The Variation of Animals and Plants under Domestication — Volume 2. Darwin Charles

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      [With respect to sterility from the crossing of domestic races, I know of no well-ascertained case with animals. This fact, seeing the great difference in structure between some breeds of pigeons, fowls, pigs, dogs, etc., is extraordinary, in contrast with the sterility of many closely allied natural species when crossed; but we shall hereafter attempt to show that it is not so extraordinary as it at first appears. And it may be well here to recall to mind that the amount of external difference between two species is not a safe guide for predicting whether or not they will breed together, — some closely allied species when crossed being utterly sterile, and others which are extremely unlike being moderately fertile. I have said that no case of sterility in crossed races rests on satisfactory evidence; but here is one which at first seems trustworthy. Mr. Youatt (16/9. 'Cattle' page 202.) and a better authority cannot be quoted, states, that formerly in Lancashire crosses were frequently made between longhorn and shorthorn cattle; the first cross was excellent, but the produce was uncertain; in the third or fourth generation the cows were bad milkers; "in addition to which, there was much uncertainty whether the cows would conceive; and full one-third of the cows among some of these half-breds failed to be in calf." This at first seems a good case: but Mr. Wilkinson states (16/10. Mr. J. Wilkinson in 'Remarks addressed to Sir J. Sebright' 1820 page 38.), that a breed derived from this same cross was actually established in another part of England; and if it had failed in fertility, the fact would surely have been noticed. Moreover, supposing that Mr. Youatt had proved his case, it might be argued that the sterility was wholly due to the two parent-breeds being descended from primordially distinct species.

      In the case of plants Gartner states that he fertilised thirteen heads (and subsequently nine others) on a dwarf maize bearing yellow seed (16/11. 'Bastarderzeugung' s. 87, 169. See also the Table at the end of volume.) with pollen of a tall maize having red seed; and one head alone produced good seed, but only five in number. Though these plants are monoecious, and therefore do not require castration, yet I should have suspected some accident in the manipulation, had not Gartner expressly stated that he had during many years grown these two varieties together, and they did not spontaneously cross; and this, considering that the plants are monoecious and abound with pollen, and are well known generally to cross freely, seems explicable only on the belief that these two varieties are in some degree mutually infertile. The hybrid plants raised from the above five seeds were intermediate in structure, extremely variable, and perfectly fertile. (16/12. 'Bastarderzeugung' s. 87, 577.) In like manner Prof. Hildebrand (16/13. 'Bot. Zeitung' 1868 page 327.) could not succeed in fertilising the female flowers of a plant bearing brown grains with pollen from a certain kind bearing yellow grains; although other flowers on the same plant, which were fertilised with their own pollen, yielded good seed. No one, I believe, even suspects that these varieties of maize are distinct species; but had the hybrids been in the least sterile, no doubt Gartner would at once have so classed them. I may here remark, that with undoubted species there is not necessarily any close relation between the sterility of a first cross and that of the hybrid offspring. Some species can be crossed with facility, but produce utterly sterile hybrids; others can be crossed with extreme difficulty, but the hybrids when produced are moderately fertile. I am not aware, however, of any instance quite like this of the maize, namely, of a first cross made with difficulty, but yielding perfectly fertile hybrids. (16/14. Mr. Shirreff formerly thought ('Gardener's Chronicle' 1858 page 771) that the offspring from a cross between certain varieties of wheat became sterile in the fourth generation; but he now admits ('Improvement of the Cereals' 1873) that this was an error.)

      The following case is much more remarkable, and evidently perplexed Gartner, whose strong wish it was to draw a broad line of distinction between species and varieties. In the genus Verbascum, he made, during eighteen years, a vast number of experiments, and crossed no less than 1085 flowers and counted their seeds. Many of these experiments consisted in crossing white and yellow varieties of both V. lychnitis and V. blattaria with nine other species and their hybrids. That the white and yellow flowered plants of these two species are really varieties, no one has doubted; and Gartner actually raised in the case of both species one variety from the seed of the other. Now in two of his works (16/15. 'Kenntniss der Befruchtung' s. 137; 'Bastarderzeugung' s. 92, 181. On raising the two varieties from seed see s. 307.) he distinctly asserts that crosses between similarly-coloured flowers yield more seed than between dissimilarly-coloured; so that the yellow-flowered variety of either species (and conversely with the white-flowered variety), when crossed with pollen of its own kind, yields more seed than when crossed with that of the white variety; and so it is when differently coloured species are crossed. The general results may be seen in the Table at the end of his volume. In one instance he gives (16/16. 'Bastarderzeugung' s. 216.) the following details; but I must premise that Gartner, to avoid exaggerating the degree of sterility in his crosses, always compares the MAXIMUM number obtained from a cross with the AVERAGE number naturally given by the pure mother-plant. The white variety of V. lychnitis, naturally fertilised by its own pollen, gave from an AVERAGE of twelve capsules ninety-six good seeds in each; whilst twenty flowers fertilised with pollen from the yellow variety of this same species, gave as the MAXIMUM only eighty-nine good seeds; so that we have the proportion of 1000 to 908, according to Gartner's usual scale. I should have thought it possible that so small a difference in fertility might have been accounted for by the evil effects of the necessary castration; but Gartner shows that the white variety of V. lychnitis, when fertilised first by the white variety of V. blattaria, and then by the yellow variety of this species, yielded seed in the proportion of 622 to 438; and in both these cases castration was performed. Now the sterility which results from the crossing of the differently coloured varieties of the same species, is fully as great as that which occurs in many cases when distinct species are crossed. Unfortunately Gartner compared the results of the first unions alone, and not the sterility of the two sets of hybrids produced from the white variety of V. lychnitis when fertilised by the white and yellow varieties of V. blattaria, for it is probable that they would have differed in this respect.

      Mr. J. Scott has given me the results of a series of experiments on Verbascum, made by him in the Botanic Gardens of Edinburgh. (16/17. The results have since been published in 'Journ. Asiatic Soc. of Bengal' 1867 page 145.) He repeated some of Gartner's experiments on distinct species, but obtained only fluctuating results, some confirmatory, the greater number contradictory; nevertheless these seem hardly sufficient to overthrow the conclusion arrived at by Gartner from experiments tried on a larger scale. Mr. Scott also experimented on the relative fertility of unions between similarly and dissimilarly-coloured varieties of the same species. Thus he fertilised six flowers of the yellow variety of V. lychnitis by its own pollen, and obtained six capsules; and calling, for the sake of comparison, the average number of good seed in each of their capsules one hundred, he found that this same yellow variety, when fertilised by the white variety, yielded from seven capsules an average of ninety-four seed. On the same principle, the white variety of V. lychnitis by its own pollen (from six capsules), and by the pollen of the yellow variety (eight capsules), yielded seed in the proportion of 100 to 82. The yellow variety of V. thapsus by its own pollen (eight capsules), and by that of the white variety (only two capsules), yielded seed in the proportion of 100 to 94. Lastly, the white variety of V. blattaria by its own pollen (eight capsules), and by that of the yellow variety (five capsules), yielded seed in the proportion of 100 to 79. So that in every case the unions of similarly-coloured varieties of the same species were more fertile than the unions of dissimilarly-coloured varieties; when all the cases are grouped together, the difference of fertility is as 100 to 86. Some additional trials were made, and altogether thirty-six similarly-coloured unions yielded thirty-five good capsules; whilst thirty-five dissimilarly- coloured unions yielded only twenty-six good capsules. Besides the foregoing experiments, the purple V. phoeniceum was crossed by a rose-coloured and a white variety of the same species; these two varieties were also crossed together, and these several unions yielded less seed than V. phoeniceum by its own pollen. Hence it follows from Mr. Scott's experiments, that in the genus Verbascum the similarly and dissimilarly-coloured varieties of the same species behave, when crossed, like closely allied but distinct species. (16/18. The following facts, given by Kolreuter in his 'Dritte Fortsetzung' ss. 34, 39, appear at first sight strongly to confirm Mr. Scott's and Gartner's statements; СКАЧАТЬ