Life Movements in Plants Part 34

THE PHOTOMETRIC RECORDER.

This difficulty I have been able to overcome by the automatic device for continuous record of the variation of light. The electric resistance of a selenium cell undergoes diminution with the intensity of light that falls on it. The photo-sensitive cell was made the fourth arm of a Wheatstone bridge, the resistance of the cell being exactly balanced when the shutter of the sensitive cell was closed. The selenium receiver was pointed upwards against the sky. Precaution was taken that it was protected from the direct action of sunlight. On opening the shutter a deflection of the index of a sensitive galvanometer was produced, and the deflection increased with increasing intensity of diffuse skylight.

The special difficulty was in securing automatic record of the galvanometer deflections. This was obtained by a special contrivance of an oscillating smoked gla.s.s plate, the up and down oscillation being at intervals of 30 minutes. A detailed account of this apparatus will, with its possibilities for meteorology, be given in a future paper. I reproduce the record obtained in my greenhouse on the 5th March (1919), which gives a general idea of the variation of the light from morning to evening (Fig. 213). The record shows that the light began to be perceptible at 5-30 a.m., and that the intensity increased rapidly and continuously till it reached a climax at noon, after which it began to decline slowly. The decline of intensity of light was very abrupt after 5 p.m., the effect being reduced to zero at 6-30 p.m.

[Ill.u.s.tration: FIG. 213.--Photometric record showing variation of intensity of light from morning to evening. Successive dots are at intervals of 30 minutes.]

THE EFFECT OF DIRECT LIGHT.

Under natural conditions, the leaf of _Mimosa_ is acted on by light from above, and it is generally supposed that the pulvinus is positively phototropic, that is to say, it curves upwards till the leaf is placed at right angles to the direction of light. My investigations show, however, that the phototropic effects vary from positive to negative through an intermediate stage of neutralisation, these depending on the intensity and duration of exposure. When light acts continuously on the upper half of the pulvinus, there follows the following sequences of reaction:

(1) The leaf is at first erected by the contraction of the upper half of the pulvinus due to direct action of light acting from above.

(2) Under continuous stimulation of the upper half of the pulvinus by light, the excitation is slowly conducted to the lower half across the pulvinus. In consequence of this transmitted excitation, the lower half begins to contract and thus neutralises the first effect of erection.

The upper half of the pulvinus is less contractile than the lower half, and the neutralisation is due to the full contraction of the upper half antagonised by slight contraction of the lower half. The horizontal position of the leaf under light is therefore the result of balance of the two antagonistic reactions. If the incident light be very strong, the more intense transmitted excitation induces greater contraction of the lower half, and bring about a resultant down-movement (_cf._ p.

331).

Let us consider the effect of daily variation of light on _Mimosa_; we have here to take account both of intensity and duration. The intensity of light is seen to undergo a continuous increase which reaches a climax at noon; it then begins to decline slowly and the diminution of intensity of light is very abrupt after 5 p.m.

Under natural conditions the following phototropic effects are observed during the course of the day: light acting from above induces an up-movement of the leaf; but this is opposed by the thermo-geotropic fall of the leaf due to rise of temperature. As the two opposing effects are nearly balanced, any fluctuation of the relative intensity of the two gives rise to the pulsatory movements often seen in the forenoon; the _Mimosa_ leaf has moreover an autonomous movement of its own. Under continued action of light neutralisation begins to take place after 1 p.m. (_cf._ _Expt._ 135). Later in the day the phototropic effect may become negative; reversal into this negative takes place under the joint action of intensity and duration of light; it takes place earlier under strong, and later under feeble, light.

THE EVENING SPASMODIC FALL OF THE LEAF.

I shall now deal with the difficult problem of the sudden fall of the leaf after 5 p.m. Pfeffer regarded this sudden fall in the evening as due to the increased mechanical moment of the secondary petioles moving forward on the withdrawal of light. But the following experiment shows that the increased mechanical moment cannot be the true explanation of the fall.

[Ill.u.s.tration: 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.

The above explanations having proved unsatisfactory we have to search for other factors to account for the fall of the leaf on the cessation of light. In this connection I was struck by the extraordinary similarity of the diurnal curve of the petiole of _Ca.s.sia alata_ with that of _Mimosa_.

[Ill.u.s.tration: FIG. 215.--Diurnal record of _Ca.s.sia_ leaf. Note similarity with diurnal record of _Mimosa_.]

DIURNAL CURVE OF THE PETIOLE OF _Ca.s.sia alata_.

_Experiment 224_.--The leaf of _Ca.s.sia_ exhibits as in the leaf of _Mimosa_ a slight erectile movement after the thermal-noon at 2 p.m., there is next a sudden fall after 5 p.m., which continues about 9 p.m.; after this the leaf exhibits a continuous rise with the fall of temperature, till the climax is reached about 6 a.m. in the morning; the leaf then undergoes a fall with rise of temperature, there being a number of pulsatory movements in the forenoon, evidently due to unstable balance under the opposing effects of light and of rise of temperature (Fig. 215).

The reason of this similarity between the records of _Ca.s.sia_ and _Mimosa_ was found in the fact:

(1) That the main pulvinus of the leaf of _Ca.s.sia_ is, like the pulvinus of _Mimosa_, differentially excitable, the lower half being more excitable than the upper. This is demonstrated by sending a diffuse electric shock through the leaf, the response being by a fall of the leaf due to the greater contraction of the lower half of the pulvinus.

The leaf recovered after an interval of 20 minutes, the curve of response being similar to that of _Mimosa_. The only difference between the two organs is in the lesser excitability of the pulvinus of _Ca.s.sia_, on account of which a greater intensity of shock is necessary for producing the responsive fall.

(2) The responses to light are the same in both as will be seen in the following experiment.

[Ill.u.s.tration: FIG. 216.--Post-maximum after-effect of light on response of leaf of _Ca.s.sia_. There is an over-shooting on cessation of light at arrow within a circle.]

_Experiment 225._--In _Ca.s.sia_, as in _Mimosa_, light acting from above induces at first an erectile movement which reaches a maximum; after this there is a neutralisation and reversal. In the record given in figure 216, light from a small arc lamp acting on the upper half of the pulvinus for 48 minutes gave the maximum positive curvature; this was completely neutralised by further exposure to light for 20 minutes.

Light was cut off at neutralisation and there was a sudden fall beyond the equilibrium position, which was more rapid than the movement under light. The after-effect of prolonged exposure is thus an 'over-shooting'

beyond the normal position of equilibrium.

RESPONSE OF _Mimosa_ TO DARKNESS AT DIFFERENT PARTS OF THE DAY.

I now tried the effect of darkness on the movement of _Mimosa_, and was surprised to find that while artificial darkness caused a sudden fall of the leaf in the afternoon, it had no such effect in the forenoon.

_Experiment 226._--Successive records were taken of the effect of artificial darkness for two hours, alternating with exposure to light for two hours. The plant was subjected to darkness by placing a piece of black cloth over the gla.s.s cover from 12 to 2 p.m., it was exposed to light from 2 to 4 p.m. and darkened once more from 4 to 6 p.m.

The record given in figure 217 shows that the leaf had been moving upwards under the action of light (positive phototropism); darkness commenced at the point marked with a thick dot. _The after-effect on the stoppage of light is seen to be in the same direction as under light_; this persisted for ten minutes followed by recovery which was complete by 2 p.m., as seen in the horizontal character of the curve. On restoration of light (at the point marked with the second thick dot) the leaf moved upwards till the positive phototropic movement attained a maximum in the course of an hour and twenty minutes, after which neutralisation set in, and by 4 p.m. the positive phototropic effect had become partially neutralised. Artificial darkness at the third thick dot caused a rapid down-movement which overshot the position of equilibrium.

The difference of after-effect in the forenoon and in the afternoon lies in the fact that in the first case it was the pre-maximum after-effect; but in the second case the after-effect was post-maximum. I have already shown in the previous chapter that the pre-maximum after-effect of light is a short-lived movement in the same direction as under light, while post-maximum after-effect was a rapid over-shooting downwards beyond the equilibrium position. These characteristics are also found in the after-effects of light in _Mimosa_.

[Ill.u.s.tration: FIG. 217.--Effect of periodic alternation of light L, and of darkness D, on the response of _Mimosa_ leaf. The first darkness causes the pre-maximal after-effect of slight erection followed by recovery. The subsequent application of light from 2 to 4 p.m. caused erectile movement followed by partial neutralisation by 4 p.m. Stoppage of light at the third thick dot caused a sudden fall of leaf _below_ the position of equilibrium.]

The responses of _Mimosa_ on the cessation of light described above took place in the course of experiments which lasted for more than six hours.

Objection may be raised that during this long period the temperature variation must have produced certain effects on the response. In order to meet this difficulty, I carried out the following experiments which were completed in a relatively short time. I have already explained how the period of experiment could be shortened by suitable increase of the intensity of light. The experiment was commenced inside a room at noon and completed by 2 p.m.; the temperature variation during this period was less than 1C.

[Ill.u.s.tration:

FIG. 218.--Pre-maximum after-effect of light in _Mimosa_.

FIG. 219.--After-effect at maximum.

FIG. 220.--Post-maximum after-effect exhibiting an 'over-shooting'

below position of equilibrium.

In the above records light was applied at arrow, and stopped at the second arrow enclosed in a circle.]

_After-effect at pre-maximum: Experiment 227._--Light from an 100 c.p.

incandescent lamp was focussed on the upper half of the pulvinus of _Mimosa_ for 8 minutes, after which the light was turned off. The after-effect was a persistence of previous movement followed by recovery (Fig. 218).

_After-effect at maximum: Experiment 228._--Continued action of light for 18 minutes induced maximum positive curvature as seen in the upper part of the curve becoming horizontal. On the stoppage of light, there was a recovery to the original position of equilibrium (Fig. 219).