Here then we have the inexplicable phenomenon of a particular tissue, itself incapable of response, yet arresting the movement in a neighbouring tissue.
The problem before us may be thus stated: Is the movement of the tendril due to certain specific sensibility of the organ, on account of which its reactions are characteristically different from other tropic movements? Or, does the twining of tendril come under the law of tropic curvature that has been established, namely that it is brought about by the contraction of the directly stimulated proximal side, and the expansion of the indirectly stimulated distal side?
I shall now describe my investigations on the effects of direct and indirect stimulus on the growth of tendril; I have in this investigation studied the effect not merely of mechanical, but also of other forms of stimuli. I shall also describe the diverse effects induced by mechanical stimulus under different conditions. From the results of these experiments I shall be able to show that the twining of the tendril comes under the general law of tropic curvature; that the curvature results from the contraction of the proximal and expansion of the distal side. Finally I shall be able to offer a satisfactory explanation of the inhibition of response of the tendril by the stimulation of the opposite side of the organ.
GENERAL EFFECTS OF INDIRECT AND DIRECT ELECTRIC STIMULATION ON THE GROWTH OF TENDRIL.
For this experiment I took a growing tendril of _Cucurbita_ in which the sensitiveness is more or less uniform on all sides. The tendril was suitably mounted on the Balanced Crescograph, which records the variation of the rate of growth induced by immediate and after-effect of stimulus. The specimen is held in a clamp as in the diagram (Fig. 102), the tip being suitably attached to the recording lever. For indirect stimulation feeble shock from an induction coil is applied at the two electric connections below the clamp. Direct stimulus is applied by means of electric connections one above and the other below the clamp.
[Ill.u.s.tration: FIG. 102.--Diagrammatic representation of indirect and direct stimulation of tendril.]
_Effect of Indirect Stimulus: Experiment 106._--The growth of the tendril was exactly balanced, and the record became horizontal. Indirect stimulus was next applied below the clamp; this is seen to upset the balance, with the resulting up-curve which indicates a sudden acceleration of growth above the normal. This acceleration took place within ten seconds of the application of stimulus, and persisted for three minutes; after this the normal rate of growth became restored, as seen by the balanced record once more becoming horizontal (Fig. 103).
[Ill.u.s.tration: FIG. 103.--Record by Method of Balance, showing acceleration of growth of tendril (up-curve) induced by indirect stimulation. (_Cucurbita._)]
_Effect of Direct Stimulus: Experiment 107._--The incipient contraction induced by direct stimulation is so great that the record obtained by the delicate method of balance cannot be kept within the plate. I, therefore, took the ordinary growth-curve on a moving plate. The first part of the curve represents normal growth; stimulus of feeble electric shock was applied at the highest point of the curve. This is seen (Fig.
104) to induce an immediate contraction and reversal of the curve which persisted for two and half minutes, after which growth was slowly renewed. The most interesting fact regarding the after-effect of stimulus is that the rate of growth became actually enhanced to three times the normal. This is clearly seen in the record (upper half of the figure) taken 20 minutes after stimulation, where the curve is far more erect than that of the normal rate of growth before stimulation.
[Ill.u.s.tration: FIG. 104.--Variation of growth induced by direct stimulation. First part of the curve shows normal rate of growth. Direct stimulation induces contraction (reversal of curve). After-effect of stimulus seen in highly erect curve in upper part of record, taken 20 minutes after.]
The effects of Indirect and Direct stimulation of the tendril are summarised below:
(1) Indirect stimulation induces a sudden enhancement of rate of growth, followed by a recovery of the normal rate.
(2) Direct stimulation induces a r.e.t.a.r.dation of the rate of growth which may culminate into an actual contraction. _The after-effect of direct stimulus of moderate intensity is an enhancement of the rate of growth._
The experiments described above demonstrate the effects of direct and indirect electrical stimulus. I shall now proceed to show that mechanical stimulus induces effects which are similar to those of electric stimulus.
EFFECTS OF DIRECT AND INDIRECT MECHANICAL STIMULUS.
_Effect of Direct mechanical stimulus: Experiment 108._--In this case I took a tendril of _Cucurbita_, and attached it to the ordinary High Magnification Crescograph, the record of which gives the absolute rate of its normal growth, and the induced variation of that rate. The tendril was stimulated mechanically by simultaneous friction of its different sides. The immediate effect was a r.e.t.a.r.dation of growth, the reduced rate being less than half the normal. There was a recovery on the cessation of the stimulus; the rate of growth was even slightly enhanced after an interval of 15 minutes. Table XXVI shows the immediate and after-effects of mechanical stimulation on growth.
TABLE XXVI.--SHOWING THE IMMEDIATE AND AFTER-EFFECT OF MECHANICAL STIMULATION ON TENDRIL (_Cucurbita_).
+--------------------------------------------------------------+Normal rate of growth 044 per sec.r.e.t.a.r.ded rate immediately after stimulation 020 " "Recovery and enhancement after 15 minutes 050 " "+--------------------------------------------------------------+
The immediate and after-effects of mechanical stimulus on the tendril are therefore the same as that of electric stimulus. The incipient contraction under direct mechanical stimulus, moreover, is not the special characteristic of tendrils, but of growing plants in general.
For I have shown (page 203) that the growth of flower stalk of _Zephyranthes_ is also r.e.t.a.r.ded after mechanical friction, from the normal rate 048 to 011 after stimulation. We shall find later that different plant organs, after moderate stimulation, exhibit acceleration of growth as an after-effect. The phenomenon of responsive reaction of tendril is therefore not unique, but similar to that of other organs under all forms of stimulation. The only speciality in tendril is that owing to anatomical peculiarities, the perceptive power of the organ for mechanical stimulus is highly developed.
We are now in a position to offer an explanation of the induced concavity of the stimulated side of the tendril, and its recovery after brief contact. The experiments that have been described show that:
(1) the proximal side contracts because it is directly stimulated, and the distal side, being indirectly stimulated, expands; the curvature is thus due to the joint effects of contraction of one side, and expansion of the opposite side, and
(2) the recovery of the tendril after brief contact is hastened by the after-effect of stimulus, which is expansion and acceleration of growth.
The results given above will also be found to explain Fitting"s important observations[5] that (_a_) the stimulated side of the tendril undergoes transient contraction with subsequent acceleration of growth, and that (_b_) the distal or convex side undergoes an immediate enhancement of growth.
[5] Pfeffer--_Ibid_--Vol. III, p. 57.
I give below a record given by a tendril of _Cucurbita_ in response to unilateral contact of short duration (Fig. 105). Successive dots in the record are at intervals of three seconds. The latent period was ten seconds, and the maximum curvature was attained in the course of two and a half minutes. The curvature persisted for a further period of two minutes after which recovery was completed in the course of 12 minutes.
Feeble stimulation is attended by a recovery within a short period, but under strong stimulus the induced curvature becomes more persistent.
[Ill.u.s.tration: FIG. 105.--Positive curvature of tendril of Cucurbita under unilateral stimulus of contact at x.]
INHIBITORY ACTION OF STIMULUS.
I have referred to the remarkable observation of Fitting that though the application of stimulus on the upper side of the tendril of _Pa.s.siflora_ did not induce any response, yet it inhibited the normal response of the under side.
The results of experiments which I have described will, however, afford a satisfactory explanation of this curious inhibition. It has been explained, that the curvature of the tendril is due to the joint effects of diminished turgor and contraction at the directly stimulated side, and an enhancement of turgor and expansion on the opposite side. In the diagram seen in figure 106, the left is the more excitable side, and contraction will induce concavity of the stimulated side. But if the opposite or less excitable side of the tendril be stimulated at the same time, then the transmitted effect of indirect stimulus will induce enhancement of turgor and expansion on the left side, and thus neutralise the previous effect of direct stimulus. An inhibition of response will thus result from the stimulation of the opposite side.
[Ill.u.s.tration: FIG. 106.--Diagrammatic representation of effects of Indirect and Direct unilateral stimulation of the tendril. Indirect stimulation, I, induces movement away from stimulated side (negative curvature) represented by continuous arrow. Direct stimulation, D, induces movement towards stimulus (positive curvature) indicated by dotted arrow.]
A difficulty arises here from the fact that the upper side of the tendril (the right side in Fig. 106) is supposed to be inexcitable and non-contractile. Hence there may be a misgiving that the stimulation of the non-motile side may not induce the effect of indirect stimulus (an increase of turgor and expansion) at the opposite side, which is to inhibit the response. But I have shown that even a non-contractile organ under stimulus generates both the impulses, positive and negative. This is seen ill.u.s.trated in figure 100, where the rigid stem of _Mimosa_ was subjected to unilateral stimulation; the effect of indirect stimulus was found to induce an enhancement of turgor at the diametrically opposite side, and thus caused an erectile movement of the motile leaf. Electric investigations which I have carried out also corroborate the results given above. Here also stimulation of a non-motile organ at any point, induces at a diametrically opposite point, a positive electric variation indicative of enhanced turgor. It will thus be seen that inhibition is possible even in the absence of contraction of the upper side of the tendril; hence the contraction of the directly stimulated side is neutralised by the effect of indirect stimulation of the distal side.
RESPONSE OF LESS EXCITABLE SIDE OF THE TENDRIL.
It is generally supposed that the upper side of the tendril of _Pa.s.siflora_ is devoid of contractility. This is however not the case, for my experiments show that stimulation of the upper side also induces contraction and concavity of that side, though the actual movement is relatively feeble.
_Experiment 109._--In order to subject the question to quant.i.tative test I applied feeble stimulus of the same intensity on upper and lower side alternately. Successive stimuli were kept more or less uniform by employing the following device. I took a flat strip of wood 1 cm. in breadth, and coated 2 cm. of its length with sh.e.l.lac varnish mixed with fine emery powder. On drying the surface became rough, the flat surface was gently pressed against the area of the tendril to be stimulated, and quickly drawn so that the rough surface 2 cm.1 cm. was rubbed against the tendril in each experiment. Stimulation, thus produced, induced a responsive movement of each side of the organ. The extent of the maximum movement was measured by the microscope micrometer. The following results were obtained with four different specimens.
TABLE XXVII.--SHOWING THE RELATIVE INTENSITIES OF RESPONSES OF THE UPPER AND UNDER SIDE OF TENDRIL (_Pa.s.siflora_).
+------------------------------------------------------------+Movement induced byMovement induced byBstimulation of understimulation of upperRatio ---.side, A.side, B.A+----------------------+-------------------------------------+(1) 85 divisions14 divisions1/6(2) 106 "15 "1/7(3) 60 "8 "1/7(4) 80 "10 "1/8+------------------------------------------------------------+
It will thus be seen that the upper side of the tendril is not totally inexcitable, its power of contraction being about one-seventh that of the under side.
NEGATIVE CURVATURE OF THE TENDRIL.
I shall now describe certain remarkable results which show that under certain definite conditions the tendril moves away from the stimulated side. I have explained, how in growing organs the effect of unilateral stimulus longitudinally transmitted, induces an expansion higher up on the same side to which the stimulus is applied, resulting in convexity and movement away from the stimulus (cf. Laws of Tropic Curvatures, p.
286). As the reaction of tendril is in no way different from that of growing organs in general, it occurred to me that it would be possible to induce in it a negative curvature by application of indirect unilateral stimulus.
_Experiment 110._--A tendril of _Pa.s.siflora_ was held in a clamp, as in the diagram (Fig. 106) in which the left is the more excitable side of the organ. The responsive movement of the tendril is observed by focussing a reading microscope on a mark on the upper part of the tendril. Direct mechanical stimulation at the dotted arrow makes the tendril move in the same direction, the response being _positive_. But if stimulus be applied on the same side below the clamp the tendril is found to move away from stimulus, the response being now _negative_.
This reversal of response, as previously stated, is due to the fact that the transmitted effect of indirect stimulus induces an acceleration of growth higher up on the same side, which now becomes convex. The result though unexpected, is in every way parallel to the response of the flower bud of _Crinum_, in which the normal positive response was converted into negative by changing the point of application of stimulus, so that it became indirect (p. 216).
SUMMARY.
The response of tendril is in no way different from that of growing organs in general.
Direct stimulus, electrical or mechanical, induces an incipient contraction; the after-effect of a feeble stimulus is an acceleration of growth above the normal. Indirect stimulus induces an enhancement of the rate of growth.
Under unilateral mechanical stimulus of short duration the directly excited proximal side undergoes contraction, the indirectly stimulated distal side exhibits the opposite effect of expansion. The induced curvature is thus due to the joint effects of the contraction of one side, and the expansion of the opposite side.