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1 Laboratory for Neuronal Circuit Dynamics, Brain Science Insitute, RIKEN, Wako-shi, Saitama, Japan
2 Obata Research Unit, Brain Science Insitute, RIKEN, Wako-shi, Saitama, Japan
* To whom correspondence should be addressed. E-mail: tknopfel{at}brain.riken.jp.
The deep cerebellar nuclei (DCN) integrate inputs from the brain stem, the inferior olive, and the spinal cord with Purkinje cell output from cerebellar cortex and provide the major output of the cerebellum. Despite their crucial function in motor control and learning, the various populations of neurons in the DCN are poorly defined and characterized. Most importantly, differences in electrophysiological properties between glutamatergic and GABAergic cells of the DCN have been largely elusive. Here, we used glutamate decarboxylase (GAD) 67-green fluorescent protein (GFP) knock-in mice to unambiguously identify GABAergic (GAD-positive) and non-GABAergic (GAD-negative, most likely glutamatergic) neurons of the DCN. Morphological analysis of DCN neurons patch-clamped with biocytin-containing electrodes revealed a significant overlap in the distributions of the soma sizes of GAD-positive and GAD-negative cells. Compared with GAD-negative DCN neurons, GAD-positive DCN neurons fire broader action potentials, display stronger frequency accommodation and do not reach as high firing frequencies during depolarizing current injections. Furthermore, GAD-positive cells display slower spontaneous firing rates and have a more shallow current-to-frequency relationship than the GAD-negative cells but exhibit a longer-lasting rebound depolarization and associated spiking following a transient hyperpolarization. In contrast to the rather homogenous population of GAD-positive cells, the GAD-negative cells were found to comprise of two distinct populations as defined by cell size and electrophysiological features. We conclude that GABAergic DCN neurons are specialized to convey phasic spike rate information, whereas tonic spike rates are more faithfully relayed by the large non-GABAergic cells.
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