Neurons of the Deep Cerebellar Nuclei

The cerebellar nuclei (CN) are the final processing unit of the cerebellar circuitry, essentially combining most of cerebellar afferent inputs with the output of cerebellar cortex, transmitting the result of this integration to the extracerebellar targets

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46

€pfel Marylka Yoe Uusisaari and Thomas Kno

Abstract

The cerebellar nuclei (CN) are the final processing unit of the cerebellar circuitry, essentially combining most of cerebellar afferent inputs with the output of cerebellar cortex, transmitting the result of this integration to the extracerebellar targets, placing them at a key position in understanding cerebellar function on system level. Until recently, the nuclei have been treated as a simple relay nucleus with little neuronal diversity or computational capability. However, with the advent of genetically encoded fluorescent labels, the complexity and diversity of the CN neuronal circuits has started to become clear, rising doubts on the simplistic view of CN role in cerebellar computation. In this chapter, the currently known CN neuronal types – in all, four projecting neuron and two interneuron types – are described, and recommendations on their electrophysiological identification (in mouse slice preparation) are provided.

M.Y. Uusisaari (*) Laboratory for Neuronal Circuit Dynamics, RIKEN Brain Science Institute, 2-1 Hirosawa, Wako City, Saitama, 351-0198, Japan and Theoretical and Experimental Neurobiology Unit, Okinawa Institute of Science and Technology (OIST), 1919-1, Tancha, Onna–Son, Okinawa, 904-2423, Japan e-mail: [email protected] T. Kn€opfel Laboratory for Neuronal Circuit Dynamics, RIKEN Brain Science Institute, 2-1 Hirosawa, Wako City, Saitama, 351-0198, Japan e-mail: [email protected] M. Manto, D.L. Gruol, J.D. Schmahmann, N. Koibuchi, F. Rossi (eds.), 1101 Handbook of the Cerebellum and Cerebellar Disorders, DOI 10.1007/978-94-007-1333-8_46, # Springer Science+Business Media Dordrecht 2013

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M.Y. Uusisaari and T. Kn€ opfel

Introduction The afferent and efferent synaptic connections of the cerebellar nuclei (CN) places them in a key position where they can integrate signals from the brain stem, the inferior olive, and the spinal cord with the output from cerebellar cortex and to provide the major efferent pathway of the cerebellum. While this conclusion can be derived based on purely anatomical knowledge, electrophysiological data are required to define the level of complexity at which the CN integrate incoming signals to generate their output. It is largely unknown to which extent the CN contributes to the generation of signals that are attributed to cerebellar function such as timing signals and temporal patterns. Arguments against a simple relay function of the CN include the indications that at least a portion of motor memory is stored in the CN (Wada et al. 2007). As a first step toward understanding operations that are accomplished by the neuronal circuits of the CN, the major circuit elements – the major CN neuronal types – are now identified and characterized. The aim of this chapter is to summarize the efforts made in several laboratories toward this goal. We mostly focus on the lateral CN and data described in a series of works (Uusisaari et al. 2007; Uusisaari and Kn€ opfel 2008, 2009, 2010; Bagnall et al. 2009) that are ba