Between sound and perception: reviewing the search for a neural code

This review investigates the roles of representation, transformation and coding as part of a hierarchical process between sound and perception. This is followed by a survey of how speech sounds and elements thereof are represented in the activity patterns along the auditory pathway. Then the evidence for a place representation of texture features of sound, comprising frequency, periodicity pitch, harmonicity in vowels, and direction and speed of frequency modulation, and for a temporal and synchrony representation of sound contours, comprising onsets, offsets, voice onset time, and low rate amplitude modulation, in auditory cortex is reviewed. Contours mark changes and transitions in sound and auditory cortex appears particularly sensitive to these dynamic aspects of sound. Texture determines which neurons, both cortical and subcortical, are activated by the sound whereas the contours modulate the activity of those neurons. Because contours are temporally represented in the majority of neurons activated by the texture aspects of sound, each of these neurons is part of an ensemble formed by the combination of contour and texture sensitivity. A multiplexed coding of complex sound is proposed whereby the contours set up widespread synchrony across those neurons in all auditory cortical areas that are activated by the texture of sound.