The membrane of giant molluscan neurons: electrophysiologic properties and the origin of the resting potential.

The molluscan neuron, because of its large size and accessibility, has been an important model for studying the electrophysiology of nerve cells. This review catalogs data about specific molluscan neurons, but the greater importance of this material is in the broad picture of how a neuronal membrane maintains internal potential and is responsive to changes in the environment. Electrical properties of the membrane. The mechanisms which contribute to the resting potential in molluscan neurons can be separated into ionic and metabolic components. When the electrogenic sodium pump is eliminated experimentally, the ionic component of the potential follows the constant field equation quite closely. Many of the "constants" and "parameters" which characterize the membrane of molluscan neurons are actually variables which depend upon temperature, ionic environment, and membrane potential. The evaluation of the electrical parameters is complicated by extensive infoldings of the somatic membrane, and by large axons which drain current from the soma. Most molluscan neurons have a very high specific membrane resistance and a correspondingly low potassium permeability. Membrane capacitance is close to the 1 microF/cm2 value which characterizes biological membranes. The current-voltage relation of molluscan neurons may be complicated by inward-going rectification, but if that is inhibited the I-V curve follows the prediction of either the constant field equation or a simple electrical model. Factors which modify membrane behavior. The resting potential of molluscan neurons is very sensitive to changes in temperature and Ko, through a combination of effects upon the electrogenic sodium pump, inward-going rectification, and the membrane "parameters". Inward-going rectification depends upon a rectifying K conductance, and can be eliminated by cold or the removal of Ko. Strong or prolonged currents have time-dependent effects upon the membrane, and excessive polarization leads to a "high conductance state". The underlying (non-rectifying) K permeability of the membrane is relatively insensitive to temperature and ionic changes, whereas the Na permeability increases with warming. Membrane resistance varies with both temperature and ions (because the I-V curve is sensitive to these conditions) but membrane capacitance is relatively insensitive to external factors. Electrogenic sodium transport. Sodium transport is electrogenic in molluscan neurons. It can be stimulated by warm temperatures and an excess of substrate (e.g. high Nai); it can be inhibited by cold, by an absence of substrate (e.g. low Ko), or by pharmacologic agents such as cyanide or ouabain.(ABSTRACT TRUNCATED AT 400 WORDS)