Irritability: A Physiological Analysis of the General Effect of Stimuli in Living Substance

by Max Verworn

Fig. 49.

Fig. 50.

It is from the beginning apparent that the response to the second stimulus is determined by the intensity of the second stimulus in relation to the degree of irritability which exists at the moment when this is effective. This relation is dependent first upon the absolute intensity of the second stimulus. In the following diagram the intensity of the existing threshold value is fixed for convenience as ordinates beneath the abscissa. If, for example, at the time point x, a stimulus of weak intensity R₁ acts, this stimulus being under the existing threshold, produces no perceptible effect. (Figure [48].) If now instead of a weak stimulus, one of stronger intensity acts at the time point x, this stimulus will produce an appreciable response. (Figure [49].) If the second stimulus is of the same strength as the first, this second stimulus will bring about relatively less disintegration, because the system is then in a state in which irritability is still reduced. But this lessened disintegration in that it summates the excitation still existing as the result of the first stimulus can produce an absolute increase of the height above that of the abscissa. Here then we see the possibility of an increase of response resulting from summation. Accordingly the increase of disintegration must occur simultaneously with a diminution of irritability and this must fall below the level of the reduction of irritability produced by the first stimulus. This augmentation of the response through summation above the level of that produced by the first stimulus acting upon an unexcitated system is, however, connected with another condition. The above example refers to systems in which weak stimuli bring about weak response and strong stimuli strong response, that is, the response is capable of increase. In systems in which the “all or none law” is applicable, such an alteration in the absolute height of excitation, as results in summation, is not possible. In order to characterize these two types of living systems by a short expression rather than by a long sentence, we will call the first a “heterobolic system,” the latter in which the “all or none law” is operative an “isobolic system.” The former term expresses various degrees of discharge depending upon the intensity of the stimulus, the latter term refers to the constancy of discharge following stimuli of various intensities. Isobolic systems are in contradistinction to the heterobolic systems not capable of summation. The response to the second stimulus of equal intensity cannot be greater than that of the first, it may be equal to the first (Figure [50]) or be less in extent, but it can never be greater than that resulting when a single stimulus is applied. These facts have been known for a long time in the case of the heart muscle. A word is necessary, however, concerning the effect of stimuli beneath the threshold in heterobolic systems. We must here distinguish between the “ideal” threshold, beneath which the influence of a stimulus is nil, and the threshold of perceptible effect, beneath which a stimulus apparently has no effect; nevertheless a weak effect does occur, as is shown by succeeding reactions. This effect is manifested by a sub-threshold disintegration and a corresponding slight reduction of irritability. (Figure [51].) The presence of such a sub-threshold effect is recognized by various facts as, for example, the summation of the sub-threshold stimuli to production of a perceptible result. Thus stimulation of a sensory spinal cord root with a single sub-threshold induction shock will not produce any evidence of a reflex excitation, whereas, when induction shocks of the same strength and of sufficient frequency are applied, a strong reflex contraction results. The fact that sub-threshold stimuli can bring about sub-threshold effects is also important in consideration of the result of interference. The relation between the intensity of the second stimulus and the degree of irritability of the system, the intensity of the stimulus being absolutely constant, depends, secondly, upon the momentary amount of irritability which exists just at the time when the second stimulus produces its effects. It is, therefore, clear that the response produced by interference must also alter with the momentary degree of irritability in a manner analogous with variations of the intensity of the second stimulus. One must, therefore, know the factors which control the momentary degree of excitation.

Fig. 51.

Effect of sub-threshold stimuli. o—Level of the ideal threshold. s—Level of the threshold of perceptible effect.

Fig. 52.

The first factor to be considered is the moment of time in which the second stimulus is applied, that is, the interval between the first and the second stimulus. If, for example, a weak second stimulus follows very quickly after the first, the stimulus will bring about no response, as the system at the time of its application is in a relative refractory period. (Figure [48].) The stimulus is, therefore, under the threshold. If, however, a stimulus of the same strength is applied somewhat later, when the irritability has already increased to a somewhat greater extent, then at this moment the stimulus is above that of the threshold and a response is obtained which, on account of the state of irritability existing, is summated. (Figure [52].) But further, it is not a question of the absolute interval between the stimuli, but rather to the relative interval to the specific rapidity of the reaction of the living substance under consideration. There are living substances, as we have seen, in which the refractory period is unusually short, as, for instance, the nerve. There are other substances wherein this period lasts a considerable time after stimulation, that is, before the irritability returns to the original level, as, for example, the smooth muscle. Indeed, depending upon the specific properties of a system, a short or a long interval is required before a stimulus of a given intensity is again operative. Finally, in one and the same living system the duration of the refractory period can be very different, depending upon the momentary state of the system. Above all we know that the refractory period is considerably prolonged in fatigue and likewise after the influence of other agents, as narcotics, lowering of the temperature, etc. In such states a second stimulus remains inoperative when it follows at a definite interval from the first, whereas under normal conditions the same stimulus applied at the same interval would be operative.