Now, if a plant of this kind were converted into one that slept, one side of one of the several ellipses which each leaf daily describes, would have to be greatly increased in length in the evening, until the leaf stood vertically, when it would go on circumnutating about the same spot. On the following morning, the side of another ellipse would have to be similarly increased in length so as to bring the leaf back again into its diurnal position, when it would again circumnutate [page 411] until the evening. If the reader will look, for instance, at the diagram (Fig. 142, p. 351), representing the nyctitropic movements of the terminal leaflet of Trifolium subterraneum, remembering that the curved broken lines at the top ought to be prolonged much higher up, he will see that the great rise in the evening and the great fall in the morning together form a large ellipse like one of those described during the daytime, differing only in size. Or, he may look at the diagram (Fig. 103, p. 236) of the 3 ½ ellipses described in the course of 6 h. 35 m. by a leaf of Lupinus speciosus, which is one of the species in this genus that does not sleep; and he will see that by merely prolonging upwards the line which was already rising late in the evening, and bringing it down again next morning, the diagram would represent the movements of a sleeping plant.

With those sleeping plants which describe several ellipses in the daytime, and which travel in a strongly zigzag line, often making in their course minute loops, triangles, etc., if as soon as one of the ellipses begins in the evening to be greatly increased in size, dots are made every 2 or 3 minutes and these are joined, the line then described is almost strictly rectilinear, in strong contrast with the lines made during the daytime. This was observed with Desmodium gyrans and Mimosa pudica. With this latter plant, moreover, the pinnae converge in the evening by a steady movement, whereas during the day they are continually converging and diverging to a slight extent. In all such cases it was scarcely possible to observe the difference in the movement during the day and evening, without being convinced that in the evening the plant saves the expenditure of force by not moving laterally, and that its whole energy is now expended [page 412] in gaining quickly its proper nocturnal position by a direct course. In several other cases, for instance, when a leaf after describing during the day one or more fairly regular ellipses, zigzags much in the evening, it appears as if energy was being expended, so that the great evening rise or fall might coincide with the period of the day proper for this movement.

The most complex of all the movements performed by sleeping plants, is that when leaves or leaflets, after describing in the daytime several vertically directed ellipses, rotate greatly on their axes in the evening, by which twisting movement they occupy a wholly different position at night to what they do during the day. For instance, the terminal leaflets of Cassia not only move vertically downwards in the evening, but twist round, so that their lower surfaces face outwards. Such movements are wholly, or almost wholly, confined to leaflets provided with a pulvinus. But this torsion is not a new kind of movement introduced solely for the purpose of sleep; for it has been shown that some leaflets whilst describing their ordinary ellipses during the daytime rotate slightly, causing their blades to face first to one side and then to another. Although we can see how the slight periodical movements of leaves in a vertical plane could be easily converted into the greater yet simple nyctitropic movements, we do not at present know by what graduated steps the more complex movements, effected by the torsion of the pulvini, have been acquired. A probable explanation could be given in each case only after a close investigation of the movements in all the allied forms.

From the facts and considerations now advanced we may conclude that nyctitropism, or the sleep of leaves [page 413] and cotyledons, is merely a modification of their ordinary circumnutating movement, regulated in its period and amplitude by the alternations of light and darkness.

Charles Darwin

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