(8/28. The case of Asclepias was described by R. Brown. Baillon asserts 'Adansonia' tome 2 1862 page 58, that with many plants the tubes are emitted from pollen-grains which have not come into contact with the stigma; and that they may be seen advancing horizontally through the air towards the stigma. I have observed the emission of the tubes from the pollen-masses whilst still within the anthers, in three widely distinct Orchidean genera namely Aceras, Malaxis, and Neottia: see 'The Various Contrivances by which Orchids are Fertilised' 2nd edition page 258.) It is, however, a wonderful sight to behold the tubes directing themselves in a straight line to the stigma, when this is at some little distance from the anthers. As soon as they reach the stigma or the open passage leading into the ovarium, no doubt they penetrate it, guided by the same means, whatever these may be, as in the case of ordinary flowers. I thought that they might be guided by the avoidance of light: some pollen-grains of a willow were therefore immersed in an extremely weak solution of honey, and the vessel was placed so that the light entered only in one direction, laterally or from below or from above, but the long tubes were in each case protruded in every possible direction.

As cleistogamic flowers are completely closed they are necessarily self- fertilised, not to mention the absence of any attraction to insects; and they thus differ widely from the great majority of ordinary flowers. Delpino believes that cleistogamic flowers have been developed in order to ensure the production of seeds under climatic or other conditions which tend to prevent the fertilisation of the perfect flowers. (8/29. 'Sull' Opera la Distribuzione dei Sessi nelle Piante' 1867 page 30.) I do not doubt that this holds good to a certain limited extent, but the production of a large supply of seeds with little consumption of nutrient matter or expenditure of vital force is probably a far more efficient motive power. The whole flower is much reduced in size; but what is much more important, an extremely small quantity of pollen has to be formed, as none is lost through the action of insects or the weather; and pollen contains much nitrogen and phosphorus. Von Mohl estimated that a single cleistogamic anther-cell of Oxalis acetosella contained from one to two dozen pollen-grains; we will say 20, and if so the whole flower can have produced at most 400 grains; with Impatiens the whole number may be estimated in the same manner at 250; with Leersia at 210; and with Viola nana at only 100. These figures are wonderfully low compared with the 243,600 pollen-grains produced by a flower of Leontodon, the 4,863 by an Hibiscus, or the 3,654,000 by a Paeony. (8/30. The authorities for these statements are given in my 'Effects of Cross and Self-Fertilisation' page 376.) We thus see that cleistogamic flowers produce seeds with a wonderfully small expenditure of pollen; and they produce as a general rule quite as many seeds as the perfect flowers.

That the production of a large number of seeds is necessary or beneficial to many plants needs no evidence. So of course is their preservation before they are ready for germination; and it is one of the many remarkable peculiarities of the plants which bear cleistogamic flowers, that an incomparably larger proportion of them than of ordinary plants bury their young ovaries in the ground;--an action which it may be presumed serves to protect them from being devoured by birds or other enemies. But this advantage is accompanied by the loss of the power of wide dissemination. No less than eight of the genera in the list at the beginning of this chapter include species which act in this manner, namely, several kinds of Viola, Oxalis, Vandellia, Linaria, Commelina, and at least three genera of Leguminosae. The seeds also of Leersia, though not buried, are concealed in the most perfect manner within the sheaths of the leaves. Cleistogamic flowers possess great facilities for burying their young ovaries or capsules, owing to their small size, pointed shape, closed condition and the absence of a corolla; and we can thus understand how it is that so many of them have acquired this curious habit.

Charles Darwin

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