Biophysical Journal 12: 1583-1629 (1972)
© 1972 the Biophysical Society

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The Excitable Membrane

A Physiochemical Model

ABSTRACT

The model of the excitable membrane assumes common channels for Na⁺ and K⁺; the two ion species interact within the pores through their electrostatic forces. The electric field varies across the membrane and with time, as a result of ionic redistribution. Ionic flow is primarily controlled by energy barriers at the two interfaces and by Ca⁺⁺ adsorption at the external interface. When the membrane is polarized, the high electric field at the external interface acting on the membrane fixed charge keeps the effective channel diameter small, so that only dihydrated ions can cross the interface. The higher energy required to partially dehydrate Na⁺ accounts for its lower permeability when polarized. Depolarized, the channel entrance can expand, permitting quadrihydrated ions to pass; the large initial Na⁺ flow is the result of the large concentration ratio across the interface. The effect at the internal interface is symmetric; Na⁺ crosses with greater difficulty when the membrane is depolarized. Na⁺ inactivation occurs when the ion distribution within the membrane has assumed its new steady-state value. Calculations based on parameters consistent with physicochemical data agree generally with a wide range of experiments. The model does not obey the two fundamental Hodgkin-Huxley (HH) postulates (independence principle, ion flow proportional to thermodynamic potential). In several instances the model predicts experimental results which are not predicted by the HH equations.