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Physiological Reviews, Vol. 80, No. 2, April 2000, pp. 555-592
Copyright ©2000 by the American Physiological Society
Departments of Physiology and Anesthesiology, University of California at Los Angeles, School of Medicine, Los Angeles, California
Bezanilla, Francisco
The Voltage Sensor in Voltage-Dependent Ion Channels. Physiol. Rev. 80: 555-592, 2000.
In voltage-dependent Na, K, or Ca channels, the
probability of opening is modified by the membrane potential. This is
achieved through a voltage sensor that detects the voltage and
transfers its energy to the pore to control its gate. We present here
the theoretical basis of the energy coupling between the electric field
and the voltage, which allows the interpretation of the gating charge
that moves in one channel. Movement of the gating charge constitutes
the gating current. The properties are described, along with
macroscopic data and gating current noise analysis, in relation to the
operation of the voltage sensor and the opening of the channel.
Structural details of the voltage sensor operation were resolved
initially by locating the residues that make up the voltage sensor
using mutagenesis experiments and determining the number of charges per
channel. The changes in conformation are then analyzed based on the
differential exposure of cysteine or histidine-substituted
residues. Site-directed fluorescence labeling is then analyzed as
another powerful indicator of conformational changes that allows time
and voltage correlation of local changes seen by the fluorophores with
the global change seen by the electrophysiology of gating currents and
ionic currents. Finally, we describe the novel results on
lanthanide-based resonance energy transfer that show small distance
changes between residues in the channel molecule. All of the
electrophysiological and the structural information are finally
summarized in a physical model of a voltage-dependent channel in
which a change in membrane potential causes rotation of the S4 segment
that changes the exposure of the basic residues from an internally
connected aqueous crevice at hyperpolarized potentials to an externally
connected aqueous crevice at depolarized potentials.
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