The Variation of Animals and Plants Under Domestication, Volume II (of 2). Darwin Charles

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 dissimilarly-coloured varieties of the same species were less fertile than the unions of similarly-coloured varieties; when all the cases are grouped together, the difference of fertility is as 86 to 100. 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. phœniceum 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. phœniceum 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.229

      This remarkable fact of the sexual affinity of similarly-coloured varieties, as observed by Gärtner and Mr. Scott, may not be of very rare occurrence; for the subject has not been attended to by others. The following case is worth giving, partly to show how difficult it is to avoid error. Dr. Herbert230 has remarked that variously-coloured double varieties of the hollyhock (Althæa rosea) may be raised with certainty by seed from plants growing close together. I have been informed that nurserymen who raise seed for sale do not separate their plants; accordingly I procured seed of eighteen named varieties; of these, eleven varieties produced sixty-two plants all perfectly true to their kind; and seven produced forty-nine plants, half of which were true and half false. Mr. Masters of Canterbury has given me a more striking case; he saved seed from a great bed of twenty-four named varieties planted in closely adjoining rows, and each variety reproduced itself truly with only sometimes a shade of difference in tint. Now in the hollyhock the pollen, which is abundant, is matured and nearly all shed before the stigma of the same flower is ready to receive it;231 and as bees covered with pollen incessantly fly from plant to plant, it would appear that adjoining varieties could not escape being crossed. As, however, this does not occur, it appeared to me probable that the pollen of each variety was prepotent on its own stigma over that of all other varieties. But Mr. C. Turner of Slough, well known for his success in the cultivation of this plant, informs me that it is the doubleness of the flowers which prevents the bees gaining access to the pollen and stigma; and he finds that it is difficult even to cross them artificially. Whether this explanation will fully account for varieties in close proximity propagating themselves so truly by seed, I do not know.

      The following cases are worth giving, as they relate to monœcious forms, which do not require, and consequently have not been injured by, castration. Girou de Buzareingues crossed what he designates three varieties of gourd,232 and asserts that their mutual fertilisation is less easy in proportion to the difference which they present. I am aware how imperfectly the forms in this group were until recently known; but Sageret,233 who ranked them according to their mutual fertility, considers the three forms above alluded to as varieties, as does a far higher authority, namely, M. Naudin.234 Sageret235 has observed that certain melons have a greater tendency, whatever the cause may be, to keep true than others; and M. Naudin, who has had such immense experience in this group, informs me that he believes that certain varieties intercross more readily than others of the same species; but he has not proved the truth of this conclusion; the frequent abortion of the pollen near Paris being one great difficulty. Nevertheless, he has grown close together, during seven years, certain forms of Citrullus, which, as they could be artificially crossed with perfect facility and produced fertile offspring, are ranked as varieties; but these forms when not artificially crossed kept true. Many other varieties, on the other hand, in the same group cross with such facility, as M. Naudin repeatedly insists, that without being grown far apart they cannot be kept in the least true.

      Another case, though somewhat different, may be here given, as it is highly remarkable, and is established on excellent evidence. Kölreuter minutely describes five varieties of the common tobacco,236 which were reciprocally crossed, and the offspring were intermediate in character and as fertile as their parents: from this fact Kölreuter inferred that they are really varieties; and no one, as far as I can discover, seems to have doubted that such is the case. He also crossed reciprocally these five varieties with N. glutinosa, and they yielded very sterile hybrids; but those raised from the var. perennis, whether used as the father or mother plant, were not so sterile as the hybrids from the four other varieties.237 So that the sexual capacity of this one variety has certainly been in some degree modified, so as to approach in nature that of N. glutinosa.238

      These facts with respect to plants show that in some few cases certain varieties have had their sexual powers so far modified, that they cross together less readily and yield less seed than other varieties of the same species. We shall presently see that the sexual functions of most animals and plants are eminently liable to be affected by the conditions of life to which they are exposed; and hereafter we shall briefly discuss the conjoint bearing of this and other facts on the difference in fertility between crossed varieties and crossed species.

Domestication eliminates the tendency to Sterility which is general with Species when crossed

      This hypothesis was first propounded by Pallas,239 and has been adopted by several authors. I can find hardly any direct facts in its support; but unfortunately no one has compared, in the case of either animals or plants, the fertility of anciently domesticated varieties, when crossed with a distinct species, with that of the wild parent-species when similarly crossed. No one has compared, for instance, the fertility of Gallus bankiva and of the domesticated fowl, when crossed with a distinct species of Gallus or Phasianus; and the experiment would in all cases be surrounded by many difficulties. Dureau de la Malle, who has so closely studied classical literature, states240 that in the time of the Romans the common mule was produced with more difficulty than at the present day; but whether this statement may be trusted I know not. A much more important, though somewhat different, case is given by M. Groenland,241 namely, that plants, known from their intermediate character and sterility to be hybrids between Ægilops and wheat, have perpetuated themselves under culture since 1857, with a rapid but varying increase of fertility in each generation. In the fourth generation the plants, still retaining their intermediate character, had become as fertile as common cultivated wheat.

      The indirect evidence in favour of the Pallasian doctrine appears to me to be extremely strong. In the earlier chapters I have attempted to show that our various breeds of dogs are descended from several wild species; and this probably is the case with sheep. There can no longer be any doubt that the Zebu or humped Indian ox belongs to a distinct species from European cattle: the latter, moreover, are descended from two or three forms, which may be called either species or wild races, but which co-existed in a state of nature and kept distinct. We have good evidence that our domesticated pigs belong to at least two specific types, S. scrofa and Indica, which probably lived together in a wild state in South-eastern Europe. Now, a widely-extended analogy leads to the belief that if these several allied species, in the wild state or when first reclaimed, had been crossed, they would have exhibited, both in their first unions and in their hybrid offspring, some degree of sterility. Nevertheless the several domesticated races descended from them are now all, as far as can be ascertained, perfectly fertile together. If this reasoning be trustworthy, and it is apparently sound, we must admit the Pallasian doctrine that long-continued domestication tends to eliminate that sterility which is natural to species when crossed in their aboriginal state.

On increased Fertility from Domestication and Cultivation

      Increased fertility from domestication, without any reference to crossing, may be

229

The following facts, given by Kölreuter in his 'Dritte Fortsetzung,' s. 34, 39, appear at first sight strongly to confirm Mr. Scott's and Gärtner's statements; and to a certain limited extent they do so. Kölreuter asserts, from innumerable observations, that insects incessantly carry pollen from one species and variety of Verbascum to another; and I can confirm this assertion; yet he found that the white and yellow varieties of Verbascum lychnitis often grew wild mingled together: moreover, he cultivated these two varieties in considerable numbers during four years in his garden, and they kept true by seed; but when he crossed them, they produced flowers of an intermediate tint. Hence it might have thought that both varieties must have a stronger elective affinity for the pollen of their own variety than for that of the other; this elective affinity, I may add, of each species for its own pollen (Kölreuter, 'Dritte Forts.,' s. 39, and Gärtner, 'Bastarderz.,' passim) being a perfectly well-ascertained power. But the force of the foregoing facts is much lessened by Gärtner's numerous experiments, for, differently from Kölreuter, he never once got ('Bastarderz.,' s. 307) an intermediate tint when he crossed the yellow and white flowered varieties of Verbascum. So that the fact of the white and yellow varieties keeping true to their colour by seed does not prove that they were not mutually fertilised by the pollen carried by insects from one to the other.

230

'Amaryllidaceæ,' 1837, p. 366. Gärtner has made a similar observation.

231

Kölreuter first observed this fact. 'Mém. de l'Acad. St. Petersburg,' vol. iii. p. 197. See also C. K. Sprengel, 'Das Entdeckte Geheimniss,' s. 345.

232

Namely, Barbarines, Pastissons, Giraumous: 'Annal. des Sc. Nat.,' tom. xxx., 1833, pp. 398 and 405.

233

'Mémoire sur les Cucurbitaceæ,' 1826, pp. 46, 55.

234

'Annales des Se. Nat.,' 4th series, tom. vi. M. Naudin considers these forms as undoubtedly varieties of Cucurbita pepo.

235

'Mém. Cucurb.,' p. 8.

236

'Zweite Forts.,' s. 53, namely, Nicotiana major vulgaris; (2) perennis; (3) Transylvanica; (4) a sub-var. of the last; (5) major latifol. fl. alb.

237

Kölreuter was so much struck with this fact that he suspected that a little pollen of N. glutinosa in one of his experiments might have accidentally got mingled with that of var. perennis, and thus aided its fertilising power. But we now know conclusively from Gärtner ('Bastarderz.,' s. 34, 431) that two kinds of pollen never act conjointly on a third species; still less will the pollen of a distinct species, mingled with a plant's own pollen, if the latter be present in sufficient quantity, have any effect. The sole effect of mingling two kinds of pollen is to produce in the same capsule seeds which yield plants, some taking after the one and some after the other parent.

238

Mr. Scott has made some observations on the absolute sterility of a purple and white primrose (Primula vulgaris) when fertilised by pollen from the primrose ('Journal of Proc. of Linn. Soc.,' vol. viii., 1864, p. 98); but these observations require confirmation. I raised a number of purple-flowered long-styled seedlings from seed kindly sent me by Mr. Scott, and, though they were all some degree sterile, they were much more fertile with pollen taken from the common primrose than with their own pollen. Mr. Scott has likewise described a red equal-styled cowslip (P. veris, idem, p. 106), which was found by him to be highly sterile when crossed with the common cowslip; but this was not the case with several equal-styled red seedlings raised by me from his plant. This variety of the cowslip presents the remarkable peculiarity of combining male organs in every respect like those of the short-styled form, with female organs resembling in function and partly in structure those of the long-styled form; so that we have the singular anomaly of the two forms combined in the same flower. Hence it is not surprising that these flowers should be spontaneously self-infertile in a high degree.

239

'Act. Acad. St. Petersburg,' 1780, part ii., pp. 84, 100.

240

'Annales des Sc. Nat.,' tom. xxi. (1st series), p. 61.

241

'Bull. Bot. Soc. de France,' Dec. 27th, 1861, tom. viii. p. 612.