I have hitherto spoken of the excitatory effect of the upper layer; there must be some physiological reaction on the lower perceptive layer, though of a different character, represented diagrammatically by vertical shading. Had the physiological reaction on the lower side of a radial organ been the same as on the upper, geotropic curvature would have been an impossibility, for similar reactions on opposite sides would, by their antagonistic effects, have neutralised each other.

After this preliminary explanation, I shall give a detailed account of the experiments and results. It is to be borne in mind that the investigation I am going to describe presupposes no hypothesis of geotropic action. I start with the observed fact that an organ under the stimulus of gravity, exhibits responsive movement. I ascertain the nature of the underlying reaction by electric tests; I have, in my previous works, fully demonstrated that the excitatory contractile reaction is detected by electro-motive change of galvanometric negativity, and the opposite expansive reaction by a change of galvanometric positivity. With the electric probe I ascertain whether geotropic irritation is diffuse, or whether it is localised at any particular depth of the organ. I map out the contour lines of physiological reaction with its heights and depths of excitation.

I shall now proceed to describe the results of electric exploration into the interior of the organ. The trouble I foresaw, related to the irritation caused by the passage of the probe, and the after-effect of wound on variation of excitability.

Fig. 175.—The Electric Probe. Figure to the left represents one electric contact made with sepal of Nymphæa, and the other, with the flower-stalk by means of the probe; the included galvanometer is represented by a circle. Figure to the right an enlarged view of the probe.

THE ELECTRIC PROBE.

The wound-irritation is, however, reduced to a minimum by making the probe exceedingly thin. A fine platinum wire 0·06 mm. in diameter passes through a glass tubing drawn out into a fine capillary, and fused round one end of the platinum wire which protrudes very slightly beyond the point of fusion; the exploring electrode is thus insulated except at the protruded sharp point of the platinum wire. The length of the capillary is about 6 mm., just long enough to pass the experimental plant-organ transversely from one end to the other; the average diameter of the capillary is about 0·15 mm. The other end of the platinum wire comes out of the side of the tubing and is led to one terminal of the galvanometer, the other being connected with an indifferent point in the organ. The probe can be gradually pushed into the plant-organ by rotation of a screw head, one complete rotation causing a forward movement through 0·2 mm. (Fig. 175).

Wound-reaction.—I have shown that a prick acts as a mechanical stimulus, and in normal excitable tissues induces an excitatory change of galvanometric negativity. This wound-reaction increases with the extent of the wound, and the suddenness with which it is inflicted. On account of the fineness of the probe, it insinuates itself into the tissue rather than make any marked rupture; the probe again is introduced very gradually; with these precautions the wound-reaction is found to be greatly reduced. The immediate effect of the prick is a negative deflection of the galvanometer, which declines and attains a steady value in the course of about 5 minutes.

Effect of wound on excitability.—I have shewn (p. 81) that severe wound caused by transverse section induced a temporary abolition of irritability in Mimosa, but that the normal excitability was restored in the course of an hour. A prick from a thick pin was shown to depress temporarily the rate of growth, the normal rate being restored after an interval of 15 minutes (p. 202). In the case of geo-electric excitability, the depressing effect of the passage of the probe, I find, to disappear in the course of about 10 minutes.

For a choice of experimental material we have to find specimens which are not merely geotropically sensitive, but also exhibit large electric response under stimulus. In both these respects the shoot of Bryophyllum and the flower stalk of Nymphæa give good results.