ABSTRACT
Recent psychophysical studies suggest that normal-hearing (NH) listeners can use acoustic temporal-fine-structure (TFS) cues for accurately discriminating shifts in the fundamental frequency (F0) of complex tones, or equal shifts in all component frequencies, even when the components are peripherally unresolved. The present study quantified both envelope (ENV) and TFS cues in single auditory-nerve (AN) fiber responses (henceforth referred to as neural ENV and TFS cues) from NH chinchillas in response to harmonic and inharmonic complex tones similar to those used in recent psychophysical studies. The lowest component in the tone complex (i.e., harmonic rank N) was systematically varied from 2 to 20 to produce various resolvability conditions in chinchillas (partially resolved to completely unresolved). Neural responses to different pairs of TEST (F0 or frequency shifted) and standard or reference (REF) stimuli were used to compute shuffled cross-correlograms, from which cross-correlation coefficients representing the degree of similarity between responses were derived separately for TFS and ENV. For a given F0 shift, the dissimilarity (TEST vs. REF) was greater for neural TFS than ENV. However, this difference was stimulus-based; the sensitivities of the neural TFS and ENV metrics were equivalent for equal absolute shifts of their relevant frequencies (center component and F0, respectively). For the F0-discrimination task, both ENV and TFS cues were available and could in principle be used for task performance. However, in contrast to human performance, neural TFS cues quantified with our cross-correlation coefficients were unaffected by phase randomization, suggesting that F0 discrimination for unresolved harmonics does not depend solely on TFS cues. For the frequency-shift (harmonic-versus-inharmonic) discrimination task, neural ENV cues were not available. Neural TFS cues were available and could in principle support performance in this task; however, in contrast to human-listeners’ performance, these TFS cues showed no dependence on N. We conclude that while AN-fiber responses contain TFS-related cues, which can in principle be used to discriminate changes in F0 or equal shifts in component frequencies of peripherally unresolved harmonics, performance in these two psychophysical tasks appears to be limited by other factors (e.g., central processing noise).
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REFERENCES
Bernstein JG, Oxenham AJ (2003) Pitch discrimination of diotic and dichotic tone complexes: harmonic resolvability or harmonic number? J Acoust Soc Am 113:3323–3334
Bernstein JG, Oxenham AJ (2005) An autocorrelation model with place dependence to account for the effect of harmonic number on fundamental frequency discrimination. J Acoust Soc Am 117:3816–3831
Cariani PA, Delgutte B (1996) Neural correlates of the pitch of complex tones. I. Pitch and pitch salience. J Neurophysiol 76:1698–1716
Cedolin L, Delgutte B (2005) Pitch of complex tones: rate-place and interspike interval representations in the auditory nerve. J Neurophysiol 94:347–362
Cedolin L, Delgutte B (2010) Spatiotemporal representation of the pitch of harmonic complex tones in the auditory nerve. J Neurosci 30:12712–12724
Chintanpalli A, Heinz MG (2007) The effect of auditory-nerve response variability on estimates of tuning curves. J Acoust Soc Am 122:EL203–EL209
Goldstein J (1973) An optimum processor theory for the central formation of the pitch of complex tones. J Acoust Soc Am 54:1496–1516
Goldstein JL, Srulovicz P (1977) Auditory-nerve spike intervals as an adequate basis for aural frequency measurement. In: Evans EF, Wilson JP (eds) Psychophysics and physiology of hearing. Academic, London, pp 337–346
Guinan JJ Jr, Peake WT (1967) Middle-ear characteristics of anesthetized cats. J Acoust Soc Am 41:1237–1261
Hartmann WM (1997) Signals, sound, and sensation. American Institute of Physics, Woodbury
Heinz MG, Swaminathan J (2009) Quantifying envelope and fine-structure coding in auditory-nerve responses to chimaeric speech. J Assoc Res Otolaryngol 10:407–423
Heinz MG, Colburn HS, Carney LH (2001) Evaluating auditory performance limits: I. One-parameter discrimination using a computational model for the auditory nerve. Neural Comput 13:2273–2316
Henry KS, Heinz MG (2013) Effects of sensorineural hearing loss on temporal coding of narrowband and broadband signals in the auditory periphery. Hear Res 303:39–47
Hopkins K, Moore BCJ (2007) Moderate cochlear hearing loss leads to a reduced ability to use temporal fine structure information. J Acoust Soc Am 122:1055–1068
Houtsma AJM, Smurzynski J (1990) Pitch identification and discrimination for complex tones with many harmonics. J Acoust Soc Am 87:304–310
Jackson HM, Moore BCJ (2014) The role of excitation-pattern, temporal-fine-structure, and envelope cues in the discrimination of complex tones. J Acoust Soc Am. In press
Johnson DH (1980) The relationship between spike rate and synchrony in responses of auditory-nerve fibers to single tones. J Acoust Soc Am 68:1115–1122
Joris PX (2003) Interaural time sensitivity dominated by cochlea-induced envelope patterns. J Neurosci 23:6345–6350
Kale S, Heinz MG (2010) Envelope coding in auditory nerve fibers following noise-induced hearing loss. J Assoc Res Otolaryngol 11:657–673
Kale S, Micheyl C, Heinz MG (2013) Effects of sensorineural hearing loss on temporal coding of harmonic and inharmonic tone complexes in the auditory nerve. Adv Exp Med Biol 787:109–118
Louage DH, Van Der Heijden M, Joris PX (2004) Temporal properties of responses to broadband noise in the auditory nerve. J Neurophysiol 91:2051–2065
Meddis R, Hewitt MJ (1992) Modeling the identification of concurrent vowels with different fundamental frequencies. J Acoust Soc Am 91:233–245
Meddis R, O'Mard L (1997) A unitary model of pitch perception. J Acoust Soc Am 102:1811–1820
Micheyl C, Dai H, Oxenham AJ (2010) On the possible influence of spectral- and temporal-envelope cues in tests of sensitivity to temporal fine structure. J Acoust Soc Am 127:1809–1810
Micheyl C, Xiao L, Oxenham AJ (2012) Characterizing the dependence of pure-tone frequency difference limens on frequency, duration, and level. Hear Res 292:1–13
Moore BCJ (2012) Introduction to the psychology of hearing, 6th edn. Brill, Leiden
Moore BCJ, Glasberg BR (2010) The role of temporal fine structure in harmonic segregation through mistuning. J Acoust Soc Am 127:5–8
Moore BCJ, Gockel HE (2011) Resolvability of components in complex tones and implications for theories of pitch perception. Hear Res 276:88–97
Moore BCJ, Sek A (2009a) Sensitivity of the human auditory system to temporal fine structure at high frequencies. J Acoust Soc Am 125:3186–3193
Moore BCJ, Sek A (2009b) Development of a fast method for determining sensitivity to temporal fine structure. Int J Audiol 48:161–171
Moore BCJ, Sek A (2011) Effect of level on the discrimination of harmonic and frequency-shifted complex tones at high frequencies. J Acoust Soc Am 129:3206–3212
Moore BCJ, Glasberg BR, Hopkins K (2006a) Frequency discrimination of complex tones by hearing-impaired subjects: evidence for loss of ability to use temporal fine structure. Hear Res 222:16–27
Moore BCJ, Glasberg BR, Flanagan HJ, Adams J (2006b) Frequency discrimination of complex tones; assessing the role of component resolvability and temporal fine structure. J Acoust Soc Am 119:480–490
Moore BCJ, Hopkins K, Cuthbertson S (2009) Discrimination of complex tones with unresolved components using temporal fine structure information. J Acoust Soc Am 125:3214–3222
Nelson DA, Stanton ME, Freyman RL (1983) A general equation describing frequency discrimination as a function of frequency and sensation level. J Acoust Soc Am 73:2117–2123
Oxenham AJ, Micheyl C, Keebler MV (2009) Can temporal fine structure represent the fundamental frequency of unresolved harmonics? J Acoust Soc Am 125:2189–2199
Plack CJ, Oxenham AJ (2005) The psychophysics of pitch. In: Plack CJ, Fay RR, Oxenham AJ, Popper AN (eds) Pitch: neural coding and perception. Springer, New York, pp 7–55
Plack CJ, Fay RR, Oxenham AJ, Popper AN (2005) Pitch: neural coding and perception. Springer, New York
Rose JE, Brugge JF, Anderson DJ, Hind JE (1969) Some possible neural correlates of combination tones. J Neurophysiol 32:402–423
Santurette S, Dau T (2011) The role of temporal fine structure information for the low pitch of high-frequency complex tones. J Acoust Soc Am 129:282–292
Schouten JF, Ritsma RJ, Cardozo BL (1962) Pitch of the residue. J Acoust Soc Am 34:1418–1424
Shackleton TM, Carlyon RP (1994) The role of resolved and unresolved harmonics in pitch perception and frequency modulation discrimination. J Acoust Soc Am 95:3529–3540
Shera CA, Guinan JJ Jr, Oxenham AJ (2010) Otoacoustic estimation of cochlear tuning: validation in the chinchilla. J Assoc Res Otolaryngol 11:343–365
Shofner WP (2011) Spectral processing does not give rise to behaviorally relevant cues for pitch perception in mammals. J Acoust Soc Am 129:2592–2593
Siebert WM (1970) Frequency discrimination in the auditory system: place or periodicity mechanisms? Proc IEEE 58:723–750
Srulovicz P, Goldstein JL (1983) A central spectrum model: a synthesis of auditory-nerve timing and place cues in monaural communication of frequency spectrum. J Acoust Soc Am 73:1266–1276
Tan Q, Carney LH (2003) A phenomenological model for the responses of auditory-nerve fibers. II. Nonlinear tuning with a frequency glide. J Acoust Soc Am 114:2007–2020
Terhardt E (1974) Pitch, consonance, and harmony. J Acoust Soc Am 55:1061–1069
Wier CC, Jesteadt W, Green DM (1977) Frequency discrimination as a function of frequency and sensation level. J Acoust Soc Am 61:178–184
Wightman FL (1973) The pattern-transformation model of pitch. J Acoust Soc Am 54:407–416
Zhang X, Heinz MG, Bruce IC, Carney LH (2001) A phenomenological model for the responses of auditory-nerve fibers: I. Nonlinear tuning with compression and suppression. J Acoust Soc Am 109:648–670
Zilany MSA, Bruce IC (2006) Modeling auditory-nerve responses for high sound pressure levels in the normal and impaired auditory periphery. J Acoust Soc Am 120:1446–1466
Zilany MSA, Bruce IC (2007) Representation of the vowel /ε/ in normal and impaired auditory nerve fibers: model predictions of responses in cats. J Acoust Soc Am 122:402–417
Zilany MSA, Bruce IC, Nelson PC, Carney LH (2009) A phenomenological model of the synapse between the inner hair cell and auditory nerve: long-term adaptation with power-law dynamics. J Acoust Soc Am 126:2390–2412
Acknowledgments
This research was supported by National Institutes of Health (NIH) grants R01-DC009838 (SK and MGH) and R01-DC05216 (CM). The authors thank Kenneth Henry and Jon Boley for help with data collection. We also acknowledge the helpful and thorough reviews from Associate Editor Chris Plack, Brian Moore, and an anonymous reviewer.
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Kale, S., Micheyl, C. & Heinz, M.G. Implications of Within-Fiber Temporal Coding for Perceptual Studies of F0 Discrimination and Discrimination of Harmonic and Inharmonic Tone Complexes. JARO 15, 465–482 (2014). https://doi.org/10.1007/s10162-014-0451-2
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DOI: https://doi.org/10.1007/s10162-014-0451-2