Fig. 194.—Diurnal record of the terminal leaflet of Desmodium gyrans. Up-curve represents movement of closure.
The diurnal curve (Fig. 194) exhibits a sudden flexure at about 5 p.m. on the rapid waning of afternoon light till, by 6-30 p.m., it becomes closely pressed against the petiole, by the rapid fall of the leaflet. The discriminating test, between effects of variation of temperature and of light, lies in the fact that the flexure of the diurnal curve takes place in the former at about 2 p.m. when temperature undergoes change from ascent to descent; in the case of light, the change in the intensity of light begins to be marked about three hours later. In the diurnal curve of Desmodium the record shows little change at 2 p.m., showing that the leaflet is not affected to any great extent by the variation of temperature; it is, however, strongly affected by change in light as seen in the rapid closure movement about 5 p.m. The leaflet remains tightly closed throughout the night and begins to open and spread out early in the morning at about 5 a.m. This up-movement is also very rapid and the leaflet assumes the fullest outspread position by 7 a.m. It remains in this position till the afternoon, after which the cycle becomes repeated. As the leaflet is very sensitive to light, the position of equilibrium of the leaflet is liable to be disturbed by the slightest variation of light and the fluctuation of light from the sky often gives rise to a wavy outline in the record. The leaflet, moreover, has a tendency to exhibit rhythmic pulsations.
In the leaflets of Cassia and Desmodium, the daily movement is thus brought about by the predominant action of recurrent light and darkness.
MIDDAY SLEEP.
I shall here briefly recapitulate the results given in greater detail in an earlier paper (p. 352). I have shown that the midday closure of leaflets is brought about by the excitatory action of strong sunlight. The responsive movement of motile pulvinus under diffuse stimulus is determined by the greater contraction of the more excitable half of the organ. Under the action of the midday sun the leaflets of Mimosa undergo a folding upwards, whereas the leaflets of Averrhoa carambola a folding downwards. The explanation of the difference lies in the fact that in the leaflets of Mimosa it is the upper half, and in Averrhoa it is the lower half of the pulvinule, that is the more excitable. This difference may be demonstrated by the action of diffuse electric shock under which the leaflets of Mimosa exhibit an upward, and those of Averrhoa a downward, closure. I have also shown that conduction of excitation takes place across the pulvinule; hence the strong excitation caused by sunlight becomes internally diffused, and brings about the responsive movements, the direction of which is determined by the more excitable half of the pulvinule.
SUMMARY.
Rise of temperature induces a movement of closure of the leaflet of Cassia, fall of temperature inducing the opposite movement.
Artificial darkness induces a closure of the leaflets, the closure being completed in the course of an hour. On readmission of light, the leaflets become fully expanded in the course of one hour and a quarter. The leaflets are extremely sensitive to light, closure movement being induced by the transitory passage of a cloud.
The effect of rise of temperature is antagonistic to the action of light. The movement of opening during the course of the day is due to the effect of light overpowering the effect of rise of temperature.
Under daily variation of light and darkness, the movement of closure is initiated at about 5 p.m., when the light is undergoing a rapid diminution. The movement of closure is complete by 9 p.m. The leaflets remain closed till about 5 a.m. next morning, after which they begin to open and become fully expanded by 9 a.m.