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J Neurophysiol (March 23, 2005). doi:10.1152/jn.01114.2004
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Submitted on October 26, 2004
Accepted on March 17, 2005

Pitch of Complex Tones: Rate-Place and Interspike-Interval Representations in the Auditory Nerve

Leonardo Cedolin1* and Bertrand Delgutte2

1 Massachusetts Eye and Ear Infirmary, Eaton-Peabody Laboratory, Boston, MA, USA; Harvard-MIT Division of Health Sciences and Technology, Speech and Hearing Bioscience and Technology Program, Cambridge, MA, USA
2 Massachusetts Eye and Ear Infirmary, Eaton-Peabody Laboratory, Boston, MA, USA; Harvard-MIT Division of Health Sciences and Technology, Speech and Hearing Bioscience and Technology Program, Cambridge, MA, USA; Massachusetts Institute of Technology, Research Laboratory of Electronics, Cambridge, MA, USA

* To whom correspondence should be addressed. E-mail: cedro{at}mit.edu.

Harmonic complex tones elicit a pitch sensation at their fundamental frequency (F0), even when their spectrum contains no energy at F0, a phenomenon known as "pitch of the missing fundamental". The strength of this pitch percept depends upon the degree to which individual harmonics are spaced sufficiently apart to be "resolved" by the mechanical frequency analysis in the cochlea. We investigated the resolvability of harmonics of missing-fundamental complex tones in the auditory nerve (AN) of anesthetized cats at low and moderate stimulus levels, and compared the effectiveness of two representations of pitch over a much wider range of F0s (110-3520 Hz) than in previous studies. We found that individual harmonics are increasingly well resolved in rate responses of AN fibers as the characteristic frequency (CF) increases. We obtained rate-based estimates of pitch dependent upon harmonic resolvability by matching harmonic templates to profiles of average discharge rate against CF. These estimates were most accurate for F0s above 400-500 Hz, where harmonics were sufficiently resolved. We also derived pitch estimates from all-order interspike-interval distributions, pooled over our entire sample of fibers. Such interval-based pitch estimates, which are dependent upon phase-locking to the harmonics, were accurate for F0s below 1300 Hz, consistent with the upper limit of the pitch of the missing fundamental in humans. The two pitch representations are complementary with respect to the F0 range over which they are effective; however, neither is entirely satisfactory in accounting for human psychophysical data.




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