Irritability : $b a physiological analysis of the general effect of stimuli in living substanceVerworn, Max
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Irritability : $b a physiological analysis of the general effect of stimuli in living substance
Verworn, Max
Irritability
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 R_{1} 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
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