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
As a result of all these investigations, linked together in a
systematic series, the proof has now been obtained that the nerve like
all other living substances is fatigable. Its fatigue is solely the
manifestation of a prolonged refractory period and the extension of the
latter by continuous stimulation is, as in all aërobic substances, a
result of relative deficiency of oxygen.
[Illustration: Fig. 38.
Curve of irritability as demonstrated by action current of two nerves
in nitrogen, which are alternatively stimulated (plain line) and at
rest (dotted line). Recovery in nitrogen is always merely partial
and relative. It only increases on introduction of oxygen. (After
_Thörner_.) ]
To briefly summarize in conclusion, I will repeat that just as all
living systems show a refractory period after an excitation, in which
irritability is reduced, all living systems are likewise capable of
fatigue. Both are most intimately connected and are based fundamentally
on the facts of metabolism.
An excitating stimulus disturbs the metabolic equilibrium of rest
by suddenly bringing about increased decomposition of certain
substances. During and directly after the breaking down, irritability
is reduced in the same degree as the amount of substances required
for disintegration in response to a succeeding stimulus is decreased
and the quantity of the decomposition products is increased. This is
the refractory period. By the metabolic self-regulation in accordance
with the principle of chemical equilibrium, the original metabolic
equilibrium is restored after every excitation. Irritability,
therefore, increases in the same measure as this occurs, that is, in
the form of a logarithmic curve, until it again reaches the specific
degree of irritability of the particular system. The refractory period
diminishes. If the processes of disintegration and self-regulation are
delayed, either by want of substance necessary for breaking down or
the accumulation of decomposition substances, the refractory period is
prolonged and the response to every further stimulation decreased, that
is, the system is fatigued. In all aërobic organisms the retardation
of the course of excitation and self-regulation under a continuous
influence of stimuli is the result of the relative want of oxygen. The
processes of oxydative disintegration are prolonged and restricted by
relative deficiency of oxygen and merge more and more into anoxydative
decomposition. The products of incomplete oxydative and anoxydative
decomposition accumulate. Both factors decrease the strength of the
response after every stimulation. Thus the want of oxygen leads to
reduced activity. In the anaërobic organisms the refractory period and
symptoms of fatigue are, of course, produced by the relative deficiency
of other substances. Fatigue in the anaërobic systems has, however, so
far not been investigated. We advance very slowly, step by step, in
physiology, and, as in every science, an acquirement of a new knowledge
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