first immigrated into that area.

It is well known, for instance, that several species appeared somewhat earlier in the palaeozoic beds of North America than in those of Europe; time having apparently been required for their migration from the American to the European seas.

Americas
Americas

europe
europe


In examining the latest deposits of various quarters of the world, it has everywhere been noted, that some few still existing species are common in the deposit, but have become extinct in the immediately surrounding sea; or, conversely, that some are now abundant in the neighbouring sea, but are rare or absent in this particular deposit.

It is an excellent lesson to reflect on the ascertained amount of migration of the inhabitants of Europe during the Glacial period, which forms only a part of one whole geological period; and likewise to reflect on the great changes of level, on the inordinately great change of climate, on the prodigious lapse of time, all included within this same glacial period.

Yet it may be doubted whether in any quarter of the world, sedimentary deposits, including fossil remains, have gone on accumulating within the same area during the whole of this period.

fossil
fossil


It is not, for instance, probable that sediment was deposited during the whole of the glacial period near the mouth of the Mississippi, within that limit of depth at which marine animals can flourish; for we know what vast geographical changes occurred in other parts of America during this space of time.

When such beds as were deposited in shallow water near the mouth of the Mississippi during some part of the glacial period shall have been upraised, organic remains will probably first appear and disappear at different levels, owing to the migration of species and to geographical changes.

And in the distant future, a geologist examining these beds, might be tempted to conclude that the average duration of life of the embedded fossils had been less than that of the glacial period, instead of having been really far greater, that is extending from before the glacial epoch to the present day.

In order to get a perfect gradation between two forms in the upper and lower parts of the same formation, the deposit must have gone on accumulating for a very long period, in order to have given sufficient time for the slow process of variation; hence the deposit will generally have to be a very thick one; and the species undergoing modification will have had to live on the same area throughout this whole time.

But we have seen that a thick fossiliferous formation can only be accumulated during a period of subsidence; and to keep the depth approximately the same, which is necessary in order to enable the same species to live on the same space, the supply of sediment must nearly have counterbalanced the amount of subsidence.

But this same movement of subsidence will often tend to sink the area whence the sediment is derived, and thus diminish the supply whilst the downward movement continues.

In fact, this nearly exact balancing between the supply of sediment and the amount of subsidence is probably a rare contingency; for it has been observed by more than one palaeontologist, that very thick deposits are usually barren of organic remains, except near their upper or lower limits.

It would seem that each separate formation, like the whole pile of formations in any country, has generally been intermittent in its accumulation.

When we see, as is so often the case, a formation composed of beds of different mineralogical composition, we may reasonably suspect that the process of deposition has been much interrupted, as a change in the currents of the sea and a supply of sediment of a different nature will generally have been due to geographical changes requiring much time.

Nor will the closest inspection of a formation give any idea of the time which its deposition has consumed.

Many instances could be given of beds only a few feet in thickness, representing formations, elsewhere thousands of feet in thickness, and which must have required an enormous period for their accumulation; yet no one ignorant of this fact would have suspected the vast lapse of time represented by the thinner formation.

Many cases could be given of the lower beds of a formation having been upraised, denuded, submerged, and then re-covered by the upper beds of the same formation, facts, showing what wide, yet easily overlooked, intervals have occurred in its accumulation.

In other cases we have the plainest evidence in great fossilised trees, still standing upright as they grew, of many long intervals of time and changes of level during the process of deposition, which would never even have been suspected, had not the trees chanced to have been preserved: thus, Messrs Lyell and Dawson found carboniferous beds 1400 feet thick in Nova Scotia, with ancient root-bearing strata, one above the other, at no less than sixty-eight different levels.

Sir Charles Lyell
Sir Charles Lyell


Hence, when the same species occur at the bottom, middle, and top of a formation, the probability is that they have not lived on the same spot during the whole period of deposition, but have disappeared and reappeared, perhaps many times, during the same geological period.

So that if such species were to undergo a considerable amount of modification during any one geological period, a section would not probably include all the fine intermediate gradations which must on my theory have existed between them, but abrupt, though perhaps very slight, changes of form.