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Faced with the bewildering characteristics of the ionic currents that cause the action potential in squid axon, Hodgkin and Huxley in 1952 developed an elegant model (1) that remains one of the most insightful descriptions of the functional properties of voltage-gated ion channels. One ingredient in this conceptual wizardry was the strict separation between gating, the process responsible for activation and inactivation of sodium and potassium channels, and permeation, the nearly ohmic behavior of fully activated channels. This clean separation between gating and permeation has been a bedrock principle in the biophysical characterization of voltage-dependent ion channels and is supported by many single-channel studies in which the opening and closing of channels is distinct from the properties of open channels. The maxim is that the gates open and close channels and pay scant attention to the flow of ions through the open channel. This dogma has some major exceptions, however, notably because of effects of permeant and pore-blocking ions on gating. Although such effects are quite variable among various classes of ion channels, the customary observation is that raising the concentration of either permeant or pore-blocking ions inhibits the gates from closing (2–15). The experimental data strongly suggest that, if an ion can bind deeply within the permeation pathway, it will tend to obstruct gate closure. This is the “foot-in-the-door” phenomenon originally described by Clay Armstrong to account for the effects of intracellular pore blockers on potassium channel gating (2, 3). The two papers from Armstrong’s laboratory …

The dual role of calcium: Pore blocker and modulator of gating