Whether membrane capacitance is the determining factor in real neurons
is, of course, a matter of speculation. Quite a controversy is raging
over membrane capacity measurements (see Rall (21)), but the evidence
indicates that the capacity in the soma is considerably greater than
that in the axon (6), (22).
It should be added that increasing the capacitance until the membrane
model becomes inexcitable has little effect on the variety of available
simulated synaptic responses. Facilitation, antifacilitation, and
rebound are still present and still depend on the transmitter
inactivation rate. Thus, in the model, we can have a truly inexcitable
membrane which nevertheless utilizes the active membrane conductances
to provide facilitation or antifacilitation, and rebound. The simulated
subthreshold pacemaker potentials are much more realistic with the
increased capacitance, being lower in frequency and more natural in
form.
In one case, the electronic model predicted behavior which was
subsequently reported in real neurons. This was in respect to the
interaction of synaptic potentials and pacemaker potential. It was
noted in early experiments that when the model was set in a pacemaker
mode, and periodic spikes were applied to the simulated inhibitory
synapse, the pacemaker frequency could be modified; and, in fact,
it would tend to lock on to the stimulus frequency. This produced
a paradoxical effect whereby the frequency of spontaneous spikes
was actually increased by increasing the frequency of inhibitory
synaptic stimuli. At very low stimulus frequencies, the spontaneous
pacemaker frequency was not appreciably perturbed. As the stimulus
frequency was increased, and approached the basic pacemaker frequency,
the latter tended to lock on and follow further increases in the
stimulus frequency. When the stimulus frequency became too high for
the pacemaker to follow, the latter decreased abruptly in frequency
and locked on to the first subharmonic. As the stimulus frequency was
further increased, the pacemaker frequency would increase, then skip to
the next harmonic, then increase again, _etc._ This type of behavior
was observed by Moore _et al._ (23) in _Aplysia_ and reported at the
San Diego Symposium for Biomedical Electronics shortly after it was
observed by the author in the electronic model.
Public-domain text, read in full here on John Shaqi.
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