The Journal of Neuroscience, February 1, 2001, 21(3):1022-1032
Dynamics of Low-Threshold Spike Activation in Relay Neurons of
the Cat Lateral Geniculate Nucleus
Carolina
Gutierrez1,
Charles L.
Cox1,
John
Rinzel2, and
S. Murray
Sherman1
1 Department of Neurobiology, State University of New
York, Stony Brook, New York 11794-5230, and 2 Center for
Neural Science and Courant Institute of Mathematical Sciences, New York
University, New York, New York 10003
The low-threshold spike (LTS), generated by the transient
Ca2+ current IT,
plays a pivotal role in thalamic relay cell responsiveness and thus in
the nature of the thalamic relay. By injecting depolarizing current
ramps at various rates to manipulate the slope of membrane
depolarization (dV/dt), we found that an LTS occurred only if dV/dt exceeded a
minimum value of ~5-12 mV/sec. We injected current ramps of variable
dV/dt into relay cells that were
sufficiently hyperpolarized to de-inactivate
IT completely. Higher values of
dV/dt activated an LTS. However, lower
values of dV/dt eventually led to tonic
firing without ever activating an LTS; apparently, the inactivation of
IT proceeded before
IT could be recruited. Because the
maximum rate of rise of the LTS decreased with slower activating ramps
of injected current, we conclude that slower ramps allow increasing
inactivation of IT before the
threshold for its activation gating is reached, and when the injected
ramps have a sufficiently low dV/dt, the
inactivation is severe enough to prevent activation of an LTS. In the
presence of Cs+, we found that even the lowest
dV/dt that we applied led to LTS activation, apparently because Cs+ reduced the
K+ "leak" conductance and increased neuronal
input resistance. Nonetheless, under normal conditions, our data
suggest that there is neither significant window current (related to
the overlap of the inactivation and activation curves for
IT), rhythmogenic properties, nor
bistability properties for these neurons. Our theoretical results using
a minimal model of LTS excitability in these neurons are consistent with the experimental observations and support our conclusions. We
suggest that inputs activating very slow EPSPs (i.e., via metabotropic receptors) may be able to inactivate
IT without generating sizable IT and a spurious burst of action
potentials to cortex.
Key words:
low-threshold spike; thalamus; burst firing; T channel; window current; lateral geniculate nucleus
Copyright © 2001 Society for Neuroscience 0270-6474/01/2131022-11$05.00/0