Cerebellum violates Marr-Albus predictions to train synapses on long-term anticipatory goals

studies, these theories predict synaptic adaptation based on temporal precision detected in time windows of 0ms to ∼100ms. These predictions have not been tested in intact animals. Using two-photon imaging from cerebellar Crus I in intact, awake mice, we show here that coincident stimulation of the PF and CF inputs does not initiate plasticity. Rather, long-term depression (LTD) is reliably evoked by ramping activity of the PF pathway that precedes a CF burst by 400ms. These observations demonstrate a cerebellar plasticity that is not about precision in coincidence with CF signaling. Rather, it shows that cerebellar learning centers on the evaluation of anticipatory PF signals by the CF input.