By 8 A.M. on the following morning the leaflets had risen to their proper height. We have seen in the fourth chapter, that the leaves of Lupinus speciosus, which do not sleep, circumnutate to an extraordinary extent, making many ellipses in the course of the day.
Cytisus (Tribe 2), Trigonella and Medicago (Tribe 3).--Only
Fig. 139. Medicago marina: A, leaves during the day; B, leaves asleep at night.
a few observations were made on these three genera. The petioles on a young plant, about a foot in height, of Cytisus fragrans rose at night, on one occasion 23o and on another 33o. The three leaflets also bend upwards, and at the same time [page 345] approach each other, so that the base of the central leaflet overlaps the bases of the two lateral leaflets. They bend up so much that they press against the stem; and on looking down on one of these young plants from vertically above, the lower surfaces of the leaflets are visible; and thus their upper surfaces, in accordance with the general rule, are best protected from radiation. Whilst the leaves on these young plants were thus behaving, those on an old bush in full flower did not sleep at night.
Trigonella Cretica resembles a Melilotus in its sleep, which will be immediately described. According to M. Royer,* the leaves of Medicago maculata rise up at night, and "se renversent un peu de manière à presenter obliquement au ciel leur face inférieure." A drawing is here given (Fig. 139) of the leaves of M. marina awake and asleep; and this would almost serve for Cytisus fragrans in the same two states.
Melilotus (Tribe 3).--The species in this genus sleep in a remarkable manner. The three leaflets of each leaf twist through an angle of 90o, so that their blades stand vertically at night with one lateral edge presented to the zenith (Fig. 140). We shall best understand the other and more complicated movements, if we imagine ourselves always to hold the leaf with the tip of the terminal leaflet pointed to the north. The leaflets in becoming vertical at night could of course twist so that their upper surfaces should face to either side; but the two lateral leaflets always twist so that this surface tends to face the north, but as they move at the same time towards the terminal leaflet, the upper surface of the one faces about N.N.W., and that of the other N.N.E. The terminal leaflet behaves differently, for it twists to either side, the upper surface facing sometimes east and sometimes west, but rather more commonly west than east. The terminal leaflet also moves in another and more remarkable manner, for whilst its blade is twisting and becoming vertical, the whole leaflet bends to one side, and invariably to the side towards which the upper surface is directed; so that if this surface faces the west the whole leaflet bends to the west, until it comes into contact with the upper and vertical surface of the western lateral leaflet. Thus the upper surface of the terminal and of one of the two lateral leaflets is well protected.
The fact of the terminal leaflet twisting indifferently to either
* 'Annales des Sc. Nat. Bot.' (5th series), ix. 1868, p. 368. [page 346]
side and afterwards bending to the same side, seemed to us so remarkable, that we endeavoured to discover the cause. We imagined that at the commencement of the movement it might be determined by one of the two halves of the leaflet being a little heavier than the other. Therefore bits of wood were gummed on one side of several leaflets, but this produced no effect; and they continued to twist in the same direction as
Fig. 140. Melilotus officinalis: A, leaf during the daytime. B, another leaf asleep. C, a leaf asleep as viewed from vertically above; but in this case the terminal leaflet did not happen to be in such close contact with the lateral one, as is usual.
they had previously done. In order to discover whether the same leaflet twisted permanently in the same direction, black threads were tied to 20 leaves, the terminal leaflets of which twisted so that their upper surfaces faced west, and 14 white threads to leaflets which twisted to the east. These were observed occasionally during 14 days, and they all continued, with a single exception, to twist and bend in the same direction; for [page 347] one leaflet, which had originally faced east, was observed after 9 days to face west.