The calyx of Held is an axosomatic terminal in the auditory brainstem that has attracted anatomists because of its giant size and physiologists because of its accessibility to patch-clamp recordings. The calyx allows the principal neurons in the medial nucleus of the trapezoid body (MNTB) to provide inhibition that is both well timed and sustained to many other auditory nuclei. The special adaptations that allow the calyx to drive its principal neuron even when frequencies are high include a large number of release sites with low release probability, a large readily releasable pool, fast presynaptic calcium clearance and little delayed release, a large quantal size, and fast AMPA-type glutamate receptors. The transformation from a synapse that is unremarkable except for its giant size into a fast and reliable auditory relay happens in just a few days. In rodents this transformation is essentially ready when hearing starts.