“…Purkinje cells are ideally suited to read out the strength of inhibitory synapses even in the absence of excitation, because they are spontaneously active (Häusser and Clark, 1997;Bean, 1997, 1999) and because inhibitory inputs have a powerful influence on Purkinje cell spiking (Midtgaard, 1992;Häusser and Clark, 1997). Moreover, PFs activate a feedforward inhibitory microcircuit, generating a precise, stereotyped excitatory-inhibitory temporal sequence (Eccles et al, 1966;Brunel et al, 2004;Mittmann et al, 2005), such that the strength of synaptic inhibition can control the timing and number of spikes activated by PFs (Mittmann et al, 2005). Thus, the efficacy of parallel fiber input depends on the "context" of inhibition provided by the feedforward inhibitory microcircuit.…”