Origin of Species Variorum

Comparison with 1866

of facts seem to be connected together by some common but unknown bond, which is essentially related to the principle of life; this principle apparently being that life, as Mr. Herbert Spencer has remarked, depends on, or consists in, the incessant action and reaction of various forces, which, as throughout nature, are always tending towards an equilibrium; and when this ten- dency is slightly disturbed by any change, the vital forces apparently gain in power.

Reciprocal Dimorphism and Trimorphism .

The infertility which may be observed in various dimorphic and trimorphic plants, when they are illegitimately fertilised, that is by pollen taken from stamens not corresponding in height with the pistil, differs much in degree, up to absolute and utter sterility; just in the same manner as occurs in crossing distinct species. As the degree of sterility in the latter case depends in an eminent degree on the conditions of life being more or less favourable, so I have found it with illegitimate unions. It is well known that if pollen of a distinct species be placed on the stigma of a flower, and its own pollen be afterwards, even after a considerable interval of time, placed on the same stigma, its action is so strongly prepotent that it generally annihilates the effect of the foreign pollen; so it is with the pollen of the several forms of the same species, for legitimate pollen is strongly prepotent over illegitimate pollen, when both are placed on the same stigma. I ascertained this by fertilising several flowers, first illegitimately, and twenty-four hours afterwards legitimately, with pollen taken from a peculiarly coloured variety, and all the seedlings were similarly coloured; this shows that the legitimate pollen, though applied twenty-four hours subsequently, had wholly destroyed or prevented the action of the previously applied illegitimate pollen. Again, as in making reciprocal crosses between the same two species, there is occasionally a great difference in the result, so something analogous occurs with dimorphic plants; for a short-styled cowslip yields more seed when fertilised by the long-styled form, and less seed when fertilised by its own form, than does a long-styled cowslip when fertilised in the two corresponding methods.

In all these respects the forms of the same undoubted species when illegitimately united behave in exactly the same manner as do two distinct species when crossed. This led me carefully to observe during four years many seedlings, raised from several illegitimate unions. The chief result is that these illegitimate plants, as they may be called, are not fully fertile. It is possible to raise form dimorphic species, both long-styled and short-styled illegitimate plants, and from trimorphic plants all three illegitimate forms; so that these can be properly united in a legitimate manner. When this is done, there is no apparent reason why they should not yield as many seeds as did their parents when legitimately fertilised. But such is not the case; they are all infertile, but in various degrees; some being so utterly and incurably sterile that they did not yield during four seasons a single seed or even a seed-capsule. These illegitimate plants, which are so sterile, although united with each other in a legitimate manner, may be strictly compared with hybrids when crossed inter se, and we all know how sterile these latter generally are. When on the other hand a hybrid is crossed with either pure parent-species, the sterility is usually much lessened: and so it is when an illegitimate plant is fertilised by a legitimate plant. In the same manner as the sterility of hybrids does not always run parallel with the difficulty of making the first cross between its two parent-species, so the sterility of certain illegitimate plants was unusually great, whilst the sterility of the union from which they were derived was by no means great. With hybrids raised from the same seed-capsule the degree of sterility is innately variable, so it is in a marked manner with illegitimate plants. Lastly, many hybrids are profuse and persistent flowerers, whilst other and more sterile hybrids produce few flowers, and are weak, miserable dwarfs; exactly similar cases occur with the illegitimate offspring of various dimorphic and trimorphic plants.

Altogether there is the closest identity in character and behaviour between illegitimate plants and hybrids. It is hardly an exaggeration to maintain that the former are hybrids, but produced within the limits of the same species by the improper union of certain forms, whilst ordinary hybrids are produced from an improper union between so-called distinct species. We have also already seen that there is the closest similarity in all respects between first illegitimate unions and first crosses between distinct species. All this will perhaps be made more fully apparent by an illustration: we may suppose that a botanist found two well-marked varieties (and such occur) of the long-styled form of the trimorphic Lythrum salicaria, and that he determined to try by crossing whether they were specifically distinct. He would find that they yielded only about one-fifth of the proper number of seed, and that they behaved in all the other above specified respects as if they had been two distinct species. But to make the case sure, he would raise plants from his supposed hybridised seed, and he would find that the seedlings were miserably dwarfed and utterly sterile, and that they behaved in all other respects like ordinary hybrids. He might then maintain that he had actually proved, in accordance with the common view, that his two varieties were as good and as distinct species as any in the world; but he would be completely mistaken.

The facts now given on dimorphic and trimorphic plants are of importance, because they show us, firstly, that the physiological test of lessened fertility, both in first crosses and in hybrids, is no safe criterion of specific distinction; secondly, because we are thus led to infer, as previously remarked, that there must be some unknown law or bond connecting the infertility both of illegitimate unions and of first crosses, with the infertility of their illegitimate and hybrid offspring; thirdly, because we find, and this seems to me of especial im- portance, that two or three forms of the same species may exist and may differ in no respect, except in certain characters in their reproductive organs,—such as in the relative lengths of the stamens and pistils, in the size, form, and colour of the pollen-grains, in the structure of the stigma, and in the number and size of the seeds. ↑ ↑ With these differences and with no others, either in organisation or constitution, between the several forms, which are all hermaphrodites, we find that their illegitimate unions and their illegitimate progeny are more or less sterile, and closely resemble in a whole series of relations the first unions and the hybrid offspring of distinct species. We are thus led to infer that the sterility of species when crossed and of their hybrid progeny is likewise in all probability exclusively due to similar differences confined to their reproductive systems. ↑ We are indeed led to his same conclusion from considering reciprocal crosses between the same two species, in which the male of one cannot be united, or can be united with great difficulty, with the female of the other species, whilst the converse cross can be effected with perfect facility; for this difference in the facility of making reciprocal crosses and in the fertility of their offspring must be attributed either to the male or to the female element in the one species having been differentiated, with reference to the other sexual element, in a higher degree than in the second species. That excellent observer, Gärtner, likewise came on general grounds to the same conclusion, namely, that species when crossed are sterile owing to differences confined to their reproductive systems.

Finally, we are naturally led to inquire for what useful end have plants been rendered reciprocally dimorphic and trimorphic? A wide-spread analogy clearly gives us the answer as far as the immediate cause is concerned, namely, to prevent the pollen of each flower acting on the stigma of that flower. We see this effected in a host of flowers by the most curious mechanical contrivances, as I have shown with Orchids, and as could be shown with many plants of many other orders. There are also numerous plants, called dicho-gamous by C. K. Sprengel, in which the pollen and stigma are never mature at the same time, so that these plants can never fertilise themselves. There are many flowers, which, though they have their stigmas and pollen mature together, and which do not present any obstacle to self-impregnation, yet nevertheless are almost always fertilised by surrounding varieties when growing in the vicinity, as shown by the character of their seedlings. Then, again, we have many flowers with separated sexes borne on distinct plants, or on the same, which inevitably prevents self-fertilisation. Lastly, in accordance with the great principle prevailing throughout nature, of the same end being gained by the most diversified means, we find in dimorphic and trimorphic plants, in which self-fertilisation is not checked by any of the above-specified means, that this has been effected by the pollen of each flower, and consequently of all the flowers of the same form, having been rendered more or less impotent on their own stigmas; so that its action is easily and wholly obliterated by pollen habitually brought by insects from other individuals and forms of the same species.

In searching for the cause of dimorphism and trimorphism in plants, we may, in my opinion, safely go one step further, and conclude that the pollen has been prevented acting on the stigma of the same flower, in order to give vigour to the offspring by leading to the union of two distinct individuals. But on this view it is not a little remarkable that the end has been gained, in the case of dimorphic and trimorphic plants, at the expense of all the plants of the same form being rendered more or less sterile when united, and producing more or less sterile offspring. With respect to the steps by which it is probable that plants have been rendered dimorphic and trimorphic, want of space prevents my entering on the subject; but I will add that there is no special difficulty in this having been effected through variability, through the good gained by the prepotency of one sort of pollen over another, and through the accumulative action of natural selection.

Fertility of Varieties when Crossed, and of their Mongrel Offspring .

It may be urged, as a most forcible argu- ment, that there must be some essential distinction between species and varieties, and that there must be some error in all the foregoing remarks, inasmuch as varieties, however much they may differ from each other in external appearance, cross with perfect facility, and yield perfectly fertile offspring. With some exceptions, presently to be given, I fully admit that this is the rule. ↑ But the subject is surrounded by difficulties, for, looking to varieties produced under nature, we are immediately involved in hopeless difficulties; for if two hitherto reputed varieties be found in any degree sterile together, they are at once ranked by most naturalists as species. For instance, the blue and red pimpernel, ... which are considered by most botanists as varieties, are said by Gärtner not to be quite fertile when crossed, and he consequently ranks them as undoubted species. If we thus argue in a circle, the fertility of all varieties produced under nature will assuredly have to be granted.

If we turn to varieties, produced, or supposed to have been produced, under domestication, we are still in- volved in doubt. For when it is stated, for instance, that the German Spitz dog crosses more easily with the fox than do other dogs, or that certain South American indigenous domestic dogs do not readily unite with European dogs, the explanation which will occur to every one, and probably the true one, is that these dogs have descended from .. aboriginally distinct species. Nevertheless the perfect fertility of so many domestic

of facts seem to be connected together by some common but unknown bond, which is essentially related to the principle of life. ↑ ↑

31 blocks not present in 1859 1860 1861; present in 1866 1869 1872

This subject may be here briefly discussed, and will be found to throw considerable light on hybridism. Several plants belonging to distinct orders present two forms, existing together in about equal numbers, which differ in no respect except in their reproductive organs; one form having a long pistil with short stamens, the other a short pistil with long stamens; both with differently sized pollen-grains. With trimorphic plants there are three forms likewise differing in the lengths of their pistils and stamens, in the size and colour of the pollen-grains, and in some other respects; and as in each of the three forms there are two sets of stamens, there are altogether six sets of stamens and three kinds of pistils. These organs are so proportioned in length to each other, that, in any two of the forms, half the stamens in each stand on a level with the stigma of the third form. Now I have shown, and the result has been confirmed by other observers, that, in order to obtain full fertility with these plants, it is necessary that the stigma of the one form should be fertilised by pollen taken form the stamens of corresponding height in the other form. So that with dimorphic species two unions, which may be called legitimate, are fully fertile, and two, which may be called illegitimate, are more or less infertile. With trimorphic species six unions are legitimate or fully fertile, and twelve are illegitimate or more or less infertile. The infertility which may be observed in various dimorphic and trimorphic plants, when they are illegitimately fertilised, that is by pollen taken from stamens not corresponding in height with the pistil, differs much in degree, up to absolute and utter sterility; just in the same manner as occurs in crossing distinct species. As the degree of sterility in the latter case depends in an eminent degree on the conditions of life being more or less favourable, so I have found it with illegitimate unions. It is well known that if pollen of a distinct species be placed on the stigma of a flower, and its own pollen be afterwards, even after a considerable interval of time, placed on the same stigma, its action is so strongly prepotent that it generally annihilates the effect of the foreign pollen; so it is with the pollen of the several forms of the same species, for legitimate pollen is strongly prepotent over illegitimate pollen, when both are placed on the same stigma. I ascertained this by fertilising several flowers, first illegitimately, and twenty-four hours afterwards legitimately, with pollen taken from a peculiarly coloured variety, and all the seedlings were similarly coloured; this shows that the legitimate pollen, though applied twenty-four hours subsequently, had wholly destroyed or prevented the action of the previously applied illegitimate pollen. Again, as in making reciprocal crosses between the same two species, there is occasionally a great difference in the result, so something analogous occurs with dimorphic plants; for a short-styled cowslip yields more seed when fertilised by the long-styled form, and less seed when fertilised by its own form, than does a long-styled cowslip when fertilised in the two corresponding methods. In all these respects the forms of the same undoubted species when illegitimately united behave in exactly the same manner as do two distinct species when crossed. This led me carefully to observe during four years many seedlings, raised from several illegitimate unions. The chief result is that these illegitimate plants, as they may be called, are not fully fertile. It is possible to raise form dimorphic species, both long-styled and short-styled illegitimate plants, and from trimorphic plants all three illegitimate forms; so that these can be properly united in a legitimate manner. When this is done, there is no apparent reason why they should not yield as many seeds as did their parents when legitimately fertilised. But such is not the case; they are all infertile, but in various degrees; some being so utterly and incurably sterile that they did not yield during four seasons a single seed or even a seed-capsule. These illegitimate plants, which are so sterile, although united with each other in a legitimate manner, may be strictly compared with hybrids when crossed inter se, and we all know how sterile these latter generally are. When on the other hand a hybrid is crossed with either pure parent-species, the sterility is usually much lessened: and so it is when an illegitimate plant is fertilised by a legitimate plant. In the same manner as the sterility of hybrids does not always run parallel with the difficulty of making the first cross between its two parent-species, so the sterility of certain illegitimate plants was unusually great, whilst the sterility of the union from which they were derived was by no means great. With hybrids raised from the same seed-capsule the degree of sterility is innately variable, so it is in a marked manner with illegitimate plants. Lastly, many hybrids are profuse and persistent flowerers, whilst other and more sterile hybrids produce few flowers, and are weak, miserable dwarfs; exactly similar cases occur with the illegitimate offspring of various dimorphic and trimorphic plants. Altogether there is the closest identity in character and behaviour between illegitimate plants and hybrids. It is hardly an exaggeration to maintain that the former are hybrids, but produced within the limits of the same species by the improper union of certain forms, whilst ordinary hybrids are produced from an improper union between so-called distinct species. We have also already seen that there is the closest similarity in all respects between first illegitimate unions and first crosses between distinct species. All this will perhaps be made more fully apparent by an illustration: we may suppose that a botanist found two well-marked varieties (and such occur) of the long-styled form of the trimorphic Lythrum salicaria, and that he determined to try by crossing whether they were specifically distinct. He would find that they yielded only about one-fifth of the proper number of seed, and that they behaved in all the other above specified respects as if they had been two distinct species. But to make the case sure, he would raise plants from his supposed hybridised seed, and he would find that the seedlings were miserably dwarfed and utterly sterile, and that they behaved in all other respects like ordinary hybrids. He might then maintain that he had actually proved, in accordance with the common view, that his two varieties were as good and as distinct species as any in the world; but he would be completely mistaken. The facts now given on dimorphic and trimorphic plants are of importance, because they show us, firstly, that the physiological test of lessened fertility, both in first crosses and in hybrids, is no safe criterion of specific distinction; secondly, because we are thus led to infer, as previously remarked, that there must be some unknown law or bond connecting the infertility both of illegitimate unions and of first crosses, with the infertility of their illegitimate and hybrid offspring; thirdly, because we find, and this seems to me of especial im- portance, that two or three forms of the same species may exist and may differ in no respect, except in certain characters in their reproductive organs,—such as in the relative lengths of the stamens and pistils, in the size, form, and colour of the pollen-grains, in the structure of the stigma, and in the number and size of the seeds.

↑ ↑ ↑ ↑

Fertility of Varieties when crossed , and of their Mongrel offspring .—

It may be urged, as a most forcible argument, that there must be some essential distinction between species and varieties, and that there must be some error in all the foregoing remarks, inasmuch as varieties, however much they may differ from each other in external appearance, cross with perfect facility, and yield perfectly fertile offspring. With some exceptions, presently to be given, I fully admit that this is very generally the rule. But the subject is surrounded by difficulties, for looking to varieties produced under nature, ... if two forms hitherto reputed to be varieties be found in any degree sterile together, they are at once ranked by most naturalists as species. For instance, the blue and red pimpernel, the primrose and cowslip, which are considered by many of our best botanists as varieties, are said by Gärtner not to be quite fertile when crossed, and he consequently ranks them as undoubted species. If we thus argue in a circle, the fertility of all varieties produced under nature will assuredly have to be granted.

If we turn to varieties, produced, or supposed to have been produced, under domestication, we are still involved in doubt. For when it is stated, for instance, that the German Spitz dog unites more easily than other dogs with foxes, or that certain South American indigenous domestic dogs do not readily cross with European dogs, the explanation which will occur to every one, and probably the true one, is that these dogs have descended from several aboriginally distinct species. Nevertheless the perfect fertility of so many domestic