Fig. 214.—Record of leaf of Mimosa after amputation of sub-petioles. The leaf fell up to 2-30 p.m., and rose till 5 p.m., after which there is a spasmodic fall. (Successive dots at intervals of 15 minutes.)
Diurnal movement of the amputated petiole: Experiment 223.—In my present experiment the possibility of variation of mechanical movement was obviated by cutting off the end of the petiole, which carried the sub-petioles. The cut end was coated with collodion flexile to prevent evaporation. The intense stimulus caused by amputation induced the excitatory fall of the leaf, but it recovered its normal activity after a period of three hours or so. The diurnal record of the leaf was commenced shortly after 1 p.m.; it will be noticed that the leaf, though deprived of the weight of its sub-petioles, still exhibited a sudden fall at about 5 p.m. (Fig. 214). The fall of the leaf cannot therefore be due to increased mechanical moment. The effect of weight was, moreover, eliminated in torsional response (Expt. 221). In spite of this the leaf exhibited a sudden movement after 5 p.m.
Pfeffer has in his 'Entstehung der Schlafbewegung' (1907) offered another explanation of the sudden fall of the leaf of Mimosa. This, according to him, is not the direct effect of diminished intensity of light in the evening, but is due to the release of the leaf from the phototropic action of light, which, so long as it is sufficiently intense, holds the leaf in the normal position with its upper surface at right angles to the incident rays. Thus, on being set free from the strong action of light, the leaf moves in accordance with the preceding condition of tension; and as this is low the leaf falls, soon to rise again as the tension increases in prolonged darkness.
The above explanation presupposes: (1) that the tension was continuously decreasing till the evening, and (2) that as soon as the phototropic restraint which held the leaf up was removed it fell down in accordance with the prevailing diminished tension.
Referring to the first point, an inspection of the diurnal curve of Mimosa shows that the leaf had no natural tendency to fall towards the evening. There was on the contrary a movement of erection, on account of fall of temperature after the thermal-noon (Fig. 210). As the natural tendency of the leaf was to erect itself, the removal of phototropic restraint cannot therefore induce a movement of fall.
As regards the factor of light, the effect in the afternoon is a down-movement on account of transverse conduction of excitation; but the leaf is prevented from exhibiting this down-movement by the thermo-geotropic up-movement due to fall of temperature after the thermal noon. I shall presently describe experiments on the pure effect of light, which will show that the action of continued photic stimulus induces a down-movement of the leaf in the afternoon.
The results of experiments that have been described show that the sudden fall of the leaf in the evening could not be due to:
(1) increased mechanical moment,
(2) the natural tendency of the leaf to fall towards evening against phototropic action by which the leaf is held up.