Another instance has been given (11/146. 'Transact. Hort. Soc.' volume 5 pages 65 and 68. See also Prof. Hildebrand with a coloured figure in 'Bot. Zeitung' May 15, 1868 page 327. Puvis also has collected 'De La Degeneration' 1837 page 36) several other instances; but it is not in all cases possible to distinguish between the direct action of foreign pollen and bud-variations.) of two very different apple-trees growing close to each other, which bore fruit resembling each other, but only on the adjoining branches. It is, however, almost superfluous to adduce these or other cases, after that of the St. Valery apple, the flowers which, from the abortion of the stamens, do not produce pollen, but are fertilised by the girls of the neighbourhood with pollen of many kinds; and they bear fruit, "differing from one another in size, flavour, and colour, but resembling in character the hermaphrodite kinds by which they have been fertilised." (11/147. T. de Clermont-Tonnerre in 'Mem. de la Soc. Linn. de Paris' tome 3 1825 page 164.)]

I have now shown, on the authority of several excellent observers, in the case of plants belonging to widely different orders, that the pollen of one species or variety, when applied to the female of a distinct form, occasionally causes the coats of the seeds, the ovarium or fruit, including even the calyx and upper part of the peduncle of the apple, and the axis of the ear in maize, to be modified. Sometimes the whole ovarium or all the seeds are thus affected; sometimes only a certain number of the seeds, as in the case of the pea, or only a part of the ovarium, as with the striped orange, mottled grapes, and maize, is thus affected. It must not be supposed that any direct or immediate effect invariably follows the use of foreign pollen: this is far from being the case; nor is it known on what conditions the result depends. Mr. Knight (11/148. 'Transact. of Hort. Soc.' volume 5 page 68.) expressly states that he has never seen the fruit thus affected, though he crossed thousands of apple and other fruit-trees.

There is not the least reason to believe that a branch which has borne seed or fruit directly modified by foreign pollen is itself affected, so as afterwards to produce modified buds; such an occurrence, from the temporary connection of the flower with the stem, would be hardly possible. Hence, but very few, if any, of the cases of bud-variation in the fruit of trees, given in the early part of this chapter can be accounted for by the action of foreign pollen; for such fruits have commonly been propagated by budding or grafting. It is also obvious that changes of colour in flowers, which necessarily supervene long before they are ready for fertilisation, and changes in the shape or colour of leaves, when due to the appearance of modified buds, can have no relation to the action of foreign pollen.

The proofs of the action of foreign pollen on the mother-plant have been given in considerable detail, because this action, as we shall see in a future chapter, is of the highest theoretical importance, and because it is in itself a remarkable and apparently anomalous circumstance. That it is remarkable under a physiological point of view is clear, for the male element not only affects, in accordance with its proper function, the germ, but at the same time various parts of the mother-plant, in the same manner, as it affects the same part in the seminal offspring from the same two parents. We thus learn that an ovule is not indispensable for the reception of the influence of the male element. But this direct action of the male element is not so anomalous as it at first appears, for it comes into play in the ordinary fertilisation of many flowers. Gartner gradually increased the number of pollen grains until he succeeded in fertilising a Malva, and has (11/149. Beitrage zur Kenntniss der Befruchtung' 1844 s. 347-351.) proved that many grains are first expended in the development, or, as he expresses it, in the satiation, of the pistil and ovarium.

Charles Darwin

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