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

1 Eaton-Peabody Laboratory, Massachusetts Eye and Ear Infirmary, Boston; 2 Speech and Hearing Bioscience and Technology Program, Harvard-Massachusetts Institute of Technology Division of Health Sciences and Technology; and 3 Research Laboratory of Electronics, Massachusetts Institute of Technology, Cambridge, Massachusetts Submitted 26 October 2004; accepted in final form 17 March 2005 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 (110–3,520 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 on phase-locking to the harmonics, were accurate for F0s below 1,300 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. Address for reprint requests and other correspondence: L. Cedolin, Eaton-Peabody Laboratory, Massachusetts Eye and Ear Infirmary, 243 Charles St., Boston, MA 02114 (E-mail: cedro{at}mit.edu )