Response in the Living and Non-Living
Chapter 36
INORGANIC RESPONSE--EFFECT OF CHEMICAL REAGENT
Action of chemical reagents--Action of stimulants on metals--Action of depressants on metals--Effect of 'poisons' on metals--Opposite effect of large and small doses.
We have seen that the ultimate criterion of the physiological character of electric response is held to be its abolition when the substance is subjected to those chemical reagents which act as poisons.
#Action of chemical reagents.#--Of these reagents, some are universal in their action, amongst which strong solutions of acids and alkalis, and salts like mercuric chloride, may be cited. These act as powerful toxic agents, killing the living tissue, and causing electric response to disappear. (See fig. 88.) It must, however, be remembered that there are again specific poisons which may affect one kind of tissue and not others. Poisons in general may be regarded as extreme cases of depressants. As an example of those which produce moderate physiological depression, potassium bromide may be mentioned, and this also diminishes electric response. There are other chemical reagents, on the other hand, which produce the opposite effect of increasing the excitability and causing a corresponding exaltation of electric response.
We shall now proceed to inquire whether the response of inorganic bodies is affected by chemical reagents, so that their excitability is exalted by some, and depressed or abolished by others. Should it prove to be so, the last test will have been fulfilled, and that parallelism which has been already demonstrated throughout a wide range of phenomena, between the electric response of animal tissues on the one hand, and that of plants and metals on the other, will be completely established.
#Action of stimulants on metals.#--We shall first study the stimulating action of various chemical reagents. The method of procedure is to take a series of normal responses to uniform stimuli, the electrolyte being water. The chemical reagent whose effect is to be observed is now added in small quantity to the water in the cell, and a second series of responses taken, using the same stimulus as before. Generally speaking, the influence of the reagent is manifested in a short period, but there may be occasional instances where the effect takes some time to develop fully. We must remember that by the introduction of the chemical reagent some change may be produced in the internal resistance of the cell. The effect of this on the deflection is eliminated by interposing a very high external resistance (from one to five megohms) in comparison with which the internal resistance of the cell is negligible. The fact that the introduction of the reagent did not produce any variation in the total resistance of the circuit was demonstrated by taking two deflections, due to a definite fraction of a volt, before and after the introduction of the reagent. These deflections were found equal.
I first give a record of the stimulating action of sodium carbonate on tin, which will become evident by a comparison of the responses before and after the introduction of Na_2CO_3 (fig. 89). The next record shows the effect of the same reagent on platinum (fig. 90).
#Action of depressants.#--Certain other reagents, again, produce an opposite effect. That is to say, they diminish the intensity of response. The record given on the next page (fig. 91) shows the depressing action of 10 per cent. solution of KBr on tin.
#Effect of 'poison.'#--Living tissues are killed, and their electric responses are at the same time abolished by the action of poisons. It is very curious that various chemical reagents are similarly effective in killing the response of metals. I give below a record (fig. 92) to show how oxalic acid abolishes the response. The depressive effect of this reagent is so great that a strength of one part in 10,000 is often sufficient to produce complete abolition. Another notable point with reference to the action of this reagent is the persistence of after-effect. This will be clearly seen from an account of the following experiment. The two wires A and B, in the cell filled with water, were found to give equal responses. The wires were now lifted off, and one wire B was touched with dilute oxalic acid. All traces of acid were next removed by rubbing the wire with cloth under a stream of water. On replacing the wire in the cell, A gave the usual response, whereas that of B was found to be abolished. The depression produced is so great and passes in so deep that I have often failed to revive the response, even after rubbing the wire with emery paper, by which the molecular layer on the surface must have been removed.
We have seen in the molecular model (fig. 62, _d_, _e_) how the attainment of maximum is delayed, the response diminished, and the recovery prolonged or arrested by increase of friction or reduction of molecular mobility.
It would appear as if the reagents which act as poisons produced some kind of molecular arrest. The following records seen to lend support to this view. If the oxalic acid is applied in large quantities, the abolition of response is complete. But on carefully adding just the proper amount I find that the first stimulus evokes a responsive electric twitch, which is less than the normal, and the period of recovery is very much prolonged from the normal one minute before, to five minutes after, the application of the reagent (fig. 93, _a_). In another record the arrest is more pronounced, i.e. there is now no recovery (fig. 93, _b_). Note also that the maximum is attained much later. Stimuli applied after the arrest produce no effect, as if the molecular mechanism became, as it were, clogged or locked up.
In connection with this it is interesting to note that the effect of veratrine poison on muscle is somewhat similar. This reagent not only diminishes the excitability, but causes a very great prolongation of the period of recovery.
In connection with the action of chemical reagents the following points are noteworthy.
(1) The effect of these reagents is not only to increase or diminish the height of the response-curve, but also to modify the time relations. By the action of some the latent period is diminished, others produce a prolongation of the period of recovery. Some curious effects produced by the change of time relations have been noticed in the account given of diphasic variation (see p. 113).
(2) The effect produced by a chemical reagent depends to some extent on the previous condition of the wire.
(3) A certain time is required for the full development of the effect. With some reagents the full effect takes place almost instantaneously, while with others the effect takes place slowly. Again the effect may with time reach a maximum, after which there may be a slight decline.
(4) The after-effects of the reagents may be transitory or persistent; that is to say, in some cases the removal of the reagent causes the responses to revert to the normal, while in others the effect persists even after the removal of all traces of the reagent.
#Opposite effects of large and small doses.#--There remains a very curious phenomenon, known not only to students of physiological response but also known in medical practice, namely that of the opposite effects produced by the same reagent when given in large or in small doses. Here, too, we have the same phenomena reproduced in an extraordinary manner in inorganic response. The same reagent which becomes a 'poison' in large quantities may act as a stimulant when applied in small doses. This is seen in record fig. 94, in which (_a_) gives the normal responses in water; KHO solution was now added so as to make the strength three parts in 1,000, and (_b_) shows the consequent enhancement of response. A further quantity of KHO was added so as to increase the strength to three parts in 100. This caused a complete abolition (_c_) of response.
It will thus be seen that as in the case of animal tissues and of plants, so also in metals, the electrical responses are exalted by the action of stimulants, lowered by depressants, and completely abolished by certain other reagents. The parallelism will thus be found complete in every detail between the phenomena of response in the organic and the inorganic.