While the complex spike error signal from a single pull can only provide miniscule LTD/LTP of parallel fibers onto a Purkinje cell, the combined effect across the population of Purks is robust enough to influence the very next joystick pull.

Imaging complex spikes in populations of Purkinje cells during learning reveals the answer: bands of cells respond similarly. Together, these cells encode both the magnitude and the direction of the behavioral error.

Like humans wearing prism goggles, mice learn to adapt their behavior in only a few trials (dark blue). They retain a memory of this adaptation, which they need to "unlearn" when returned to control conditions (light blue).

We trained mice to use a joystick to retrieve water. To test their cerebellar learning, we imposed a perturbation (gain = 2x; lower panel) that they needed to overcome to retrieve the reward.

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