In the Hodgkin-Huxley model, is the membrane potential, the voltage inside the axon relative to outside. The space clamp assumption makes it spatially uniform. The gating variables lie between zero and one: describes sodium activation, describes availability against sodium inactivation, and describes potassium activation. Thus large means channels remain available, not that inactivation is strong.
The left side is capacitive electric current per unit membrane area. The imposed is inward-positive applied electric current. The other terms are outward-positive ionic electric currents: sodium with electrical conductance , potassium with conductance , and a passive leak with conductance . Each current is its conductance times the difference between and the corresponding reversal potential , or . The coefficients are maximal conductances; the gate factors represent independent permissive gates in an ion channel.
For each gate, is the equilibrium opening/availability fraction at fixed voltage and its exponential relaxation time. The equation follows from two-state ion-channel gating: if , thenVoltage feedback couples these otherwise elementary gating equations to the membrane potential and produces action potentials.
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