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.
Thus, we have shown that an electronic analog with all parameters except membrane capacitance fixed at values close to those of Hodgkin and Huxley, can provide all of the normal threshold or axonal behavior and also all of the subthreshold somatic and dendritic behavior outlined on [page 7]. Whether or not this is of physiological significance, it certainly provides a unifying basis for construction of electronic neural analogs. Simple circuits, based on the Hodgkin-Huxley model and providing all of the aforementioned behavior, have been constructed with ten or fewer inexpensive transistors with a normal complement of associated circuitry [(18)]. In the near future we hope to utilize several models of this type to help assess the information-processing capabilities not only of individual neurons but also of small groups or networks of neurons.
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Fields and Waves in Excitable
Cellular Structures
R. M. STEWART
Space General Corporation
El Monte, California
“Study of living processes by the physiological method only proceeded laboriously behind the study of non-living systems. Knowledge about respiration, for instance, began to become well organized as the study of combustion proceeded, since this is an analogous operation....”