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
If we assume that at the moment when the entire amount of blood is
removed from the vascular system, no oxygen remains in the cells of the
spinal cord and muscle, then disintegration of the living substance
could from this instant take place exclusively anoxydatively, and there
would be no further oxydative breaking down into carbon dioxide and
water. The energy production compared in equal number of molecules,
taking the figures of _Lesser_ for the fermentation of sugar, would
approximately amount to about 3.8 per cent. of that of the energy
production in the oxydative disintegration of dextrose into carbon
dioxide and water. In reality, however, the tetanic convulsions are
at first exactly as violent as in the frog with a normal circulation.
There simply remains the assumption, therefore, that either the
disintegration as soon as it becomes _an_oxydative involves relatively
greater number of molecules than would be the case if it were oxydative
in nature, or to suppose that even after the complete displacement
of the blood a certain, though relatively small, amount of oxygen is
present in the cells which for a short time suffices for the taking
place of oxydative disintegration and with this an almost maximal
production of energy which naturally decreases as the oxygen is
consumed. It seems to me that the latter supposition contains more
probability than the first. To return, however, from this observation
to a further consideration of the animal we are studying, we see how
the complete tetanic convulsions in the refractory period which we
assumed to be .1 second are gradually transformed into incomplete
tetanus. After a time the tetanic convulsions become shorter after each
stimulus (Figure 32, B) and permit us to distinguish their individual
movements, even though the latter at first succeed each other still
very rapidly. Gradually this incomplete tetanic convulsion assumes the
form of a short series of individual contractions, distinctly separated
from each other and soon a stage is reached in which each reaction
to a peripheral stimulus consists merely in a single contraction.
(Figure 32, C.) The refractory period is, however, even now less than
a second. Nevertheless, with a further continuation of the experiment,
the refractory period becomes more and more prolonged, so that stimuli
succeeding each other at intervals of less than a second are without
effect. It is possible at this stage, as _Tiedemann_[142] did, to
graphically record the reactions. He severed the sciatic nerve on one
side and stimulated its central stump, at the same time connecting the
triceps with a writing lever. It is then found that when the single
induction shocks follow each other at intervals of a second or more
every stimulus produces a contraction, but that on the contrary only
the first stimulus of a rhythmical series is operative and all those
succeeding ineffectual, if the stimuli follow each other at shorter
intervals.
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