M Database Inspector (cheetah)
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|07 - Instinct||07-10 - Neuter or sterile insects||10||
The subject well deserves to be discussed at great length, but I will here take only a single case, that of working or sterile ants.
How the workers have been rendered sterile is a difficulty; but not much greater than that of any other striking modification of structure; for it can be shown that some insects and other articulate animals in a state of nature occasionally become sterile; and if such insects had been social, and it had been profitable to the community that a number should have been annually born capable of work, but incapable of procreation, I can see no very great difficulty in this being effected by natural selection.
But I must pass over this preliminary difficulty.
The great difficulty lies in the working ants differing widely from both the males and the fertile females in structure, as in the shape of the thorax and in being destitute of wings and sometimes of eyes, and in instinct.
As far as instinct alone is concerned, the prodigious difference in this respect between the workers and the perfect females, would have been far better exemplified by the hive-bee.
If a working ant or other neuter insect had been an animal in the ordinary state, I should have unhesitatingly assumed that all its characters had been slowly acquired through natural selection; namely, by an individual having been born with some slight profitable modification of structure, this being inherited by its offspring, which again varied and were again selected, and so onwards.
But with the working ant we have an insect differing greatly from its parents, yet absolutely sterile; so that it could never have transmitted successively acquired modifications of structure or instinct to its progeny.
It may well be asked how is it possible to reconcile this case with the theory of natural selection?
First, let it be remembered that we have innumerable instances, both in our domestic productions and in those in a state of nature, of all sorts of differences of structure which have become correlated to certain ages, and to either sex.
We have differences correlated not only to one sex, but to that short period alone when the reproductive system is active, as in the nuptial plumage of many birds, and in the hooked jaws of the male salmon.
We have even slight differences in the horns of different breeds of cattle in relation to an artificially imperfect state of the male sex; for oxen of certain breeds have longer horns than in other breeds, in comparison with the horns of the bulls or cows of these same breeds.
Hence I can see no real difficulty in any character having become correlated with the sterile condition of certain members of insect-communities: the difficulty lies in understanding how such correlated modifications of structure could have been slowly accumulated by natural selection.
This difficulty, though appearing insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be applied to the family, as well as to the individual, and may thus gain the desired end.
Thus, a well-flavoured vegetable is cooked, and the individual is destroyed; but the horticulturist sows seeds of the same stock, and confidently expects to get nearly the same variety; breeders of cattle wish the flesh and fat to be well marbled together; the animal has been slaughtered, but the breeder goes with confidence to the same family.
I have such faith in the powers of selection, that I do not doubt that a breed of cattle, always yielding oxen with extraordinarily long horns, could be slowly formed by carefully watching which individual bulls and cows, when matched, produced oxen with the longest horns; and yet no one ox could ever have propagated its kind.
Thus I believe it has been with social insects: a slight modification of structure, or instinct, correlated with the sterile condition of certain members of the community, has been advantageous to the community: consequently the fertile males and females of the same community flourished, and transmitted to their fertile offspring a tendency to produce sterile members having the same modification.
And I believe that this process has been repeated, until that prodigious amount of difference between the fertile and sterile females of the same species has been produced, which we see in many social insects.
But we have not as yet touched on the climax of the difficulty; namely, the fact that the neuters of several ants differ, not only from the fertile females and males, but from each other, sometimes to an almost incredible degree, and are thus divided into two or even three castes.
The castes, moreover, do not generally graduate into each other, but are perfectly well defined; being as distinct from each other, as are any two species of the same genus, or rather as any two genera of the same family.
Thus in Eciton, there are working and soldier neuters, with jaws and instincts extraordinarily different: in Cryptocerus, the workers of one caste alone carry a wonderful sort of shield on their heads, the use of which is quite unknown: in the Mexican Myrmecocystus, the workers of one caste never leave the nest; they are fed by the workers of another caste, and they have an enormously developed abdomen which secretes a sort of honey, supplying the place of that excreted by the aphides, or the domestic cattle as they may be called, which our European ants guard or imprison.
It will indeed be thought that I have an overweening confidence in the principle of natural selection, when I do not admit that such wonderful and well-established facts at once annihilate my theory.
In the simpler case of neuter insects all of one caste or of the same kind, which have been rendered by natural selection, as I believe to be quite possible, different from the fertile males and females, in this case, we may safely conclude from the analogy of ordinary variations, that each successive, slight, profitable modification did not probably at first appear in all the individual neuters in the same nest, but in a few alone; and that by the long-continued selection of the fertile parents which produced most neuters with the profitable modification, all the neuters ultimately came to have the desired character.
On this view we ought occasionally to find neuter-insects of the same species, in the same nest, presenting gradations of structure; and this we do find, even often, considering how few neuter-insects out of Europe have been carefully examined.
Mr F. Smith has shown how surprisingly the neuters of several British ants differ from each other in size and sometimes in colour; and that the extreme forms can sometimes be perfectly linked together by individuals taken out of the same nest: I have myself compared perfect gradations of this kind.
It often happens that the larger or the smaller sized workers are the most numerous; or that both large and small are numerous, with those of an intermediate size scanty in numbers.
Formica flava has larger and smaller workers, with some of intermediate size; and, in this species, as Mr F. Smith has observed, the larger workers have simple eyes (ocelli), which though small can be plainly distinguished, whereas the smaller workers have their ocelli rudimentary.
Having carefully dissected several specimens of these workers, I can affirm that the eyes are far more rudimentary in the smaller workers than can be accounted for merely by their proportionally lesser size; and I fully believe, though I dare not assert so positively, that the workers of intermediate size have their ocelli in an exactly intermediate condition.
So that we here have two bodies of sterile workers in the same nest, differing not only in size, but in their organs of vision, yet connected by some few members in an intermediate condition.
I may digress by adding, that if the smaller workers had been the most useful to the community, and those males and females had been continually selected, which produced more and more of the smaller workers, until all the workers had come to be in this condition; we should then have had a species of ant with neuters very nearly in the same condition with those of Myrmica.
For the workers of Myrmica have not even rudiments of ocelli, though the male and female ants of this genus have well-developed ocelli.
I may give one other case: so confidently did I expect to find gradations in important points of structure between the different castes of neuters in the same species, that I gladly availed myself of Mr F. Smith's offer of numerous specimens from the same nest of the driver ant (Anomma) of West Africa.
The reader will perhaps best appreciate the amount of difference in these workers, by my giving not the actual measurements, but a strictly accurate illustration: the difference was the same as if we were to see a set of workmen building a house of whom many were five feet four inches high, and many sixteen feet high; but we must suppose that the larger workmen had heads four instead of three times as big as those of the smaller men, and jaws nearly five times as big.
The jaws, moreover, of the working ants of the several sizes differed wonderfully in shape, and in the form and number of the teeth.
But the important fact for us is, that though the workers can be grouped into castes of different sizes, yet they graduate insensibly into each other, as does the widely-different structure of their jaws.
I speak confidently on this latter point, as Mr Lubbock made drawings for me with the camera lucida of the jaws which I had dissected from the workers of the several sizes.
With these facts before me, I believe that natural selection, by acting on the fertile parents, could form a species which should regularly produce neuters, either all of large size with one form of jaw, or all of small size with jaws having a widely different structure; or lastly, and this is our climax of difficulty, one set of workers of one size and structure, and simultaneously another set of workers of a different size and structure; a graduated series having been first formed, as in the case of the driver ant, and then the extreme forms, from being the most useful to the community, having been produced in greater and greater numbers through the natural selection of the parents which generated them; until none with an intermediate structure were produced.
Thus, as I believe, the wonderful fact of two distinctly defined castes of sterile workers existing in the same nest, both widely different from each other and from their parents, has originated.
We can see how useful their production may have been to a social community of insects, on the same principle that the division of labour is useful to civilised man.
As ants work by inherited instincts and by inherited tools or weapons, and not by acquired knowledge and manufactured instruments, a perfect division of labour could be effected with them only by the workers being sterile; for had they been fertile, they would have intercrossed, and their instincts and structure would have become blended.
And nature has, as I believe, effected this admirable division of labour in the communities of ants, by the means of natural selection.
But I am bound to confess, that, with all my faith in this principle, I should never have anticipated that natural selection could have been efficient in so high a degree, had not the case of these neuter insects convinced me of the fact.
I have, therefore, discussed this case, at some little but wholly insufficient length, in order to show the power of natural selection, and likewise because this is by far the most serious special difficulty, which my theory has encountered.
The case, also, is very interesting, as it proves that with animals, as with plants, any amount of modification in structure can be effected by the accumulation of numerous, slight, and as we must call them accidental, variations, which are in any manner profitable, without exercise or habit having come into play.
For no amount of exercise, or habit, or volition, in the utterly sterile members of a community could possibly have affected the structure or instincts of the fertile members, which alone leave descendants.
I am surprised that no one has advanced this demonstrative case of neuter insects, against the well-known doctrine of Lamarck.
|08 - Hybridism||08-01 - Distinction between the sterility of first crosses and of hybrids||10||
THE view generally entertained by naturalists is that species, when intercrossed, have been specially endowed with the quality of sterility, in order to prevent the confusion of all organic forms.
This view certainly seems at first probable, for species within the same country could hardly have kept distinct had they been capable of crossing freely.
The importance of the fact that hybrids are very generally sterile, has, I think, been much underrated by some late writers.
On the theory of natural selection the case is especially important, inasmuch as the sterility of hybrids could not possibly be of any advantage to them, and therefore could not have been acquired by the continued preservation of successive profitable degrees of sterility.
I hope, however, to be able to show that sterility is not a specially acquired or endowed quality, but is incidental on other acquired differences.
In treating this subject, two classes of facts, to a large extent fundamentally different, have generally been confounded together; namely, the sterility of two species when first crossed, and the sterility of the hybrids produced from them.
Pure species have of course their organs of reproduction in a perfect condition, yet when intercrossed they produce either few or no offspring.
Hybrids, on the other hand, have their reproductive organs functionally impotent, as may be clearly seen in the state of the male element in both plants and animals; though the organs themselves are perfect in structure, as far as the microscope reveals.
In the first case the two sexual elements which go to form the embryo are perfect; in the second case they are either not at all developed, or are imperfectly developed.
This distinction is important, when the cause of the sterility, which is common to the two cases, has to be considered.
The distinction has probably been slurred over, owing to the sterility in both cases being looked on as a special endowment, beyond the province of our reasoning powers.
The fertility of varieties, that is of the forms known or believed to have descended from common parents, when intercrossed, and likewise the fertility of their mongrel offspring, is, on my theory, of equal importance with the sterility of species; for it seems to make a broad and clear distinction between varieties and species.
First, for the sterility of species when crossed and of their hybrid offspring.
It is impossible to study the several memoirs and works of those two conscientious and admirable observers, Koelreuter and Gaertner, who almost devoted their lives to this subject, without being deeply impressed with the high generality of some degree of sterility.
Koelreuter makes the rule universal; but then he cuts the knot, for in ten cases in which he found two forms, considered by most authors as distinct species, quite fertile together, he unhesitatingly ranks them as varieties.
Gaertner, also, makes the rule equally universal; and he disputes the entire fertility of Koelreuter's ten cases.
But in these and in many other cases, Gaertner is obliged carefully to count the seeds, in order to show that there is any degree of sterility.
He always compares the maximum number of seeds produced by two species when crossed and by their hybrid offspring, with the average number produced by both pure parent-species in a state of nature.
But a serious cause of error seems to me to be here introduced: a plant to be hybridised must be castrated, and, what is often more important, must be secluded in order to prevent pollen being brought to it by insects from other plants.
Nearly all the plants experimentised on by Gaertner were potted, and apparently were kept in a chamber in his house.
That these processes are often injurious to the fertility of a plant cannot be doubted; for Gaertner gives in his table about a score of cases of plants which he castrated, and artificially fertilised with their own pollen, and (excluding all cases such as the Leguminosae, in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired.
Moreover, as Gaertner during several years repeatedly crossed the primrose and cowslip, which we have such good reason to believe to be varieties, and only once or twice succeeded in getting fertile seed; as he found the common red and blue pimpernels (Anagallis arvensis and coerulea), which the best botanists rank as varieties, absolutely sterile together; and as he came to the same conclusion in several other analogous cases; it seems to me that we may well be permitted to doubt whether many other species are really so sterile, when intercrossed, as Gaertner believes.
|02 - Variations Under Nature||02-06 - Many of the Species included within the Larger Genera resemble Varieties in being very closely, but unequally, related to each other, and in having restricted ranges||10||
There are other relations between the species of large genera and their recorded varieties which deserve notice.
We have seen that there is no infallible criterion by which to distinguish species and well-marked varieties; and when intermediate links have not been found between doubtful forms, naturalists are compelled to come to a determination by the amount of difference between them, judging by analogy whether or not the amount suffices to raise one or both to the rank of species.
Hence the amount of difference is one very important criterion in settling whether two forms should be ranked as species or varieties.
Now Fries has remarked in regard to plants, and Westwood in regard to insects, that in large genera the amount of difference between the species is often exceedingly small.
I have endeavoured to test this numerically by averages, and, as far as my imperfect results go, they confirm the view.
I have also consulted some sagacious and experienced observers, and, after deliberation, they concur in this view.
In this respect, therefore, the species of the larger genera resemble varieties, more than do the species of the smaller genera.
Or the case may be put in another way, and it maybe said, that in the larger genera, in which a number of varieties or incipient species greater than the average are now manufacturing, many of the species already manufactured still to a certain extent resemble varieties, for they differ from each other by less than the usual amount of difference.
Moreover, the species of the larger genera are related to each other, in the same manner as the varieties of any one species are related to each other. No naturalist pretends that all the species of a genus are equally distinct from each other; they may generally be divided into sub-genera, or sections, or lesser groups.
As Fries has well remarked, little groups of species are generally clustered like satellites around other species.
|10 - On The Geological Succession of Organic Beings||10-08 - On the state of development of ancient forms||10||
There has been much discussion whether recent forms are more highly developed than ancient.
I will not here enter on this subject, for naturalists have not as yet defined to each other's satisfaction what is meant by high and low forms.
But in one particular sense the more recent forms must, on my theory, be higher than the more ancient; for each new species is formed by having had some advantage in the struggle for life over other and preceding forms.
If under a nearly similar climate, the eocene inhabitants of one quarter of the world were put into competition with the existing inhabitants of the same or some other quarter, the eocene fauna or flora would certainly be beaten and exterminated; as would a secondary fauna by an eocene, and a palaeozoic fauna by a secondary fauna.
I do not doubt that this process of improvement has affected in a marked and sensible manner the organisation of the more recent and victorious forms of life, in comparison with the ancient and beaten forms; but I can see no way of testing this sort of progress.
Crustaceans, for instance, not the highest in their own class, may have beaten the highest molluscs.
From the extraordinary manner in which European productions have recently spread over New Zealand, and have seized on places which must have been previously occupied, we may believe, if all the animals and plants of Great Britain were set free in New Zealand, that in the course of time a multitude of British forms would become thoroughly naturalized there, and would exterminate many of the natives.
On the other hand, from what we see now occurring in New Zealand, and from hardly a single inhabitant of the southern hemisphere having become wild in any part of Europe, we may doubt, if all the productions of New Zealand were set free in Great Britain, whether any considerable number would be enabled to seize on places now occupied by our native plants and animals.
Under this point of view, the productions of Great Britain, may be said to be higher than those of New Zealand.
Yet the most skilful naturalist from an examination of the species of the two countries could not have foreseen this result.
Agassiz insists that ancient animals resemble to a certain extent the embryos of recent animals of the same classes; or that the geological succession of extinct forms is in some degree parallel to the embryological development of recent forms.
I must follow Pictet and Huxley in thinking that the truth of this doctrine is very far from proved.
Yet I fully expect to see it hereafter confirmed, at least in regard to subordinate groups, which have branched off from each other within comparatively recent times.
For this doctrine of Agassiz accords well with the theory of natural selection.
In a future chapter I shall attempt to show that the adult differs from its embryo, owing to variations supervening at a not early age, and being inherited at a corresponding age.
This process, whilst it leaves the embryo almost unaltered, continually adds, in the course of successive generations, more and more difference to the adult.
Thus the embryo comes to be left as a sort of picture, preserved by nature, of the ancient and less modified condition of each animal.
This view may be true, and yet it may never be capable of full proof.
Seeing, for instance, that the oldest known mammals, reptiles, and fish strictly belong to their own proper classes, though some of these old forms are in a slight degree less distinct from each other than are the typical members of the same groups at the present day, it would be vain to look for animals having the common embryological character of the Vertebrata, until beds far beneath the lowest Silurian strata are discovered a discovery of which the chance is very small.