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
The facts just mentioned have, however, a much deeper meaning. They
show us that it is possible by means of narcosis to convert an extreme
type of a living system, with decrementless conductivity, into another
extreme type of living substance, in which excitation in its progress
meets with a strong decrement, like that seen in the rhizopods. The
same results may also be obtained by asphyxiation and other forms of
temporary and permanent injury of the nerve. We are, therefore, in the
fortunate position in the case of the medullated nerve of having a
substance to study, which, depending upon conditions which are under
our control, may become a type in which conductivity occurs with
or without the presence of a decrement. We can likewise reduce the
irritability to various degrees, producing all intermediate gradations
between the two extremes. This latter is particularly valuable in that
it permits us to study the conditions in one and the same substance
necessary to bring about the various peculiarities of conductivity. The
great differences in the conductivity of excitation are conditioned by
variations in the degree of irritability. If the irritability of the
nerve is at the normal level the wave of excitation progresses to the
end of the nerve without manifesting a decrement of its intensity or
rapidity.
If the level of irritability of the intact nerve is artificially
reduced, the wave of excitation meets with a greater decrement and
reduces in velocity, and in fact disappears the more quickly in the
stretch of nerve, as the reduction in irritability is increased.
These three factors, irritability, intensity and velocity of the
progress of the wave of excitation, are inseparable. All living
substances may be grouped according to their capability of conducting
excitation into a long series, starting with those possessing the
least irritability, as we found in the rhizopods, then those having
greater irritability, as the smooth muscle and ganglion cells, then
those with still greater irritability, as the striped muscle, and
finally those having the greatest degree of irritability, as the
medullated nerves of the warm-blooded animal. Should the processes of
excitation, as we saw, result from the energy production following the
disintegration of the labile molecules of the living substance, then
the degree of irritability is determined by the chemical constitution
of the disintegrating molecules, by the number of molecules which are
broken down in a definite space and a given time, and by the nature
of the disintegration itself. All of these individual components, if
we observe the problem from the physical standpoint, are manifested
by the quantity of energy production. The higher the irritability of
a living system, the greater is the amount of energy production in a
given time and space which the stimulus produces. This has particular
interest from the standpoint of the extreme cases of medullated nerves
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