Cerebellar Granule Cell
The cerebellar granule cells are the most numerous neurons of the brain. They are also among the most simple by emitting just four short unbranched dendrites innervated by the mossy fibers and a thin axon that branches into the parallel fibers. The granul
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Cerebellar Granule Cell
31
Egidio D’Angelo
Abstract
The cerebellar granule cells are the most numerous neurons of the brain. They are also among the most simple by emitting just four short unbranched dendrites innervated by the mossy fibers and a thin axon that branches into the parallel fibers. The granule cells are the only excitatory neurons of the cerebellum and receive inhibition uniquely through the Golgi cells. Since their early discovery by Camillo Golgi and Ramon y Cajal at the end of the nineteenth century, the granule cells have been the object of intensive investigation. The milestones have been the field recording of their activity in the 1960s, the discovery of their GABAergic inhibition in the 1970s and of their glutamatergic excitation in the 1980s, and the characterization of their membrane and synaptic mechanisms with patch-clamp techniques in the 1990s. Then in the last two decades, a series of careful electrophysiological and imaging investigations have unveiled major yet undisclosed functional properties of granule cells leading to a reevaluation of the function of the whole cerebellar cortex. The initial guess that granule cells simply retransmit incoming information (integrate and fire behavior) has been challenged by recent results showing that granule cells and their synapses are endowed with nonlinear transmission properties and are wired in a way allowing them to operate complex transformations of input signals in the spatiotemporal domain. Moreover, the prediction that the mossy fiber–granule cell synapse would lack long-term plasticity has been reversed by the discovery of complex forms of long-term potentiation and depression (LTP and LTD). These properties allow the granule neurons to provide a substantial contribution to the computational and learning properties of the cerebellum.
E. D’Angelo Department of Neuroscience, University of Pavia, Brain Connectivity Center, IRCCS C. Mondino, via Mondino 2, Pavia, I-27100, Italy e-mail: [email protected] M. Manto, D.L. Gruol, J.D. Schmahmann, N. Koibuchi, F. Rossi (eds.), 767 Handbook of the Cerebellum and Cerebellar Disorders, DOI 10.1007/978-94-007-1333-8_31, # Springer Science+Business Media Dordrecht 2013
768
E. D’Angelo
Introduction The granule cells were described initially by Camillo Golgi (1874, 1885, 1967) and subsequently analyzed in detail by Santiago Ramo´n y Cajal (1889, 1904, 1909, 1911). Cajal also gave special attention to the glomerulus, the specialized structure in which granule cell dendrites are synaptically connected with the mossy fibers and the Golgi cells (for historical review, see DAngelo et al. 2011; Galliano et al. 2010). The granule cells (Fig. 31.1a) are the smallest and most numerous neurons in the brain and show minor differences among species and across the different cerebellar sections (Eccles et al. 1967; Llinas 1969; Palay and Chan-Palay 1974; Ito 1984). The soma is spherical (about 5 mm in diameter) and is almost entirely occupied by the nucleus. The dendrites are four on average, short (about 13
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