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
If the muscle of the frog is isolated and rhythmically stimulated
with single induction shocks and the muscle contractions graphically
recorded, it will be found that the first perceptible alteration during
the course of stimulation is the increasing height in the curve,
which appears directly after the first contraction and becomes more
and more noticeable after every succeeding one. With the isolated
apex preparation of the frog’s heart an effect is produced which
_Bowditch_[145] has termed the “Treppe” and _Tiegel_,[146] _Minot_[147]
and others have obtained the same result for the skeletal muscle. The
_Treppe_ has been often regarded as an expression of increasing of
capability of the muscle following each succeeding stimulus in spite of
the fact that it is physiologically incomprehensible that an isolated
muscle can become more capable by increased demands. _Fröhlich_[148]
first threw light on this seeming contradiction by showing that the
increase in height of the muscle contraction in the _Treppe_ is in
reality the first indication of the beginning of fatigue, and _Fr.
Lee_[149] arrived at the same result. The increase in height of the
contraction curve depends upon the retardation of the course of
contraction. As the contraction extends over the muscle substance in
the form of a wave, a longer stretch of the muscle will be in a state
of contraction when the wave is more extended than when it is shorter,
that is, the shortening of the muscle will be greater, the contraction
curve higher, when the wave is more extended. With increasing fatigue
the retardation in the course of contraction, as _Rollet_[150] already
has shown, becomes continuously greater. (Figure 34.) The consequence
of this retardation in the course of contraction is, therefore,
perceptible in the rhythmically activated muscle in the form of
contracture. As fatigue increases, the muscle requires an increasing
length of time to relax to its full extent and in consequence the
period between the two stimuli is very soon insufficient for this to
occur. There remains a certain amount of shortening, when the next
contraction begins. This characteristic extension of the individual
contraction curve of the fatigued muscle is an expression of the
retardation of the oxydative disintegrating processes and of the
_Treppe_. It shows us that fatigue is perceptible to a slight degree
even after the first excitation. After every succeeding stimulus
the oxydative decomposition in the fatigued muscle is increasingly
prolonged. It is, therefore, self-evident that the capability of
action of the muscle likewise becomes less with increasing fatigue.
Every state of fatigue is, in fact, distinguished by the decrease of
response. This is perceptible in the later stages by the decline of
the height of contraction. Hence all symptoms of fatigue which we
observe form the expression of one single process; it is the constantly
increasing slowness of oxydative disintegration with increasing fatigue.
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