The Different Forms of Flowers on Plants of the Same Species, Charles Robert Darwin [tohfa e dulha read online .TXT] 📗

der Aurikel" 'Zeitschr. des Deutschen und Oest. Alpen-Vereins' Band 6 page 52. Also 'Die Primulaceen-Bastarten' 'Oest. Botanische Zeitschrift' 1835 Numbers 3, 4 and 5.)
This species is heterostyled, like the preceding ones; but amongst the varieties distributed by florists the long-styled form is rare, as it is not valued. There is a much greater relative inequality in the length of the pistil and stamens in the two forms of the auricula than in the cowslip; the pistil in the long-styled being nearly four times as long as that in the short-styled, in which it is barely longer than the ovarium. The stigma is nearly of the same shape in both forms, but is rougher in the long-styled, though the difference is not so great as between the two forms of the cowslip. In the long-styled plants the stamens are very short, rising but little above the ovarium. The pollen-grains of these short stamens, when distended with water, were barely 5/6000 of an inch in diameter, whereas those from the long stamens of the short-styled plants were barely 7/6000, showing a relative difference of about 71 to 100. The smaller grains of the long-styled plant are also much more transparent, and before distention with water more triangular in outline than those of the other form. Mr. Scott compared ten plants of both forms growing under similar conditions, and found that, although the long-styled plant produced more umbels and more capsules than the short-styled, yet they yielded fewer seeds, in the ratio of 66 to 100. (1/12. 'Journal of the Linnean Society Botany' volume 8 1864 page 86.) Three short-styled plants were protected by me from the access of insects, and they did not produce a single seed. Mr. Scott protected six plants of both forms, and found them excessively sterile. The pistil of the long-styled form stands so high above the anthers, that it is scarcely possible that pollen should reach the stigma without some aid; and one of Mr. Scott's long-styled plants which yielded a few seeds (only 18 in number) was infested by aphides, and he does not doubt that these had imperfectly fertilised it.
I tried a few experiments by reciprocally fertilising the two forms in the same manner as before, but my plants were unhealthy, so I will give, in a condensed form, the results of Mr. Scott's experiments. For fuller particulars with respect to this and the five following species, the paper lately referred to may be consulted. In each case the fertility of the two legitimate unions, taken together, is compared with that of the two illegitimate unions together, by the same two standards as before, namely, by the proportional number of flowers which produced good capsules, and by the average number of seeds per capsule. The fertility of the legitimate unions is always taken at 100.
By the first standard, the fertility of the two legitimate unions of the auricula is to that of the two illegitimate unions as 100 to 80; and by the second standard as 100 to 15.
Primula Sikkimensis.
According to Mr. Scott, the pistil of the long-styled form is fully four times as long as that of the short-styled, but their stigmas are nearly alike in shape and roughness. The stamens do not differ so much in relative length as the pistils. The pollen-grains differ in a marked manner in the two forms; "those of the long-styled plants are sharply triquetrous, smaller, and more transparent than those of the short-styled, which are of a bluntly triangular form." The fertility of the two legitimate unions to that of the two illegitimate unions is by the first standard as 100 to 95, and by the second standard as 100 to 31.
Primula cortusoides.
The pistil of the long-styled form is about thrice as long as that of the short- styled, the stigma being double as long and covered with much longer papillae. The pollen-grains of the short-styled form are, as usual, "larger, less transparent, and more bluntly triangular than those from the long-styled plants." The fertility of the two legitimate unions to that of the two illegitimate unions is by the first standard as 100 to 74, and by the second standard as 100 to 66.
Primula involucrata.
The pistil of the long-styled form is about thrice as long as that of the short- styled; the stigma of the former is globular and closely beset with papillae, whilst that of the short-styled is smooth and depressed on the apex. The pollen- grains of the two forms differ in size and transparency as before, but not in shape. The fertility of the two legitimate to that of the two illegitimate unions is by the first standard as 100 to 72; and by the second standard as 100 to 47.
Primula farinosa.
According to Mr. Scott, the pistil of the long-styled form is only about twice as long as that of the short-styled. The stigmas of the two forms differ but little in shape. The pollen-grains differ in the usual manner in size, but not in form. The fertility of the two legitimate to that of the two illegitimate unions is by the first standard as 100 to 71, and by the second standard as 100 to 44.]
SUMMARY ON THE FOREGOING HETEROSTYLED SPECIES OF PRIMULA.
TABLE 1.12. Summary on the Fertility of the two Legitimate Unions, compared with that of the two Illegitimate Unions, in the genus Primula. The former taken at 100.
Column 1: Name of Species. Column 2: Illegitimate Unions, Judged of by the Proportional Number of Flowers which produced Capsules. Column 3: Illegitimate Unions, Judged of by the Average Number (or Weight in some cases) of Seeds per Capsule.
Primula veris : 69 : 65.
Primula elatior : 27 : 75 (Probably too high).
Primula vulgaris : 60 : 54 (Perhaps too low).
Primula Sinensis : 84 : 63.
Primula Sinensis (second trial) : ? : 53. Primula Sinensis (after Hildebrand) : 100 : 42.
Primula auricula (Scott) : 80 : 15.
Primula Sikkimensis (Scott): 95 : 31.
Primula cortusoides (Scott): 74 : 66.
Primula involucrata (Scott): 72 : 48.
Primula farinosa (Scott): 71 : 44.
Average of the nine species : 88.4 : 61.8.
The fertility of the long-and short-styled plants of the above species of Primula, when the two forms are fertilised legitimately, and illegitimately with pollen of the same form taken from a distinct plant, has now been given. The results are seen in Table 1.12; the fertility being judged by two standards, namely, by that of the proportional number of flowers which yielded capsules, and by that of the average number of seeds per capsule. But for full accuracy many more observations, under varied conditions, would be requisite.
With plants of all kinds some flowers generally fail to produce capsules, from various accidental causes; but this source of error has been eliminated, as far as possible, in all the previous cases, by the manner in which the calculations have been made. Supposing, for instance, that 20 flowers were fertilised legitimately and yielded 18 capsules, and that 30 flowers were fertilised illegitimately and yielded 15 capsules, we may assume that on an average an equal proportion of the flowers in both lots would fail to produce capsules from various accidental causes; and the ratio of 18/20 to 15/30, or as 100 to 56 (in whole numbers), would show the proportional number of capsules due to the two methods of fertilisation; and the number 56 would appear in the left-hand column of Table 1.12, and in my other tables. With respect to the average number of seeds per capsule hardly anything need be said: supposing that the legitimately fertilised capsules contained, on an average, 50 seeds, and the illegitimately fertilised capsules 25 seeds; then as 50 is to 25 so is 100 to 50; and the latter number would appear in the right hand column.
It is impossible to look at the above table and doubt that the legitimate unions between the two forms of the above nine species of Primula are much more fertile than the illegitimate unions; although in the latter case pollen was always taken from a distinct plant of the same form. There is, however, no close correspondence in the two rows of figures, which give, according to the two standards, the difference of fertility between the legitimate and illegitimate unions. Thus all the flowers of P. Sinensis which were illegitimately fertilised by Hildebrand produced capsules; but these contained only 42 per cent of the number of seeds yielded by the legitimately fertilised capsules. So again, 95 per cent of the illegitimately fertilised flowers of P. Sikkimensis produced capsules; but these contained only 31 per cent of the number of seeds in the legitimate capsules. On the other hand, with P. elatior only 27 per cent of the illegitimately fertilised flowers yielded capsules; but these contained nearly 75 per cent of the legitimate number of seeds. It appears that the setting of the flowers, that is, the production of capsules whether good or bad, is not so much influenced by legitimate and illegitimate fertilisation as is the number of seeds which the capsules contain. For, as may be seen at the bottom of Table 1.12, 88.4 per cent of the illegitimately fertilised flowers yielded capsules; but these contained only 61.8 per cent of seeds, in comparison, in each case, with the legitimately fertilised flowers and capsules of the same species. There is another point which deserves notice, namely, the relative degree of infertility in the several species of the long-styled and short-styled flowers, when both are illegitimately fertilised. The data may be found in the earlier tables, and in those given by Mr. Scott in the Paper already referred to. If we call the number of seeds per capsule produced by the illegitimately fertilised long-styled flowers 100, the seeds from the illegitimately fertilised short- styled flowers will be represented by the following numbers (Table 1.a.):--
TABLE 1.a.
Primula veris : 71.
Primula elatior : 44 (Probably too low).
Primula vulgaris : 36 (Perhaps too low).
Primula Sinensis : 71.
Primula auricula : 119.
Primula Sikkimensis : 57.
Primula cortusoides : 93.
Primula involucrata : 74.
Primula farinosa : 63.
We thus see that, with the exception of P. auricula, the long-styled flowers of all nine species are more fertile than the short-styled flowers, when both forms are illegitimately fertilised. Whether P. auricula really differs from the other species in this respect I can form no opinion, as the result may have been accidental. The degree of self-fertility of a plant depends on two elements, namely, on the stigma receiving its own pollen and on its more or less efficient action when placed there. Now as the anthers of the short-styled flowers of several species of Primula stand directly above the stigma, their pollen is more likely to fall on it, or to be carried down to it by insects, than in the case of the long-styled form. It appears probable, therefore, at first sight, that the lessened capacity of the short-styled flowers to be fertilised with their own pollen, is a special adaptation for counteracting their greater liability to receive their own pollen, and thus for checking self-fertilisation. But from facts with respect to other species hereafter to be given, this view can hardly be admitted. In accordance with the above liability, when some of the species of Primula were allowed to fertilise themselves spontaneously under a net, all insects being excluded, except such minute ones as Thrips, the short-styled flowers, notwithstanding their greater innate self-sterility, yielded more seed than did the long-styled. None of the species, however, when insects were excluded, made a near approach to full fertility. But the long-styled form of P. Sinensis gave, under these circumstances, a considerable number of seeds, as the corolla in falling off drags the anthers, which are seated low down in the tube, over the stigma, and thus leaves plenty of pollen on it.