Anatomy Highly folded, surface area 500 cm2 (compared to 2300 cm2 of cerebral cortex) 3-layered: Granular, Purkinje, Molecular Voogd, Glickstein (1998)

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Anatomy Highly folded, surface area 500 cm2 (compared to 2300 cm2 of cerebral cortex) 3-layered: Granular, Purkinje, Molecular Voogd, Glickstein (1998) Trends Neurosci, 21:370-375 slice by Thomas Deerinck

Cerebellar Cortex Granule cells – most numerous in CNS, ~50 billion Axons form parallel fibers in molecular layer, up to 3 mm Purkinje cells sole output – inhibitory Interneurons: Golgi, stellate, basket PC target – vestibular and deep nuclei Rokni et al (2008) Front Neurosci, 2:192-198

Median cell counts in vertebrate cerebellar structures From data summarized in KWT Caddy and TJ Biscoe (1979) Phil Trans Roy Soc Lond Ser B 287:167-201. Additional measurements from BB Gould and P Rakic (1981) Exp Brain Res 44:195-206, R Alvarez and R Anadón (1987) J Hirnforsch 28:133-137, RJ Harvey and RMA Napper(1991) Prog Neurobiol 36:437-463, JA Heckroth (1994) J Comp Neurol 343:173-182, and R Alvarez-Otero et al (1996) J Comp Neurol 368:487-502. JC Eccles (1969) Naturwissenschaften 56:525-534

Cerebellar Output All Purkinje cells project to one of the cerebellar nuclei (dentate, interpositus, fastigial) or one of the vestibular nuclei

Purkinje cells Dendritic arbors, ~180,000 PF synapses in humans - however, at any given time up to 80% of these synapses are ‘silent’ - planar dendritic arbor - fast AMPA EPSCs, and slow mGluR1 - also receives 20-30 inhibitory stellate/basket cells GABAergic projection neuron Spontaneous pacemaking, 20-100 Hz - depends on a ‘resurgent’ sodium current Raman, Bean (1999) J Neurosci 19:1663-1674

Climbing fibers and complex spikes Strong multisynaptic connection - ~25,000 synapses - strongest afferent in CNS - glutamatergic Purkinje cell complex spikes - very large EPSP with Calcium spikes Complex results in a pause of ongoing simple spike firing Kreitzer et al. (2000) Neuron, 27:25-32 Davie et al. (2008) J Neurosci, 28:7599-7609

Apps and Garwicz (2005) Nat Rev Neurosci 6:297-311