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
[200] _Thörner_: “Weitere Untersuchungen über die Ermüdung des
markhaltigen Nerven: Die Ermüdung in Luft,” etc. Zeitschr. f. allgem.
Physiologie Bd. X, 1910.
[201] _Vészi_: “Zur Frage des Alles oder Nichtsgetzes beim
Strychninfrosche.” Zeitschr. f. allgem. Physiologie Bd. XII, 1911.
On the basis of these facts it is readily understood when a level
of equilibrium of lower intensity has been reached that excitation
produced by weak faradic stimulation must have weaker effects than when
strong stimuli are applied, for when the system assumes a heterobolic
type as the result of relative fatigue weak stimuli bring about weak,
and strong, stronger excitation. Consequently, during interference
induced by a second series of excitations, in the first case we have
the conditions favorable for inhibition, in the second for those of
summation. If we also assume that this characteristic alteration of
the isobolic character of the elementary nerve fibers which has been
shown to occur in fatigue, as seen when continued faradic stimulation
is employed, develops immediately after the beginning of stimulation
then we can readily understand the various kinds of effects produced
by interference observed in the reflex response following weak and
strong faradic stimulation to the different nerves in spite of the
fact that the nerve in the state of rest is a system isobolic in type.
Experimental evidence, therefore, must be brought forward to show that
faradic stimulation of short duration produces the above-mentioned
alteration in the character of the system. _Thörner_ in his experiments
on the nerve stimulated it faradically at least four minutes and always
found after this that excitation was reduced. After shorter intervals
of stimulation _Thörner_ made no test of the state of excitation. It
is, however, highly probable that a reduction of excitation is much
more quickly reached. Indeed, we are unavoidably compelled to accept
the assumption that even after the first single stimulus of the faradic
current, alterations of a slight degree are present which, after
repeated stimulation, become constantly greater and give to the system
a heterobolic character. As a result of fatigue, as we have already
seen, the refractory period becomes more and more prolonged. As the
individual shocks in faradic stimulation follow each other at regular
intervals, a necessary consequence is that the shocks are operative
before the refractory period has completely disappeared, otherwise
_Thörner_ could not have obtained fatigue produced by continued
stimulation. The intervals of the individual shocks must be somewhat
shorter than the duration of the refractory period, even in fatigue
of a very slight degree. It is very interesting in this connection
that _Thörner_ invariably obtained positive evidences of fatigue by
the application of stimuli at the rate of 10–12 per second. When the
number of stimuli per second was less than this the above-mentioned
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