in-phase (that is, simultaneously in the same direction) with vestibular stimulation but at a gradually greater amplitude (BOX2) . This mismatch training forces the mouse to make a compensatory eye movement during vestibular stimulation in the dark that is opposite in direction to that before the training because the error signals of retinal slip during the training in the light are reversed in direction. In line with the deficits in VOR phase reversal adaptation in Nr2amice and Nr2aCmice, local microperfusion of α7nAchR agonists into the granular layer of the flocculus, which is the main cerebellar lobule for controlling compensatory eye movements, affects VOR gain decrease adaptation34This form of VOR adaptation also occurs when a visual stimulus is constantly given in-phase with vestibular stimulation but at constant amplitudes. Together, these results suggest that NMDAR-mediated potentiation at the mossy fibre–granule cell synapse may contribute to vestibule cerebellar learning but not to basic motor performance. Interestingly, the number of mossy fibre filopodia in the granular layer may correlate with the strength of cued fear conditioning 36 , which raises the possibility that presynaptic LTP at mossy fibre terminals has a morphological substrate.