The brain stem auditory system of the chick has proven to be a useful model system for analyzing how the brain encodes temporal information. This paper reviews some of the work on a circuit in the brain stem that compares the timing of information coming from the two ears to determine the location of a sound source. The contralateral projection from the cochlear nucleus, nucleus magnocellularis (NM), to nucleus laminaris (NL) forms a delay line as it proceeds from medial to lateral across NL. NL neurons function like coincidence detectors in that they respond maximally when input from the two ears arrive simultaneously. This arrangement may allow NL to code sound space by the relative level of activity across the nucleus. The head anatomy of the chick allows for enhancement of the functional interaural time differences. Comparing the functional interaural time differences to the length of the neural delay line suggests that each NL can encode approximately one hemifield of sound space. Finally it is suggested that inhibitory input into the NM-NL circuit may provide a means to dynamically adjust the gain of the circuit to allow accurate coding of sound location despite changes in overall sound intensity.
KeywordsAuditory system; Sound localization; Nucleus magnocellularis; Nucleus laminaris; Coincidence detection; Interaural canal; GABA Temporal information is important in the auditory system both for identification of different sources of sound and for determining the location of sounds. Identification of different sound sources can be accomplished, to a large extent, using monaural cues. While localization of a sound source can also be accomplished, to some extent, with only one ear, significant localization cues are contained in the comparison of binaural information. Sounds located to one side will arrive at the near ear slightly before they arrive at the distant ear and, in general, these offset sounds will be slightly louder in the near ear than in the distant ear. By analyzing differences in the timing and intensity of information at the two ears, the brain can compute the location of a sound source.The circuitry for computing the location of a sound source begins in the brain stem. This report will review some of our work over the past several years on the circuit that begins the neural analysis of interaural time differences in the chick. Emphasis will be given to the anatomical, physiological and neurochemical adaptations in this model system to accomplish binaural sound localization. This manuscript is not meant to be an exhaustive review of the literature in the field, but rather, a story that exemplifies how adaptations at different levels of a system each contribute to enhancing the precision of an important task. Fig. 1A, provides a circuit for transforming time differences at the two ears into a place of maximal activation along an array of neurons. There are two main features of this model: 1) delay lines to an array of neurons and 2) neurons that function as coincidence detectors. As wi...