Drawing of Purkinje cells (A) and granule cells (B) from pigeon cerebellum by Santiago Ramón y Cajal, 1899. Instituto Santiago Ramón y Cajal, Madrid, Spain.
While anatomically similar, granule cells in different brain regions are functionally diverse. For instance, olfactory bulb granule cells are GABAergic and axonless, while granule cells in the dentate gyrus have glutamatergic projection axons. Interestingly, these two populations of granule cells are also the only major neuronal populations that undergo adult neurogenesis, while cerebellar and cortical granule cells do not.
Cerebellar granule cells account for nearly half of the neurons in the central nervous system. Granule cells receive excitatory input from mossy fibers originating from pontinenuclei. Cerebellar granule cells send parallel fibers up through the Purkinje layer into the molecular layer where they branch out and spread through Purkinje celldendritic arbors. These parallel fibers form thousands of excitatory synapses onto the intermediate and distal dendrites of Purkinje cells using glutamate as a neurotransmitter.
Layer 4 granule cells of the cerebral cortex receive driving inputs from thalamus and convey driving inputs largely to supragranular layers 2-3, but also to infragranular layers of the cerebral cortex.