Golgi cells are important players in the function of the cerebellar cortex, controlling the flow of incoming information from mossy fibres to the granule cells, which excite other cortical neurons. We recently showed that in anaesthetized rats most Golgi cells respond to stimulation of afferents from a very wide peripheral receptive field with a long-lasting depression of firing. These responses are mediated via a crossed ascending afferent pathway but the supraspinal part of this pathway is unknown. Here we have examined the hypothesis that the lateral reticular nucleus, a brainstem nucleus with known broad afferent convergence that projects mossy fibres to much of the cerebellum, is involved. First, we showed that single-pulse electrical microstimulation within the lateral reticular nucleus can elicit long-lasting depressions in Golgi cells, which are qualitatively similar to those evoked by peripheral afferent stimulation. Second, we showed that the amplitude of the depressions of Golgi cell firing evoked by peripheral stimulation can be reduced by pharmacological manipulation of the lateral reticular nucleus, either ipsilateral or contralateral to the stimulus site, with local injections of either the GABA(A) receptor agonist muscimol or the AMPA receptor blocker 6-cyano-7-nitroquinoxaline-2,3-dione. This evidence suggests that the lateral reticular nucleus is a relay nucleus in the brainstem for peripheral afferent information in a pathway that generates Golgi cell long-lasting depression responses.