On the Origin of Species By Means of Natural Selection

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.

SUMMARY.

I have endeavoured briefly in this chapter to show that the mental qualities of our domestic animals vary, and that the variations are inherited. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. No one will dispute that instincts are of the highest importance to each animal. Therefore I can see no difficulty, under changing conditions of life, in natural selection accumulating slight modifications of instinct to any extent, in any useful direction. In some cases habit or use and disuse have probably come into play. I do not pretend that the facts given in this chapter strengthen in any great degree my theory; but none of the cases of difficulty, to the best of my judgment, annihilate it. On the other hand, the fact that instincts are not always absolutely perfect and are liable to mistakes; – that no instinct has been produced for the exclusive good of other animals, but that each animal takes advantage of the instincts of others; – that the canon in natural history, of "natura non facit saltum" is applicable to instincts as well as to corporeal structure, and is plainly explicable on the foregoing views, but is otherwise inexplicable, – all tend to corroborate the theory of natural selection.

This theory is, also, strengthened by some few other facts in regard to instincts; as by that common case of closely allied, but certainly distinct, species, when inhabiting distant parts of the world and living under considerably different conditions of life, yet often retaining nearly the same instincts. For instance, we can understand on the principle of inheritance, how it is that the thrush of South America lines its nest with mud, in the same peculiar manner as does our British thrush: how it is that the male wrens (Troglodytes) of North America, build "cock-nests," to roost in, like the males of our distinct Kitty-wrens, – a habit wholly unlike that of any other known bird. Finally, it may not be a logical deduction, but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its foster-brothers, – ants making slaves, – the larvae of ichneumonidae feeding within the live bodies of caterpillars, – not as specially endowed or created instincts, but as small consequences of one general law, leading to the advancement of all organic beings, namely, multiply, vary, let the strongest live and the weakest die.

8. HYBRIDISM

Distinction between the sterility of first crosses and of hybrids. Sterility various in degree, not universal, affected by close interbreeding, removed by domestication. Laws governing the sterility of hybrids. Sterility not a special endowment, but incidental on other differences. Causes of the sterility of first crosses and of hybrids. Parallelism between the effects of changed conditions of life and crossing. Fertility of varieties when crossed and of their mongrel offspring not universal. Hybrids and mongrels compared independently of their fertility. Summary.

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, Kolreuter and Gartner, who almost devoted their lives to this subject, without being deeply impressed with the high generality of some degree of sterility. Kolreuter 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. Gartner, also, makes the rule equally universal; and he disputes the entire fertility of Kolreuter's ten cases. But in these and in many other cases, Gartner 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 Gartner 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 Gartner 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 Gartner 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 Gartner believes.

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, Kolreuter and Gartner, 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 Gartner 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 Gartner, 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 Gartner 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 Honourable and Reverend W. Herbert. He is as emphatic in his conclusion that some hybrids are perfectly fertile – as fertile as the pure parent-species – as are Kolreuter and Gartner 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 Gartner. 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. 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." 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. 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. 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. 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, etc., have been crossed, yet many of these hybrids seed freely. 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." 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 Rhododendron Ponticum and Catawbiense, and that this hybrid "seeds as freely as it is possible to imagine." Had hybrids, when fairly treated, gone on decreasing in fertility in each successive generation, as Gartner 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. 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. 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. 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.

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 it 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. 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 Gartner'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.