In this latter case the movement no doubt depends on alternately greater growth on the upper and lower side of the petiole, or of the blade, or of both, preceded probably by the increased turgescence of the growing cells. Such movements generally last for a very short period-- for instance, with Brassica and Githago for 4 or 5 nights, with Beta for 2 or 3, and with [page 314] Raphanus for only a single night. There are, however, some strong exceptions to this rule, as the cotyledons of Gossypium, Anoda and Ipomoea do not possess pulvini, yet continue to move and to grow for a long time. We thought at first that when the movement lasted for only 2 or 3 nights, it could hardly be of any service to the plant, and hardly deserved to be called sleep; but as many quickly-growing leaves sleep for only a few nights, and as cotyledons are rapidly developed and soon complete their growth, this doubt now seems to us not well-founded, more especially as these movements are in many instances so strongly pronounced. We may here mention another point of similarity between sleeping leaves and cotyledons, namely, that some of the latter (for instance, those of Cassia and Githago) are easily affected by the absence of light; and they then either close, or if closed do not open; whereas others (as with the cotyledons of Oxalis) are very little affected by light. In the next chapter it will be shown that the nyctitropic movements both of cotyledons and leaves consist of a modified form of circumnutation.

As in the Leguminosae and Oxalidae, the leaves and the cotyledons of the same species generally sleep, the idea at first naturally occurred to us, that the sleep of the cotyledons was merely an early development of a habit proper to a more advanced stage of life. But no such explanation can be admitted, although there seems to be some connection, as might have been expected, between the two sets of cases. For the leaves of many plants sleep, whilst their cotyledons do not do so--of which fact Desmodium gyrans offers a good instance, as likewise do three species of Nicotiana observed by us; also Sida rhombifolia, Abutilon Darwinii, and Chenopodium album. On the other [page 315] hand, the cotyledons of some plants sleep and not the leaves, as with the species of Beta, Brassica, Geranium, Apium, Solanum, and Mirabilis, named in our list. Still more striking is the fact that, in the same genus, the leaves of several or of all the species may sleep, but the cotyledons of only some of them, as occurs with Trifolium, Lotus, Gossypium, and partially with Oxalis. Again, when both the cotyledons and the leaves of the same plant sleep, their movements may be of a widely dissimilar nature: thus with Cassia the cotyledons rise vertically up at night, whilst their leaves sink down and twist round so as to turn their lower surfaces outwards. With seedlings of Oxalis Valdiviana, having 2 or 3 well-developed leaves, it was a curious spectacle to behold at night each leaflet folded inwards and hanging perpendicularly downwards, whilst at the same time and on the same plant the cotyledons stood vertically upwards.

These several facts, showing the independence of the nocturnal movements of the leaves and cotyledons on the same plant, and on plants belonging to the same genus, lead to the belief that the cotyledons have acquired their power of movement for some special purpose. Other facts lead to the same conclusion, such as the presence of pulvini, by the aid of which the nocturnal movement is continued during some weeks. In Oxalis the cotyledons of some species move vertically upwards, and of others vertically downwards at night; but this great difference within the same natural genus is not so surprising as it may at first appear, seeing that the cotyledons of all the species are continually oscillating up and down during the day, so that a small cause might determine whether they should rise or sink at night. Again, the peculiar nocturnal movement of the left-hand coty- [page 316] ledon of Trifolium strictum, in combination with that of the first true leaf.

Charles Darwin

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