BPRs are thought to reflect a stereotyped PC response to bursting granular layer inputs. The granule cells generate brief spike bursts in vitro (D’Angelo et al., 1995; Nieus et al., 2006) and in vivo (Chadderton et al., 2004; van Beugen et al., 2013; Powell et al., 2015) in response to bursts in the mossy fibers (Rancz et al., 2007). These granule cell bursts are finely regulated by GoC inhibition (Mapelli et al., 2009; Nieus et al., 2014) generating specific patterns in the number and timing of spikes (D’Angelo and De Zeeuw, 2009; Arleo et al., 2010). The granule cell bursts are then conveyed to molecular layer activating PCs and molecular layer interneurons, which in turn generate feed-forward inhibition on PCs (Santamaria et al., 2007; Rieubland et al., 2014; Zhang and Südhof, 2016). Therefore, granule cells and molecular layer interneuron set-up complex spatio-temporal patterns of activity generating PC responses that have been classified as type I, II and III (De Zeeuw et al., 2011) and are differentiated depending on Zebrin positive or negative (Z+ or Z−) PCs in different cerebellar areas (Zhou et al., 2014, 2015).