Part 27 (1/2)

The following experiment will show that the position of tropic equilibrium is not fixed but subject to variation under changes of effective stimulation.

[Ill.u.s.tration: FIG. 185.--Effect of variation of intensity of light on phototropic equilibrium. Increase of intensity of light from L to L'

produces an increased positive curvature and a new state of balance.

Diminished intensity of light _l_ brings about a new balance at a lower level. The cessation of light (_l_ within a circle) restores the normal position of the organ.]

_Experiment 198._--I have explained how a maximum tropic curvature is induced under continued action of light. Employing the pulvinus of _Erythrina indica_ I applied light on the upper half of the pulvinus: (1) of medium intensity L, (2) of strong intensity L', and (3) of feeble intensity _l_. The source of light was an arc lamp; the intensity of light was varied by means of a focussing lens, which gave a parallel, a convergent or a divergent beam, with corresponding increase or diminution of intensity of light. Light was in each case continued till equilibrium was reached. Inspection of figure 185 shows that the position of equilibrium depends on the intensity of stimulation; the balance is 'raised' under increased and 'lowered' under decreased intensity.

In the case of geotropism the stimulus is constant, but its tropic effect, we shall presently see, undergoes variation with changing temperature.

EFFECT OF VARIATION OF TEMPERATURE ON GEOTROPIC TORSION.

_Modification of geotropic torsion: Experiment 199._--The _Mimosa_ plant was placed on its side, so that the pulvinus was subjected to lateral geotropic action. In response to this it underwent torsion, the upper half of the pulvinus tending to place itself so as to face the vertical lines of gravity. This torsional response was recorded as an up-movement; on the attainment of equilibrium the record became horizontal. The plant was now subjected to a cyclic variation of temperature, and the resulting variation of torsion recorded at the same time. The temperature of the plant chamber was gradually raised from the normal 30 to 34 C. and then allowed to return to the normal; finally the temperature was lowered to 26C. Rise of temperature was effected by means of an electrical heater placed inside the chamber with a vessel of water placed above it. Care has to be taken that the rise of temperature is gradual, since a sudden variation often acts as a stimulus. The water in the vessel not only keeps the chamber in a humid condition but also prevents sudden fluctuation of temperature. After the temperature had been raised to 34C., the heating current was stopped and the door of the plant chamber gradually opened, so as to allow the temperature to be restored to the normal. Cooled air was next introduced into the chamber till the temperature fell to 26C. Figure 186 exhibits clearly the effect of variation of temperature on geotropic torsion. The maximum torsion had been attained at 30C. and the first part of the record is therefore horizontal. Warmth was applied at H, and after a latent period of ten minutes, the geotropic torsion underwent a continuous diminution till a new state of equilibrium was reached at 34C. This took place shortly after the stoppage of the heating current at (H). On return to normal temperature the torsional balance was restored to its original position of equilibrium. Application of cold at C, is seen to bring about a new state of balance with an increase of geotropic torsion.

[Ill.u.s.tration: FIG. 186.--Effect of variation of temperature on geotropic torsion. Application of warmth at H diminishes the geotropic torsion; return to normal temperature (H) restores the original torsion; cooling at C, increases the geotropic torsion.]

The position of geotropic equilibrium is thus seen to be modified by variation of temperature, the tropic effect being diminished with the rise, and enhanced with the fall of temperature.

It may be thought that the phenomenon just described may not be different from ordinary thermonasty, exhibited by the perianth leaves of _Crocus_ and _Tulip_ in which a rise of temperature induces a movement of unfolding, and a fall of temperature brings about the opposite movement of closure. In these cases the movement is determined solely by the natural anisotropy of the organ, and not by the paratonic action of a directive external force. Thus the inner side of the perianth leaves undergoes an expansion with rise of temperature attended by the opening of the flower; this movement of opening does not undergo any change on holding the flower in an inverted position.

But the torsional movement of the leaf of _Mimosa_, and the induced variation of torsion under change of temperature are not solely determined by the natural anisotropy of the organ; it is, on the contrary, regulated by the directive action of the stimulus of gravity.

The pulvinus in normal position does not exhibit any geotropic torsion and in the absence of an antecedent torsion change of temperature cannot induce any variation in it. It is only after the pulvinus had become torsioned under the lateral action of geotropic stimulus that a responsive variation is induced in it by the action of changing temperature.

The change in torsion is, moreover, determined in reference to the paratonic action of incident geotropic stimulus. This will be clearly understood from the tabular statement given below.

TABLE XLVI.--SHOWING THE EFFECT OF RISE OF TEMPERATURE ON GEOTROPIC TORSION.

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Position of the organ.

Geotropic effect.

Effect of rise of

temperature.

+----------------------+---------------------+----------------------+

Right flank above:

Right-handed torsion.

Left-handed torsional

(_a_) position.

movement (untwist).

Left flank above:

Left-handed torsion.

Right-handed torsional

(_b_) position.

movement (untwist).

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