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
From the information here gained on the nature and origin of the
refractory period the conclusion must inevitably be drawn that in all
living substance there must exist, directly following an excitation,
a period of time in which its irritability is reduced, that is, under
proper conditions a refractory period can be demonstrated for every
living organism. Every living system possessing irritability undergoes
a period of reduced irritability at the time of and subsequent to every
excitation, for every excitation momentarily decreases the amount of
products capable of disintegration and increases the disintegration
products in the unit of space. As restitution involves time, a
stimulus occurring in the phase preceding complete restitution cannot
break down the same quantity of molecules as would be the case after
the establishment of complete restitution, that is, the response is
weaker, the irritability is decreased. The refractory period during and
subsequent to excitation is as much a general property of the living
substance as irritability and metabolic self-regulation.
This conclusion appears so self-evident that it would seem hardly
to call for emphasis were it not that even at the present time the
view is still widely held that the refractory period is a special
characteristic of certain forms of living substance. This assumption
is explained on the one hand by the fact that our information
concerning the refractory period is still of comparatively recent
date and that few physiologists are in the habit of connecting
special observations with general physiological conceptions, but also
for the reason that some investigators have vainly tried to find a
refractory period in certain forms of living substance. _Langendorff_
and _Winterstein_,[135] for instance, have not succeeded in proving
a refractory period for the spinal cord of the frog. _Langendorff_
stimulated the central sciatic stump with two stimuli in quick
succession and used the contractions of the triceps as indicator of
the response. He found that when the stimuli, if consisting in either
single induction shocks or faradic shocks, followed each other even at
intervals of .004 seconds the second stimulus was still operative, this
being perceptible in an increase of the contraction or with greater
intervals of time in a summation of two contractions. _Winterstein_
concludes from this that the development of a refractory period after
a stimulation is not a general property of all nerve centers. If
the experiments of _Langendorff_ failed to show the presence of a
refractory period it is not for the reason that this does not take
place in the centers of the spinal cord but rather results from the
fact that the conditions for the investigation were not suited for its
demonstration. In fact, _Fröhlich_[136] and especially _Vészi_[137]
have incontestably proved the existence of relative refractory periods
in the normal spinal cord.
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