Modulation and control of synaptic transmission across the MNTB.

The aim of this review is to consider the various forms and functions of transmission across the calyx of Held/MNTB synapse and how its modulation might contribute to auditory processing. The calyx of Held synapse is the largest synapse in the mammalian brain which uses the conventional excitatory synaptic transmitter, glutamate. It is sometimes portrayed as the 'ultimate' in synaptic signalling: it is a synaptic relay in which a single axon forms one synaptic terminal onto one specific target neuron. Questions that are often raised are: "Why does such a large and secure synapse need any form of modulation? Surely it is built simply to guarantee firing an action potential in the target neuron? If this synapse is so secure, why is a synapse needed at all?" Investigating these questions explains some general limitations of transmission at synapses and provides insight into the ionic basis of neuronal function by bringing together in vivo and in vitro approaches. We will start by defining the firing behaviour of MNTB neurons in vitro (in response to synaptic stimulation or current injection) and in vivo (in response to sound) and examining the reasons for different types of firing under the two conditions. Then we will consider some of the mechanisms by which transmission can be regulated. We will finish by discussing the following hypothesis: modulation and adaptation of presynaptic and postsynaptic conductances at the calyx of Held relay synapse are aimed at maximising the security of sound onset encoding while providing secondary information on frequency spectrum, harmonic envelope and duration of sound throughout the later part of the response.