XLVII.--DIURNAL MOVEMENT DUE TO ALTERNATION OF LIGHT AND DARKNESS
_By_
SIR J. C. BOSE,
_a.s.sisted by_
LALIT MOHAN MUKHERJI, B.Sc. (_Nawroji Scholar_).
The nyct.i.tropic movements of the leaflet of _Ca.s.sia alata_ and of the terminal leaflet of _Desmodium gyrans_ furnish us with typical examples of the recurrent effects of light and darkness. The petiole of _Ca.s.sia_ contains a number of paired leaflets each of which is about 5 cm. long and 25 cm. broad. The leaflets are extremely sensitive to light; at night each pair of leaflets fold themselves in a forward direction (see Fig. 150). With the appearance of light they open at first in a lateral direction; later on there is a twist of the pulvinus by which the inner surface of the leaflets faces light coming from above (p. 405). I shall show that the diurnal movements of the leaflets are predominantly due to phototropic action.
Before proceeding further it will be necessary to give a general description of the experimental method employed, and of the apparatus by which diurnal movements are recorded.
EXPERIMENTAL ARRANGEMENTS.
The diurnal record is often taken continuously for several days, and it is therefore necessary to take precautions against the disturbing effect of watering the plant. The record is also liable to be affected by the twist induced by light when it acts on one side of the organ.
_Irrigation._--There is, as is well known, a periodic variation of turgor in the plant. This normal variation is, however, disturbed by watering the plant at irregular intervals. Precaution against this was taken by placing the three flower pots on a long trough filled with water (Fig. 190). The height of water in the trough is always maintained constant by a syphon.
_Vertical illumination._--The direction of sunlight changes from morning to evening, and the leaves exhibit appropriate phototropic movements or torsions under changing directions of lateral light. In order to obviate this, a special chamber was constructed, which allowed light from the sky to fall vertically on the plant through a sheet of ground gla.s.s which covered the roof. The sides and the base of the chamber are impervious to light. A narrow slit covered with red gla.s.s allows inspection of the curve during the process of record.
_The Ventilator._--A revolving ventilator, acted on by the wind, sucks the air away from the chamber, thus ensuring constant supply of fresh air, without causing any disturbances of the record.
_The Recorder._--The Oscillating Recorder employed is of the quadruplex type carrying four recording plates (Fig. 190). The first lever records the daily variation of temperature. The other three are attached to three different specimens of the same plant, or to three different plants. In the former case, three records are obtained of the same species of plant, under identical external condition. If they agree in all essentials, the periodic curve may be taken as characteristic of the given plant. A very great saving of time is thus ensured, and it is thus possible to obtain characteristic curves of numbers of different species of plants within the short period of a season. The quadruplex recorder enables us also to obtain simultaneous records under identical external condition of leaves of different age of the same plant, or of leaves of three different species of plant. I have for the last five years taken records of numerous plants at all seasons of the year. The autograph of the plant is often so characteristic that it is possible to name it by mere inspection of its daily record.
[Ill.u.s.tration: FIG. 190.--The Nyct.i.tropic Recorder with four writing levers. The flower pots are placed in a trough filled with water to a constant height. The first two levers are shown in the figure to record movements of leaves, the third to record movement of a horizontally laid shoot; the fourth lever attached to a differential thermometer, T, records diurnal variation of temperature.]
_Thermograph._--For obtaining a continuous record of diurnal variation of temperature, I use a compound strip, T, made of bra.s.s and steel.
Variation of temperature induces a curvature of the compound strip which is recorded by means of the attached lever. The oscillation of the plate takes place once in fifteen minutes, and the successive dots thus produced give time records of the diurnal curve. The record thus consists of a series of dots. An additional device makes the plate oscillate three times in rapid succession at the end of each hour; the hourly dot is thus thicker than others. The movement of the plant, corresponding to the particular variation of temperature at any period, may thus be easily determined. I shall now give a typical example of diurnal movement induced by variation of light and darkness.
DIURNAL MOVEMENT OF THE LEAFLET OF _Ca.s.sia alata_.
The leaflet of _Ca.s.sia alata_ exhibits a movement of opening in the morning, and it remains outspread throughout the day. It then begins to close before evening and remains closed throughout the night. The problem before us is to find out the relative importance of variation of temperature and of light in the diurnal movement of the leaflets.
In the daytime the light is increasing till midday; there is, on the other hand, a rapid decline of light after 5 p.m. and uninterrupted darkness at night. As regards temperature there is a continuous rise from morning till the thermal noon at 2 p.m., after which the fall of temperature is continuous till next morning. The opening of the leaflets in the daytime may therefore be due to the summated effects of rising temperature and increasing light, the closure, on the other hand, being due to falling temperature, and to darkness. The individual effect of each of these factors is not known and it is therefore necessary to determine the effects of variation of temperature and of light.
EFFECT OF VARIATION OF TEMPERATURE.
_Experiment 204._--The plant was enclosed in a gla.s.s chamber and exposed to diffuse light. The experiment was commenced at midday, when the leaflets were open; the light was kept uniform while temperature was artificially increased by means of an electric heater placed in the chamber, and decreased by introducing cold air into the plant chamber.
One of the leaflets was attached to the recording lever and its movement, up or down, indicated the movement of opening or closure. The records showed that rise of temperature induces a movement of closure, while that of fall brings about the movement of opening.
[Ill.u.s.tration: FIG. 191.--Effect of sudden darkening at arrow, produces movement of closure (up-curve). Restoration of light induces opening movement (down-curve). Successive dots at intervals of 15 minutes.
(Leaflet of _Ca.s.sia_.)]
EFFECT OF VARIATION OF LIGHT.
_Experiment 205._--This experiment was also carried out at midday, when the leaflets were open. The horizontal record in figure 191 represents the stationary expanded condition of the leaflet; a black cloth was put over the gla.s.s chamber at 1 p.m., and the effect of darkness was recorded for one hour. Darkness is seen to initiate a movement of closure which increased at a rapid rate; the black cloth was removed after an hour, and the movement of opening under light was completed in the course of five quarters of an hour. It is thus seen that the leaflets are extremely sensitive to the action of light.
The experiments that have just been described on the effects of rise of temperature, and of light, show that they are antagonistic to each other. In the forenoon the opening movement under light has to be carried out against the closure movement due to rise of temperature.
Light, therefore, is the predominant factor in the diurnal movement of the leaflets of _Ca.s.sia_. The closure effect of darkness at night, on the other hand, overpowers the tendency of movement of opening due to fall of temperature.
[Ill.u.s.tration: FIG. 192.--Diurnal movement of the leaflet of _Ca.s.sia alata_. Closure movement commenced at 5 p.m. and completed by 9 p.m.
Leaflets began to open at 5 a.m.]
DIURNAL MOVEMENT OF THE LEAFLET OF _Ca.s.sia alata_.
_Experiment 206._--I next obtained the diurnal record of the leaflet, from 4 p.m. till 1 p.m. next day. The leaflets remain open from 1 p.m.
to 4 p.m. and the record of this period is therefore omitted. In the diurnal record (Fig. 192) the first thick dot was made at 4 p.m. and successive thick dots are at intervals of an hour, the thinner dots being at intervals of 15 minutes. It will be seen that a rapid movement of closure was initiated at 5 p.m. when the light is undergoing a rapid diminution. The movement of closure is completed at about 9 p.m. The leaflets remain closed till 5 a.m. next morning, after which they begin to open; this opening may commence even an hour earlier. It should be borne in mind in this connection, that since light and rise of temperature are antagonistic in their reactions, the effects of light and fall of temperature would be concordant; and the opening in the early hours may possibly be hastened by the low temperature in the morning. The leaflets open to their utmost by 9 a.m., and they remain open till the afternoon. The plant is so extremely sensitive to light that any slight fluctuation is followed by responsive movement of the leaflet. Thus the transitory pa.s.sage of a cloud is marked in the record by a short-lived closure movement.
DIURNAL MOVEMENT OF THE TERMINAL LEAFLET OF _Desmodium gyrans_.
Both the petiole, and the terminal leaflet of this plant exhibit very marked nyct.i.tropic movement. The petiole is raised and becomes almost erect in the evening, while the pulvinus of the terminal leaflet exhibits a sharp curvature downwards (Fig. 193).
[Ill.u.s.tration: FIG. 193.--The day and night positions of the petiole and terminal leaflet of _Desmodium gyrans_.]
_Experiment 207._--The petiole was held fixed, and the terminal leaflet attached to the recording lever. I have already explained that light falling on the pulvinus from above, induces an up-movement of the leaflet, which is thus erected under light of moderate intensity. If the light be strong, the transversely conducted excitation induces a partial neutralisation; very intense light may even cause a reversal into down-movement. Under natural conditions, day-light acting from above induces an up-movement; darkness, on the other hand, induces a rapid movement of fall. The leaflets sometimes exhibit autonomous pulsations; but the diurnal movement is very strong and the daily curve appears as a single large pulse on which smaller autonomous pulsations may become superposed.
[Ill.u.s.tration: 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 a.s.sumes 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 _Ca.s.sia_ 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 _Ca.s.sia_, 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 pa.s.sage 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.