Unraveling the cerebellar cortex: cytology and cellular physiology of large-sized interneurons in the granular layer

Neuronal network behaviors emerge from complex interactions between excitatory relay cells, principal cells and inhibitory interneurons. Therefore, characterizing homogeneous cell types and their properties is an essential step towards understanding information processing in the brain. The cerebellar cortex is generally described as a repetitive circuit composed of only five cell types. However, recent studies have revealed an unexpected diversity in the morphological, neurochemical and electrophysiological properties of the large-sized granular layer interneurons. These data are reviewed here with an emphasis on the synaptic interactions of the different cell types within the cerebellar cortex. The existence of a complex network of excitatory and inhibitory interneurons controlling the spatial and temporal pattern of granule cell firing is documented, providing insights into the cellular and synaptic processes underlying oscillations and synchronization in the cerebellar cortex.