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
In 1876 _Marey_[119] found that the irritability of the heart in
response to artificial stimulation was greatly reduced during the
systole, and that recovery took place during the following diastole.
(Figure 29.) This fact was already apparent from the observations made
by _Bowditch_[120] and _Kronecker_,[121] that by stimulation of the
isolated frog’s heart with single induction shocks, an artificial
systole can only be produced with certainty when the stimuli succeed
each other at certain intervals, which must be the longer as the
strength of the stimulation is weaker. _Marey_ calls this period
of reduced irritability “_phase réfractaire_” of the heart. The
refractory period of the heart has been made the subject of a great
number of investigations, especially by _Engelmann_ and his pupils.
It was _Engelmann_[122] especially who determined more exactly the
duration of the course of the refractory period. He found, namely, that
irritability disappears immediately before each systole and reappears
shortly before the beginning of the diastole, and again reaches its
original height at the end of the diastole. For a long time, however,
this refractory period was looked upon as a special peculiarity of the
heart. It was not until _Broca_ and _Richet_,[123] twenty years after
_Marey’s_ investigations, discovered an analogous refractory period for
the motor centers of the cerebral cortex of the dog. They first made
this observation on a dog affected with chorea, in which the choreic
movements rhythmically occurred in intervals of one second. They found
that after each movement electrical stimulation of the cortex remained
without result for about .5 seconds. During the next .25 seconds
stimulation was followed by a weak response and it was not until the
last .25 seconds before the next movement that a strong effect was
produced. They also found in the normal dog a refractory period after
every artificial stimulation equal to .1 second, so that the number of
contractions brought about by rhythmical electrical stimulation were
only ten per second. Following this, numerous other investigations of
the refractory period have been made on the central nervous system.
_Zwaardemaker_[124] and _Lans_ have observed a refractory period in
the eyelid reflex of the human being which, on stimulation of the
optic nerve, amounts to about .5–1 second; on the stimulation of
the trigeminus produced by blowing on the cornea on the other hand,
it is somewhat shorter, less than .25 seconds. _Zwaardemaker_[125]
also was able to demonstrate an analogous refractory period for the
swallowing reflex of the cat. Further a refractory period was found
and closely analyzed by _Verworn_[126] for the reflexes in the spinal
cord of the strychninized frog. _Dodge_[127] found a refractory period
in the knee jerk reflex of man. _Gotch_ and _Burch_[128] showed, by
two induction shocks following each other in quick succession, a
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