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J. Gen. Physiol.,
Volume 111, Number 2, February 1, 1998 271-294
From the Department of Cellular and Molecular Physiology, Yale University School of Medicine, New Haven, Connecticut 06520
The conformational changes associated with activation gating in Shaker potassium channels are functionally characterized in patch-clamp recordings made from Xenopus laevis oocytes expressing Shaker channels with
fast inactivation removed. Estimates of the forward and backward rates for transitions are obtained by fitting exponentials to macroscopic ionic and gating current relaxations at voltage extremes, where we assume that transitions
are unidirectional. The assignment of different rates is facilitated by using voltage protocols that incorporate
prepulses to preload channels into different distributions of states, yielding test currents that reflect different subsets of transitions. These data yield direct estimates of the rate constants and partial charges associated with three
forward and three backward transitions, as well as estimates of the partial charges associated with other transitions. The partial charges correspond to an average charge movement of 0.5 e0 during each transition in the activation
process. This value implies that activation gating involves a large number of transitions to account for the total gating charge displacement of 13 e0. The characterization of the gating transitions here forms the basis for constraining a detailed gating model to be described in a subsequent paper of this series.
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