The fact of the leaves of several plants not sleeping unless they have been well illuminated during the day, made us for a time doubt whether the protection of their upper surfaces from radiation was in all cases the final cause of their well-pronounced nyctitropic movements. But we have no reason to suppose that the illumination from the open sky, during even the most clouded day, is insufficient for this purpose; and we should bear in mind that leaves which are shaded from being seated low down on the plant, and which sometimes do not sleep, are likewise protected at night from full radiation. Nevertheless, we do not wish to deny that there may exist cases in which leaves change their position considerably at night, without their deriving any benefit from such movements.
Although with sleeping plants the blades almost [page 399] always assume at night a vertical, or nearly vertical position, it is a point of complete indifference whether the apex, or the base, or one of the lateral edges, is directed to the zenith. It is a rule of wide generality, that whenever there is any difference in the degree of exposure to radiation between the upper and the lower surfaces of leaves and leaflets, it is the upper which is the least exposed, as may be seen in Lotus, Cytisus, Trifolium, and other genera. In several species of Lupinus the leaflets do not, and apparently from their structure cannot, place themselves vertically at night, and consequently their upper surfaces, though highly inclined, are more exposed than the lower; and here we have an exception to our rule. But in other species of this genus the leaflets succeed in placing themselves vertically; this, however, is effected by a very unusual movement, namely, by the leaflets on the opposite sides of the same leaf moving in opposite directions.
It is again a very common rule that when leaflets come into close contact with one another, they do so by their upper surfaces, which are thus best protected. In some cases this may be the direct result of their rising vertically; but it is obviously for the protection of the upper surfaces that the leaflets of Cassia rotate in so wonderful a manner whilst sinking downwards; and that the terminal leaflet of Melilotus rotates and moves to one side until it meets the lateral leaflet on the same side. When opposite leaves or leaflets sink vertically down without any twisting, their lower surfaces approach each other and sometimes come into contact; but this is the direct and inevitable result of their position. With many species of Oxalis the lower surfaces of the adjoining leaflets are pressed together, and are thus better protected [page 400] than the upper surfaces; but this depends merely on each leaflet becoming folded at night so as to be able to sink vertically downwards. The torsion or rotation of leaves and leaflets, which occurs in so many cases, apparently always serves to bring their upper surfaces into close approximation with one another,