Unipolar brush cell axons form a large system of intrinsic mossy fibers in the postnatal vestibulocerebellum

The unipolar brush cells (UBCs), a class of neurons recently identified in the granular layer of the vestibulocerebellum, receive excitatory synaptic input from mossy fibers (MFs) in the form of a giant glutamatergic synapse. UBCs are provided with axons that bear synaptic endings situated at the center of glomeruli, similar to cerebellar MF afferents. A single MF stimulus evokes a prolonged train of action potentials in the UBC (Rossi et al., 1995), which is presumably distributed to postsynaptic targets. Knowledge of the synaptic connections of UBC axons is essential to define the role of these cells in the integration of vestibular signals in the cerebellar circuitry. To evaluate these connections, the nodulus (folium X) was isolated from vermal slices of postnatal day 8 mice, cultured for 2-4 or 15-30 days in vitro, and studied by electron and fluorescence microscopy. The peak of degeneration of extrinsic MF terminals, which have been severed from the parent cell bodies, was observed at 2 days in vitro (DIV). Quantification of degenerating and nondegenerating (e.g., intrinsic) MF terminals indicated that about half of the MF terminals were provided by local UBC axons synapsing on dendrites of granule cells and other UBCs. The proportion of nondegenerating vs. degenerating MF terminals terminating on UBCs also indicated that approximately two-thirds of the intrinsic MFs are involved in UBC-UBC connections. In long-term cultures, the granular layer appeared well preserved and the UBC axons formed an extensive system of MF collaterals. It is suggested that UBCs may act by spatially amplifying vestibular inputs carried by extrinsic MFs.