In aggregate, these simulations showed that BPRs could fully exploit dendritic ionic channels (Llinás and Sugimori, 1980a,b) to generate voltage-dependent responses. However, this mechanism did not compromise the linearity of BPR input-output relationships supporting the predictions that PC may work as a linear perceptron (Brunel et al., 2004) and a perfect integrator (Phoka et al., 2010). Simulations supported the concept that BPRs represent a fundamental operating mode of PCs (Walter and Khodakhah, 2006, 2009; Chen et al., 2016). First, BPRs could discriminate among different input patterns coming from the mossy fibers and expanded/recoded in the granular and molecular layer (Marr, 1969). Second, BPRs sensitivity on the millisecond scale would allow PCs to operate as precise temporal devices (Eccles, 1973). Third, BPRs could discriminate among synaptic locations, generating an exquisite sensitivity to the topography of input patterns (Migliore et al., 2008). Finally, BPRs could generate complex combinations and sequences (Santamaria and Bower, 2005). By expressing these properties, BPRs would effectively integrate the spatio-temporal activity patterns generated in the cerebellar cortex into salient engrams, a prediction warranted experimental testing through electrophysiological, imaging and optogenetic recordings.