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The Journal of Neurophysiology Vol. 87 No. 4 April 2002, pp. 2031-2042
Copyright ©2002 by the American Physiological Society
Systems Neurobiology Laboratories, The Salk Institute for Biological Studies, La Jolla, California 92037
Smith, Marianne R.,
Alexandra B. Nelson, and
Sascha du Lac.
Regulation of Firing Response Gain by Calcium-Dependent
Mechanisms in Vestibular Nucleus Neurons. J. Neurophysiol. 87: 2031-2042, 2002. Behavioral reflexes can be modified
by experience via mechanisms that are largely unknown. Within the
circuitry for the vestibuloocular reflex (VOR), neurons in the medial
vestibular nucleus (MVN) show adaptive changes in firing rate responses
that are correlated with VOR gain (the ratio of evoked eye velocity to
input head velocity). Although changes in synaptic strength are
typically assumed to underlie gain changes in the VOR, modulation of
intrinsic ion channels that dictate firing could also play a role.
Little is known, however, about how ion channel function or regulation contributes to firing responses in MVN neurons. This study examined contributions of calcium-dependent currents to firing responses in MVN
neurons recorded with whole cell patch electrodes in rodent brain stem
slices. Firing responses were remarkably linear over a wide range of
firing rates and showed modest spike frequency adaptation. Firing
response gain, the ratio of evoked firing rate to input current, was
reduced by increasing extracellular calcium and increased either by
lowering extracellular calcium or with antagonists to SK- and BK-type
calcium-dependent potassium channels and N- and T-type calcium
channels. Blockade of SK channels occluded gain increases via N-type
calcium channels, while blocking BK channels occluded gain increases
via presumed T-type calcium channels, indicating specific coupling of
potassium channels and their calcium sources. Selective inhibition of
Ca2+/calmodulin-dependent kinase II and
broad-spectrum inhibition of phosphatases modulated gain via
BK-dependent pathways, indicating that firing responses are tightly
regulated. Modulation of firing response gain by phosphorylation
provides an attractive mechanism for adaptive control of VOR gain.
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