Part 7 (1/2)

COMPLEXITY OF PROBLEM OF PHOTOTROPIC REACTION.

The complexity of phototropic reaction arises from the summated effects of numerous factors; for explanation of the resultant response it is therefore necessary to take full account of the individual effect of each of them.

Among these operative factors in phototropic reaction may be mentioned:--

(1) The difference of effects induced by light at the proximal and distal sides of the organ.

(2) The modification of the latent period with the intensity of stimulus.

(3) The after-effect of stimulus.

(4) The modifying influence of tonic condition on response.

(5) The effect of direction of light.

(6) The effect of intensity of light.

(7) The effect of duration of stimulation.

(8) The transmitted effect of light.

(9) The effect of unequal excitability in different zones of the organ.

(10) The effect of transverse conduction in modification of the sign of response.

(11) The effect of temperature on phototropic action.

(12) The modification of response due to differential excitability of the organ.

(13) Nastic and tropic reactions.

(14) The torsional effect of light.

The sketch given above will give us some idea of the complexity of the problem. In this and in the following papers I shall describe the investigations I have carried out on the subjects detailed above.

ACTION OF LIGHT.

I have shown that there is no essential difference between the responses of pulvinated and growing organs, that diminution of turgor induced by stimulus brings about contraction in the one, and r.e.t.a.r.dation of the rate of growth in the other. Indirect stimulation, on the other hand, induces an expansion and acceleration of the rate of growth. The experimental investigation on the tropic effect of light may therefore be carried out both with pulvinated and growing organs.

As regards the effect of direct stimulus of light on growing organs we found (p. 208) that it induces an incipient contraction, seen in diminution of the rate of growth; this incipient contraction culminates in an actual contraction under increasing intensity of light. The contraction under direct stimulation is also observed in pulvinated organs. When light acts from above the upper half of the pulvinus undergoes contraction, resulting in erection of the motile leaf or leaflets. As regards the effect of indirect unilateral stimulus of light on the distal side of the organ, we found that its effect is an enhancement of turgor (p. 281). Hence the positive tropic curvature under light is brought about, as in the case of other forms of stimuli, by the contraction of the proximal, and expansion of the distal sides of the organ.

Various a.n.a.logies have been noticed between phototropic and geotropic reactions, and it has been supposed that the two phenomena are closely related to each other. This has even led to a.s.sumption that there are phototropic particles which function like statoliths in geotropic organs. There is, however, certain outstanding difference between the two cla.s.ses of phenomena. In the case of light, the incident energy is entirely derived from the outside. But in geotropism, the force of gravity by itself is ineffective without the intervention of the weight of cell-contents to exert pressure on the sensitive ectoplasm, and thus induce stimulation. This aspect of the subject will be treated in greater detail in a subsequent chapter.

POSITIVE PHOTOTROPIC CURVATURE.

I shall now describe the phototropic effect of unilateral light in pulvinated, and in growing organs. From the explanation that has already been given, it will be understood that the side of the organ directly acted on by light undergoes contraction and concavity.

_Tropic curvature of pulvinated organs: Experiment 117._--For this experiment I employed the terminal leaflet of the bean plant. The source of illumination was 32 c.p. electric lamp, enclosed in a metallic tube with circular aperture for pa.s.sage of light. The leaflet was attached to an Oscillating Recorder. Light was applied on the upper half of the pulvinus for 20 seconds; this induced an up-movement of the leaflet, due to the contraction of the upper half of the organ. Recovery took place in course of 8 minutes (Fig. 112).