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
following each other oftener than ten times per minute produced
merely single initial contractions in the muscle concerned, that
is, in a series of stimuli of which the intervals are less than .1
per second, only the first produces response, whereas the following
occur in the refractory period, brought about by those preceding,
and are, therefore, inoperative. The nerve is fatigued by the quick
succession of stimuli. The normal nerve on the contrary invariably
responds, as known, to an even more rapid succession of stimuli with
a rhythmical excitation corresponding to the number of stimuli and
which is manifest in the muscle by a tetanus. This again confirmed the
identity of fatigue with the prolonged refractory period, conditioned
by the relative want of oxygen. It likewise explained the conditions
of the analogous behavior that _Wedensky_[159] had observed in the
narcotized nerve, but had neither recognized as manifestation of the
prolonged refractory period nor as fatigue. A further advance was made
by the investigations of _Thörner_. He placed two nerves of the same
frog in a double chamber under completely identical conditions with
the exception that one remained in a state of rest, whilst to the
other tetanic stimuli were applied. (Figure 35.) If this took place in
nitrogen, the irritability of the stimulated nerve invariably sank with
much greater velocity than that of the nonstimulated, whereas after an
introduction of oxygen, even when the stimulation was continuous, both
again recovered. In these experiments of _Thörner_[160] the action
current and not the muscle contraction served as indicator. Here the
fatigue of the medullated nerve brought about by the deficiency of
oxygen during prolonged stimulation is demonstrated in the most obvious
manner. (Figure 36.) _Thörner_[161] further succeeded by a continuous
stimulation of the nerve in obtaining even in atmospheric air the
indications of primary fatigue. The symptoms were exactly the same
as those characterizing fatigue of the muscle; the extension of the
course of excitation and, as a consequence of this, the appearance of
a summation of excitation produced by tetanic currents and a reduction
of irritability in response to single stimuli. The form of the curve,
resulting from alteration of irritability in fatigue and recovery,
likewise shows complete conformity with that of the muscle. (Figure
37.) Finally _Thörner_[162] proved that the nerve, when fatigued by
continuous tetanic stimulation in nitrogen, could also partially
recover in the latter if the stimulation was interrupted, whereas a
complete recovery could not take place unless a supply of oxygen was
introduced. (Figure 38.) This fact is in perfect accordance with the
relations found by _Verworn_, _Lipschütz_, in fatigue of the nervous
centers. It is the expression for the accumulation and removal of
fatigue substances, the depressing effect of which _Ranke_[163] first
established for the fatigued muscle.
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