Effect of anæsthetics and poisons.—Ordinary anæsthetics, like chloroform, and poisons, like mercuric chloride, are known to produce a profound depression or abolish all signs of response in the living tissue. For the purpose of experiment, I took two groups of stalks, with leaves attached, exactly similar to each other in every respect. In order that the leaf-stalks might absorb chloroform I dipped their cut ends in chloroform-water, a certain amount of which they absorbed, the process being helped by the transpiration from the leaves. The second group of stalks was placed simply in water, in order to serve for control experiment. The narcotic action of chloroform, finally culminating in death, soon became visually evident. The leaves began to droop, a peculiar death-discolouration began to spread from the mid rib along the venation of the leaves. Another peculiarity was also observed. The aphides feeding on the leaves died even before the appearance of the discoloured patches, whereas on the leaves of the stalks placed in water these little creatures maintained their accustomed activity, nor did any discolouration occur. In order to study the effect of poison, another set was placed in water containing a small quantity of mercuric chloride. The leaves here underwent the same change of appearance, and the aphides met with the same untimely fate, as in the case of those subjected to the action of chloroform. There was hardly any visible change in the appearance of the stalks themselves, which were to all outer seeming as living as ever, indications of death being apparent only on the leaf surfaces. I give below the results of several sets of experiments, from which it would appear that whereas there was strong normal response in the group of stalks kept in water, there was practically a total abolition of all response in those anæsthetised or poisoned.

Experiments on the effect of anæsthetics and poisons. A batch of ten leaf-stalks of plane-tree was placed with the cut ends in water, and leaves in air; an equal number was immersed in chloroform-water; a third batch was placed in 5 per cent. solution of mercuric chloride.

Similarly a batch of three horse-chestnut leaf-stalks was put in water, another batch in chloroform-water, and a third batch in mercuric chloride solution.

I. Leaf-stalk of Plane-tree


The stimulus applied was a single vibration of 90°.
A. After 24 hours in
water
B. After 24 hours in
chloroform water
C. After 24 hours in
mercuric chloride
[All leaves standing up
and fresh—aphides alive]
[Leaves began to droop
in 1 hour and bent over
in 3 hours—aphides dead]
[Leaves began to droop
in 4 hours. Deep discolouration
along the veins. Aphides dead]
Electric responseElectric responseElectric response
(1)21dns.(1)1dn.(1)0dn.
(2)31" (2)1" (2)0·25"
(3)26" (3)2" (3)0·25"
(4)15" (4)0" (4)0"
(5)17" (5)1" (5)0·25"
(6)23" (6)1·5" (6)0·25"
(7)30" (7)2" (7)0"
(8)27" (8)1" (8)0·25"
(9)29" (9)1" (9)0·25"
(10)17" (10)0·5" (10)0·5"
Mean response 23·6Mean 1Mean 0·15

II. Leaf-stalk of Horse-chestnut

(1)15dns.(1)0·5dn.(1)0dn.
(2)17" (2)0·5" (2)0"
(3)10" (3)0" (3)0"
Mean 14Mean 0·3Mean 0

These results conclusively prove the physiological nature of the response.

I shall in a succeeding chapter give a continuous series of response-curves showing how, owing to progressive death from the action of poison, the responses undergo steady diminution till they are completely abolished.

Effect of high temperature.—It is well known that plants are killed when subjected to high temperatures. I took a stalk, and, using the block method, with torsional vibration as the stimulus, obtained strong responses at both ends A and B. I then immersed the same stalk for a short time in hot water at about 65° C., and again stimulated it as before. But at neither A nor B could any response now be evoked. As all the external conditions were the same in the first and second parts of this experiment, the only difference being that in one the stalk was alive and in the other killed, we have here further and conclusive proof of the physiological character of electric response in plants.

The same facts may be demonstrated in a still more striking manner by first obtaining two similar but opposite responses in a fresh stalk, at A and B, and then killing one half, say B, by immersing only that half of the stalk in hot water. The stalk is replaced in the apparatus, and it is now found that whereas the A half gives strong response, the end B gives none.

In the experiments on negative variation, it was tacitly assumed that the variation is due to a differential action, stimulus producing a greater excitation at the uninjured than at the injured end. The block method enables us to test the correctness of this assumption. The B end of the stalk is injured or killed by a few drops of strong potash, the other end being uninjured. There is a clamp between A and B. The end A is stimulated and a strong response is obtained. The end B is now stimulated, and there is little or no response. The block is now removed and the plant stimulated throughout its length. Though the stimulus now acts on both ends, yet, owing to the irresponsive condition of B, there is a resultant response, which from its direction is found to be due to the responsive action of A. This would not have been the case if the end B had been uninjured. We have thus experimentally verified the assumption that in the same tissue an uninjured portion will be thrown into a greater excitatory state than an injured, by the action of the same stimulus.