Evidence from both anatomical and physiological studies suggests that the ascending segment of the granule cell axon provides a large, driving input to overlying Purkinje cells. In the current experiments, we used dual recording electrodes to simultaneously record spike activity of Purkinje cells and multiunit field potential activity in the directly underlying granule cell layer. These dual recordings were performed both during periods of spontaneous ("background") firing and also after peripheral tactile stimulation. The results demonstrate that in the large majority of cases, there is a strong positive correlation between spontaneous Purkinje cell simple spikes and spontaneous activity in the immediately underlying granule cell layer. The strength of this correlation was dependent on both the firing rate of the Purkinje cell as well as on the rate of granule cell layer multiunit activity. In addition, for any given pair of recordings, the correlation seen during spontaneous activity accurately predicted the magnitude and time course of responses evoked by peripheral tactile stimulation. These results provide additional evidence that the synapses associated with the ascending segment of the granule cell axon have a substantial influence on Purkinje cell output. This relationship is considered in the context of our ongoing reevaluation of the physiological relationship between cerebellar granule and Purkinje cells.
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