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.

Position of the organ. Geotropic effect.Effect of rise of temperature.
Right flank above:Right-handed torsion.Left-handed torsional movement (untwist).
(a) position.
Left flank above:Left-handed torsion.Right-handed torsional movement (untwist).
(b) position.

By right flank in the above table is meant the side of the pulvinus to the right of the observer facing the leaf of the plant held in the normal position. When the plant is laid on its left side in the a-position, the right flank will be above and the responsive torsion under geotropic stimulus becomes right handed or with the hands of a clock (Cf. Fig. 179). When the plant is laid on its right side, the left flank will be above and the geotropic torsion becomes left handed or against the hands of the clock.

It will be seen from the above that in whatever way the experimental condition may be varied, the movement in response to variation of temperature is determined in relation to the antecedent geotropic torsion. The geotropic effect whether left-handed or right-handed torsion is always diminished by the rise of temperature, and enhanced by the fall of temperature.

VARIATION OF APO-GEOTROPIC CURVATURE UNDER THERMAL CHANGE.

I shall now proceed to show that variation of temperature not merely induces variation of geotropic torsion but also of geotropic curvature. I shall first demonstrate the effect of thermal change on geotropic curvature of the shoot, and then demonstrate its effect on dia-geotropic curvature of leaves.

Experiment 200.—A specimen of Tropæolum majus grown in a small flower pot, is laid on its side. Under geotropic action the shoot becomes curved, the upper side becoming concave and the lower side convex. The end of the stem is attached to the recording apparatus; when the plant is subjected to a rise of temperature, the movement induced shows that the geotropic effect has undergone a diminution, the curvature exhibiting a flattening; lowering of temperature, on the other hand, increases the geotropic curvature. Other instances of this will be found in a subsequent chapter. The diurnal movement of the 'Praying Palm' is a striking example of the effect of variation of temperature in modification of geotropic curvature (p. 30). Rise of temperature is thus shown to diminish geotropic torsion of dorsiventral organs, and the apo-geotropic curvature of radial organs. We have next to study the effect of temperature variation on the dia-geotropic equilibrium of leaves.