The Journal of Neuroscience, February 15, 2003, 23(4):1506
Slow Na+ Inactivation and Variance Adaptation in
Salamander Retinal Ganglion Cells
Kerry J.
Kim and
Fred
Rieke
Department of Physiology and Biophysics, University of Washington,
Seattle, Washington 98195
The retina adapts to the temporal contrast of the light inputs. One
component of contrast adaptation is intrinsic to retinal ganglion
cells: temporal contrast affects the variance of the synaptic inputs to
ganglion cells, which alters the gain of spike generation. Here we show
that slow Na+ inactivation is sufficient to produce
the observed variance adaptation. Slow inactivation caused the
Na+ current available for spike generation to depend
on the past history of activity, both action potentials and
subthreshold voltage variations. Recovery from slow inactivation
required several hundred milliseconds. Increased current variance
caused the threshold for spike generation to increase, presumably
because of the decrease in available Na+
current. Simulations indicated that slow Na+
inactivation could account for the observed decrease in excitability. This suggests a simple picture of how ganglion cells contribute to
contrast adaptation: (1) increasing contrast causes an increase in
input current variance that raises the spike rate, and (2) the
increased spike rate reduces the available Na+
current through slow inactivation, which feeds back to reduce excitability. Cells throughout the nervous system face similar problems
of accommodating a large range of input signals; furthermore, the
Na+ currents of many cells exhibit slow
inactivation. Thus, adaptation mediated by feedback modulation of the
Na+ current through slow inactivation could serve as
a general mechanism to control excitability in spiking neurons.
Key words:
contrast adaptation; slow Na+
inactivation; modulation of Na+ current; retinal
ganglion cell; models for spike generation; adaptation; spike-frequency
adaptation; retinal signal processing
Copyright © 2003 Society for Neuroscience 0270-6474/03/2341506-11$05.00/0