Glutamate transporters are believed to remove glutamate from the synaptic c
left only slowly because they cycle slowly. However, we show that when glut
amate binds to postsynaptic transporters at the cerebellar climbing fiber s
ynapse, it evokes a conformation change and inward current that reflect glu
tamate removal from the synaptic cleft within a few milliseconds, a time sc
ale much faster than the overall cycle time. Contrary to present models, gl
utamate removal does not require binding of an extracellular proton, and th
e time course of transporter anion conductance activation differs from that
of glutamate removal. The charge movement associated with glutamate remova
l is consistent with the majority of synaptically released glutamate being
removed from the synaptic cleft by postsynaptic transporters.