The synapse en marron between Golgi II neurons and mossy fibers in the rat's cerebellar cortex

Summary

Mossy fibers engage in synapses en marron with the somata of some Golgi II cells. These synapses resemble in all particulars the synapses en marron made by climbing fibers except for the distinctive characteristics of the presynaptic terminal. The mossy fiber, with its axial stream of neurofilaments and mitochondria and its loose aggregations of round synaptic vesicles, makes an extensive contact with the wrinkled surface of the Golgi II perikaryon. Synaptic complexes are confined to the depths and sides of the furrows in the Golgi cell. The free side of the mossy fiber terminal often articulates with large numbers of granule cell dendrites, an arrangement similar to that found in ordinary glomeruli.

These synaptic connections may be interpreted in the light of the physiological evidence that Golgi II cells inhibit granule cells that are not strongly activated by mossy fibers. Since each granule cell receives four to six mossy fibers, strong activation may require either a selected frequency pattern or synchrony of several inputs. The collateral inhibition indirectly evoked by the same mossy fiber via Golgi II cells would suppress those granule cells not receiving concurrent excitation from other mossy fibers or the favored pattern of excitation. In contrast, granule cells simultaneously activated by other mossy fibers would not be inhibited but would send impulses to the molecular layer. Thus, the glomerulus would behave as a filter that increases the signal-to-noise ratio of the excitatory input to the Purkinje cells.