Spatiotemporal Representation of the Pitch of Harmonic Complex Tones in the Auditory Nerve

The pitch of harmonic complex tones plays an important role in speech and music perception and the analysis of auditory scenes, yet traditional rate-place and temporal models for pitch processing provide only an incomplete description of the psychophysical data. To test physiologically a model based on spatiotemporal pitch cues created by the cochlear traveling wave (Shamma, 1985), we recorded from single fibers in the auditory nerve of anesthetized cat in response to harmonic complex tones with missing fundamentals and equal-amplitude harmonics. We used the principle of scaling invariance in cochlear mechanics to infer the spatiotemporal response pattern to a given stimulus from a series of measurements made in a single fiber as a function of fundamental frequency F-0. We found that spatiotemporal cues to resolved harmonics are available for F-0 values between 350 and 1100 Hz and that these cues are more robust than traditional rate-place cues at high stimulus levels. The lower F-0 limit is determined by the limited frequency selectivity of the cochlea, whereas the upper limit is caused by the degradation of phase locking to the stimulus fine structure at high frequencies. The spatiotemporal representation is consistent with the upper F-0 limit to the perception of the pitch of complex tones with a missing fundamental, and its effectiveness does not depend on the relative phase between resolved harmonics. The spatiotemporal representation is thus consistent with key trends in human psychophysics.