a, Top, task illustration. Bottom, average licking response (11 sessions in 5 mice). b–d, Trial averaged response of three example granule cells (solid traces) superimposed on licking response (dashed). Dashed and solid vertical lines indicate the time of tone onset and reward delivery, respectively. First row compares rewarded trials and randomly interspersed omitted reward trials. Second row compares rewarded trials to interspersed unexpected rewards not preceded by a tone (trial numbers in b–d: 178, 163, 163 rewarded, 26, 24, 24 omitted reward, and 26, 24, 24 unexpected reward, respectively). e, Plot of each cell’s response differences between rewarded and omitted reward trials (x-axis), and between unexpected and expected reward trials (y-axis). Colors denote lobule origin of the cells (450 cells). Example cells from b–d are outlined.