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The Journal of Neurophysiology Vol. 82 No. 4 October 1999, pp. 1697-1709
Copyright ©1999 by the American Physiological Society
Department of Neuroscience, Johns Hopkins University School of Medicine, Baltimore, Maryland 21205
Aizenman, Carlos D. and
David J. Linden.
Regulation of the Rebound Depolarization and Spontaneous Firing
Patterns of Deep Nuclear Neurons in Slices of Rat Cerebellum. J. Neurophysiol. 82: 1697-1709, 1999. Current-clamp recordings were made from the deep cerebellar nuclei
(DCN) of 12- to 15-day-old rats to understand the factors that mediate
intrinsic spontaneous firing patterns. All of the cells recorded were
spontaneously active with spiking patterns ranging continuously from
regular spiking to spontaneous bursting with the former predominating.
A robust rebound depolarization (RD) leading to a Na+ spike
burst was elicited after the offset of hyperpolarizing current
injection. The voltage and time dependence of the RD was consistent
with mediation by low-threshold voltage-gated Ca2+
channels. In addition, induction of a RD also may be affected by
activation of a hyperpolarization-activated cation current, Ih. A RD could be evoked efficiently after
brief high-frequency bursts of inhibitory postsynaptic potentials
(IPSPs) induced by stimulation of Purkinje cell axons. IPSP-driven RDs
were typically much larger and longer than those elicited by direct
hyperpolarizing pulses of approximately matched amplitude and duration.
Intracellular perfusion of the Ca2+ buffer
bis-(o-aminophenoxy)-N,N,N',N'-tetraacetic
acid (BAPTA) dramatically enhanced the RD and its associated spiking,
sometimes leading to a plateau potential that lasted several hundred
milliseconds. The effects of BAPTA could be mimicked partly by
application of apamin, a blocker of small conductance
Ca2+-gated K+ channels, but not by paxilline,
which blocks large conductance Ca2+-gated K+
channels. Application of both BAPTA and apamin, but not paxilline, caused cells that were regularly spiking to burst spontaneously. Taken
together, our data suggest that there is a strong relationship between
the ability of DCN cells to elicit a RD and their tendency burst
spontaneously. The RD can be triggered by the opening of T-type
Ca2+ channels with an additional contribution of
hyperpolarization-activated current Ih. RD
duration is regulated by small-conductance Ca2+-gated
K+ channels. The RD also is modulated tonically by
inhibitory inputs. All of these factors are in turn subject to
alteration by extrinsic modulatory neurotransmitters and are, at least
in part, responsible for determining the firing modes of DCN neurons.
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