Irritability : $b a physiological analysis of the general effect of stimuli in living substanceVerworn, Max
Science
Irritability : $b a physiological analysis of the general effect of stimuli in living substance
Verworn, Max
Irritability
result was not always obtained. From this we can easily estimate the
refractory period of the nerve, which is present after reaching a state
of equilibrium under certain conditions. If we assume ten stimuli
per second to be the number required to produce slight fatigue when
stimulation is prolonged, we can conclude that the refractory period in
this state is somewhat longer than one tenth of a second. Even though
_Gotch_ in his investigations already cited placed the refractory
period of the normal nerve at about .005 second, this statement is in
no way contradictory to the figure which we have just given. _Gotch_
measured simply the duration of the absolute refractory period of
the normal nerve, in other words, the duration of the period in
which no excitation at all could be brought about. On the contrary,
my estimate, based upon the investigations of _Thörner_, refers to
the _total_ refractory period of the nerve, that is, to the point
of _complete_ recovery of the equilibrium of metabolism and of the
specific irritability. Experimental proof of this assumption is already
under way.
I have endeavored to show the elementary principles at the basis
of these extremely varied interference effects and to make a few
generalizations concerning the complicated conditions here concerned.
It has been shown that a great number of interference effects possess
characteristics in common if one takes into consideration the process
occurring in the course of a single excitation. The altered state
which exists in living substance until the complete disappearance of
excitation is the basis upon which to explain the altered effects
produced by a second stimulus. This state alters during the whole
course of the first stimulus until the original equilibrium of the
metabolism of rest is, by self-regulation, again reached. It is,
therefore, self-evident that the second stimulus must have different
effects depending upon the momentary state of the living system at the
time of its application. The state of the system differs depending on
the length of the interval in which the second stimulation follows the
first. The most important factor is the phase of the excitation period
and the reduction of irritability. The second important factor is the
intensity of the second stimulus; the relation of the two with each
other determines the response. But the specific properties of the given
systems must also be taken into consideration. It is important to know
if the living system possesses isobolic properties, that is, every
intensity of stimulation produces a _maximal_ liberation of energy, or
if it possesses a heterobolic character, that is, stimuli of different
strength bring about the liberation of _different_ amounts of energy.
It is further important to know the rapidity of reaction, whether the
system rapidly or slowly fatigues. In all cases it depends whether the
second stimulus produces a perceptible excitation or whether it occurs
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