Quantifying the effects of the cochlear amplifier on temporal and average-rate information in the auditory nerve

An active mechanism referred to as the cochlear amplifier is believed to be responsible for sharp tuning and excellent sensitivity in the normal auditory system, and is impaired or absent in many common forms of sensorineural hearing loss. The cochlear amplifier is thought to benefit normal-hearing listeners, especially in complex listening environments in which hearing-impaired listeners have difficulty even with hearing aids. A modeling approach was developed to relate nonlinear physiological response properties associated with the cochlear amplifier to human psychophysical performance. Quantitative methods combined analytical and computational population models of the auditory-nerve (AN) with statistical decision theory to evaluate performance limits imposed by the random nature of AN discharges (modeled by a nonstationary Poisson process). A new theoretical approach was developed to predict performance for psychophysical tasks that use random-noise stimuli to mask signal information. The ability of temporal and average-rate information in the AN to account for human performance was evaluated for several listening tasks for which the cochlear amplifier has been suggested to be significant. The benefit of the cochlear amplifier for extending the auditory system’s dynamic range was evaluated in terms of AN information available for encoding changes in stimulus level. An analytical model included nonlinear gain, level-dependent phase, and high-, mediumand low-spontaneous-rate AN fibers. Both nonlinear-phase and average-rate information were required to encode levels up to 120 dB SPL based on a narrow range of AN characteristic frequencies. A physiologically realistic mechanism to decode nonlinear gain and phase cues is monaural, across-frequency coincidence detection. Level-discrimination performance of a coincidence-counter population matched human performance across the entire dynamic range of hearing at both low and high frequencies. Results suggest that the cochlear amplifier is beneficial for encoding sound level within narrow frequency regions, and has only a small influence on simple listening tasks in quiet. The cochlear amplifier alters tuning within the normal auditory system based on the spectral and temporal configuration of the stimulus. The influences of compression and suppression on psychophysical measures of auditory frequency selectivity were evaluated. Implications for the interpretation of psychophysical methods for estimating auditory filters are discussed. Thesis Supervisor: Laurel H. Carney, Ph.D. Title: Associate Professor of Biomedical Engineering, Boston University

[1]  E. de Boer,et al.  Masking and Discrimination , 1966 .

[2]  M B Sachs,et al.  Dynamic range of neural rate responses in the ventral cochlear nucleus of awake cats. , 1992, Journal of neurophysiology.

[3]  R H Gilkey,et al.  Effects of masker waveform and signal-to-masker phase relation on diotic and dichotic masking by reproducible noise. , 1985, The Journal of the Acoustical Society of America.

[4]  H. V. Trees Detection, Estimation, And Modulation Theory , 2001 .

[5]  Eric D Young,et al.  Effects of high sound levels on responses to the vowel /ε/ in cat auditory nerve , 1998, Hearing Research.

[6]  T Dau,et al.  A quantitative model of the "effective" signal processing in the auditory system. I. Model structure. , 1996, The Journal of the Acoustical Society of America.

[7]  E R Hafter,et al.  Just-noticeable differences of frequency for masked tones. , 1980, The Journal of the Acoustical Society of America.

[8]  M. Charles Liberman,et al.  Structure-Function Correlation in Noise-Damaged Ears: A Light and Electron-Microscopic Study , 1986 .

[9]  A Robert,et al.  A composite model of the auditory periphery for simulating responses to complex sounds. , 1999, The Journal of the Acoustical Society of America.

[10]  J. Allen,et al.  Using acoustic distortion products to measure the cochlear amplifier gain on the basilar membrane. , 1992, The Journal of the Acoustical Society of America.

[11]  J. Lentz,et al.  Different auditory filter bandwidth estimates based on profile analysis, notched noise, and hybrid tasks. , 1999, The Journal of the Acoustical Society of America.

[12]  G B Henning A model for auditory discrimination and detection. , 1967, The Journal of the Acoustical Society of America.

[13]  J. O. Pickles,et al.  Responses of auditory nerve fibres in the guinea pig to noise bands of different widths , 1980, Hearing Research.

[14]  M. L. Hicks,et al.  Effects of aspirin on psychophysical measures of frequency selectivity, two-tone suppression, and growth of masking. , 1999, The Journal of the Acoustical Society of America.

[15]  S. Bacon,et al.  Psychophysical measures of auditory nonlinearities as a function of frequency in individuals with normal hearing. , 1999, The Journal of the Acoustical Society of America.

[16]  N. Kiang,et al.  Tails of tuning curves of auditory-nerve fibers. , 1973, The Journal of the Acoustical Society of America.

[17]  A. Oxenham,et al.  A behavioral measure of basilar-membrane nonlinearity in listeners with normal and impaired hearing. , 1997, The Journal of the Acoustical Society of America.

[18]  William Bialek,et al.  Spikes: Exploring the Neural Code , 1996 .

[19]  R. Patterson Auditory filter shapes derived with noise stimuli. , 1976, The Journal of the Acoustical Society of America.

[20]  M F Dorman,et al.  Susceptibility to intraspeech spread of masking in listeners with sensorineural hearing loss. , 1983, The Journal of the Acoustical Society of America.

[21]  S Buus,et al.  Psychometric functions for level discrimination in cochlearly impaired and normal listeners with equivalent-threshold masking. , 1995, The Journal of the Acoustical Society of America.

[22]  J. E. Rose,et al.  Some neural mechanisms in the inferior colliculus of the cat which may be relevant to localization of a sound source. , 1966, Journal of neurophysiology.

[23]  Donald L. Snyder,et al.  Random Point Processes in Time and Space , 1991 .

[24]  Laurel H. Carney,et al.  Monaural, cross‐frequency coincidence detection as a mechanism for decoding perceptual cues provided by the cochlear amplifier , 1999 .

[25]  B. Delgutte,et al.  Neural correlates of the pitch of complex tones. II. Pitch shift, pitch ambiguity, phase invariance, pitch circularity, rate pitch, and the dominance region for pitch. , 1996, Journal of neurophysiology.

[26]  D G Sinex,et al.  Neural mechanisms of tone-on-tone masking: patterns of discharge rate and discharge synchrony related to rates of spontaneous discharge in the chinchilla auditory nerve. , 1986, Journal of neurophysiology.

[27]  J. M. Pickett,et al.  Some Masking Effects Produced by Low-Frequency Vowel Formants in Persons with Sensorineural Hearing Loss , 1975 .

[28]  Neal F. Viemeister,et al.  Intensity coding and the dynamic range problem , 1988, Hearing Research.

[29]  David Kay Ryugo,et al.  The Auditory Nerve: Peripheral Innervation, Cell Body Morphology, and Central Projections , 1992 .

[30]  A. Nuttall,et al.  Steady-state sinusoidal velocity responses of the basilar membrane in guinea pig. , 1996, The Journal of the Acoustical Society of America.

[31]  B. Moore Perceptual Consequences of Cochlear Damage , 1995 .

[32]  Allyn E. Hubbard,et al.  Computational Analysis of Hair Cell and Auditory Nerve Processes , 1996 .

[33]  Raimond L Winslow,et al.  Single-tone intensity discrimination based on auditory-nerve rate responses in backgrounds of quiet, noise, and with stimulation of the crossed olivocochlear bundle , 1988, Hearing Research.

[34]  B. Moore,et al.  Modeling the additivity of nonsimultaneous masking , 1994, Hearing Research.

[35]  W. S. Rhode Observations of the vibration of the basilar membrane in squirrel monkeys using the Mössbauer technique. , 1971, The Journal of the Acoustical Society of America.

[36]  K. Payton Vowel processing by a model of the auditory periphery: A comparison to eighth‐nerve responses , 1988 .

[37]  Philip H Smith,et al.  Coincidence Detection in the Auditory System 50 Years after Jeffress , 1998, Neuron.

[38]  T. Yin,et al.  Interaural time sensitivity in medial superior olive of cat. , 1990, Journal of neurophysiology.

[39]  J. Pickles An Introduction to the Physiology of Hearing , 1982 .

[40]  E. F. Evans,et al.  Comparisons of Physiological and Behavioural Properties: Auditory Frequency Selectivity , 1992 .

[41]  John J. Rosowski,et al.  Models of External- and Middle-Ear Function , 1996 .

[42]  W. M. Siebert,et al.  Frequency discrimination in the auditory system: Place or periodicity mechanisms? , 1970 .

[43]  M. Liberman,et al.  Auditory-nerve response from cats raised in a low-noise chamber. , 1978, The Journal of the Acoustical Society of America.

[44]  R L Freyman,et al.  Frequency discrimination as a function of tonal duration and excitation-pattern slopes in normal and hearing-impaired listeners. , 1986, The Journal of the Acoustical Society of America.

[45]  L H Carney,et al.  Effects of interaural time delays of noise stimuli on low-frequency cells in the cat's inferior colliculus. III. Evidence for cross-correlation. , 1987, Journal of neurophysiology.

[46]  M G Heinz,et al.  Masked detection thresholds and temporal integration for noise band signals. , 1994, The Journal of the Acoustical Society of America.

[47]  A. Oxenham,et al.  Basilar-membrane nonlinearity estimated by pulsation threshold. , 2000, The Journal of the Acoustical Society of America.

[48]  B C Moore,et al.  Psychoacoustic consequences of compression in the peripheral auditory system. , 1998, Psychological review.

[49]  I. Whitfield Discharge Patterns of Single Fibers in the Cat's Auditory Nerve , 1966 .

[50]  D. M. Green,et al.  Signal detection theory and psychophysics , 1966 .

[51]  D. M. Green,et al.  Intensity discrimination as a function of frequency and sensation level. , 1977, The Journal of the Acoustical Society of America.

[52]  S. McKee,et al.  Precise velocity discrimination despite random variations in temporal frequency and contrast , 1986, Vision Research.

[53]  E D Young,et al.  Discriminability of vowel representations in cat auditory-nerve fibers after acoustic trauma. , 1999, The Journal of the Acoustical Society of America.

[54]  Brian R Glasberg,et al.  Derivation of auditory filter shapes from notched-noise data , 1990, Hearing Research.

[55]  D. H. Johnson,et al.  The relationship between spike rate and synchrony in responses of auditory-nerve fibers to single tones. , 1980, The Journal of the Acoustical Society of America.

[56]  M B Sachs,et al.  Nonlinearities in auditory-nerve fiber responses to bandlimited noise. , 1980, The Journal of the Acoustical Society of America.

[57]  L A JEFFRESS,et al.  A place theory of sound localization. , 1948, Journal of comparative and physiological psychology.

[58]  L H Carney,et al.  Enhancement of neural synchronization in the anteroventral cochlear nucleus. I. Responses to tones at the characteristic frequency. , 1994, Journal of neurophysiology.

[59]  John J. Guinan,et al.  FREQUENCY DEPENDENCE OF STIMULUS-FREQUENCY-EMISSION PHASE: IMPLICATIONS FOR COCHLEAR MECHANICS , 2000 .

[60]  David M. Green,et al.  Profile Analysis: Auditory Intensity Discrimination , 1987 .

[61]  M. Sachs,et al.  Encoding of steady-state vowels in the auditory nerve: representation in terms of discharge rate. , 1979, The Journal of the Acoustical Society of America.

[62]  W. S. Rhode,et al.  Mechanical responses to two-tone distortion products in the apical and basal turns of the mammalian cochlea. , 1997, Journal of neurophysiology.

[63]  B. Kollmeier,et al.  Modeling auditory processing of amplitude modulation. I. Detection and masking with narrow-band carriers. , 1997, The Journal of the Acoustical Society of America.

[64]  J. Guinan,et al.  Evoked otoacoustic emissions arise by two fundamentally different mechanisms: a taxonomy for mammalian OAEs. , 1999, The Journal of the Acoustical Society of America.

[65]  G. A. Miller,et al.  Sensitivity to Changes in the Intensity of White Noise and Its Relation to Masking and Loudness , 1947 .

[66]  H S Colburn,et al.  Detection of tones in reproducible narrow-band noise. , 1991, The Journal of the Acoustical Society of America.

[67]  Mario A. Ruggero,et al.  Physiology and Coding of Sound in the Auditory Nerve , 1992 .

[68]  L. Robles,et al.  Two-tone suppression in the basilar membrane of the cochlea: mechanical basis of auditory-nerve rate suppression. , 1992, Journal of neurophysiology.

[69]  L. Robles,et al.  Basilar-membrane responses to tones at the base of the chinchilla cochlea. , 1997, The Journal of the Acoustical Society of America.

[70]  Steven Greenberg,et al.  Physiology of the Cochlear Nuclei , 1992 .

[71]  M. Sachs,et al.  Representation of steady-state vowels in the temporal aspects of the discharge patterns of populations of auditory-nerve fibers. , 1979, The Journal of the Acoustical Society of America.

[72]  Harvey b. Fletcher,et al.  Speech and hearing in communication , 1953 .

[73]  T. F. Weiss,et al.  A comparison of synchronization filters in different auditory receptor organs , 1988, Hearing Research.

[74]  D A Nelson,et al.  Extension of a temporal model of frequency discrimination: intensity effects in normal and hearing-impaired listeners. , 1985, The Journal of the Acoustical Society of America.

[75]  Julius L. Goldstein,et al.  Quantifying 2-factor phase relations in non-linear responses from low characteristic-frequency auditory-nerve fibers , 1995, Hearing Research.

[76]  J. Goldberg,et al.  Response of binaural neurons of dog superior olivary complex to dichotic tonal stimuli: some physiological mechanisms of sound localization. , 1969, Journal of neurophysiology.

[77]  H. Duifhuis Consequences of peripheral frequency selectivity for nonsimultaneous masking. , 1973, The Journal of the Acoustical Society of America.

[78]  B C Moore,et al.  Speech reception thresholds in noise with and without spectral and temporal dips for hearing-impaired and normally hearing people. , 1998, The Journal of the Acoustical Society of America.

[79]  W A Yost,et al.  Measures of intensity discrimination. , 1982, The Journal of the Acoustical Society of America.

[80]  R V Harrison,et al.  Rate-versus-intensity functions and related AP responses in normal and pathological guinea pig and human cochleas. , 1981, The Journal of the Acoustical Society of America.

[81]  Mario A. Ruggero,et al.  Auditory-nerve responses to low-frequency tones: Intensity dependence , 1996 .

[82]  Dianne J. Van Tasell,et al.  Hearing Loss, Speech, and Hearing Aids , 1993 .

[83]  H S Colburn,et al.  Theory of binaural interaction based on auditory-nerve data. I. General strategy and preliminary results on interaural discrimination. , 1973, The Journal of the Acoustical Society of America.

[84]  B. Moore,et al.  Gap detection and masking in hearing-impaired and normal-hearing subjects. , 1987, The Journal of the Acoustical Society of America.

[85]  J. Guinan,et al.  Auditory-nerve-fiber responses to high-level clicks: interference patterns indicate that excitation is due to the combination of multiple drives. , 2000, The Journal of the Acoustical Society of America.

[86]  H. Steven Colburn,et al.  Internal and external noise in binaural detection , 1978, Hearing Research.

[87]  M J Osberger,et al.  Discrimination of formant frequency transitions in synthetic vowels. , 1973, Journal of speech and hearing research.

[88]  H. Steven Colburn,et al.  Computational Models of Binaural Processing , 1996 .

[89]  B C Moore,et al.  Short-term temporal integration: evidence for the influence of peripheral compression. , 1997, The Journal of the Acoustical Society of America.

[90]  D J Van Tasell,et al.  Sensorineural hearing loss and the discrimination of vowel-like stimuli. , 1984, The Journal of the Acoustical Society of America.

[91]  D M Green,et al.  Successive versus simultaneous comparison in auditory intensity discrimination. , 1983, The Journal of the Acoustical Society of America.

[92]  Brian C. J. Moore,et al.  Chapter 5 – Frequency Analysis and Masking , 1995 .

[93]  M C Teich,et al.  Pulse-number distribution for the neural spike train in the cat's auditory nerve. , 1983, The Journal of the Acoustical Society of America.

[94]  B. Delgutte Physiological Models for Basic Auditory Percepts , 1996 .

[95]  M. Sachs,et al.  Two-tone inhibition in auditory-nerve fibers. , 1968, The Journal of the Acoustical Society of America.

[96]  Dianne J. Van Tasell,et al.  Perception of second-formant transitions by hearing-impaired persons. , 1980 .

[97]  H. Steven Colburn Erratum: ''Theory of binaural interaction based on auditory-nerve data. II. Detection of tones in noise'' [J. Acoust. Soc. Am. 61, 525-533 (1977)] , 1977 .

[98]  C. Plack,et al.  Beneficial effects of notched noise on intensity discrimination in the region of the "severe departure". , 1998, The Journal of the Acoustical Society of America.

[99]  Fan-Gang Zeng,et al.  On the role of the olivocochlear bundle in hearing : 16 case studies , 2013 .

[100]  Eric Javel,et al.  Physiological and psychophysical correlates of temporal processes in hearing , 1988, Hearing Research.

[101]  B. Delgutte,et al.  Physiological mechanisms of psychophysical masking: observations from auditory-nerve fibers. , 1990, The Journal of the Acoustical Society of America.

[102]  G. K. Yates,et al.  Chapter 2 – Cochlear Structure and Function , 1995 .

[103]  M Florentine Effects of cochlear impairment and equivalent-threshold masking on psychoacoustic tuning curves. , 1992, Audiology : official organ of the International Society of Audiology.

[104]  B C Moore,et al.  Inter-relationship between different psychoacoustic measures assumed to be related to the cochlear active mechanism. , 1999, The Journal of the Acoustical Society of America.

[105]  M C Teich,et al.  A neural-counting model incorporating refractoriness and spread of excitation. I. Application to intensity discrimination. , 1979, The Journal of the Acoustical Society of America.

[106]  R. Gifford,et al.  Contributions of suppression and excitation to simultaneous masking: effects of signal frequency and masker-signal frequency relation. , 2000, The Journal of the Acoustical Society of America.

[107]  B C Moore,et al.  The temporal course of masking and the auditory filter shape. , 1987, The Journal of the Acoustical Society of America.

[108]  R. FitzHugh A STATISTICAL ANALYZER FOR OPTIC NERVE MESSAGES , 1958, The Journal of general physiology.

[109]  E D Young,et al.  Effects of acoustic trauma on the representation of the vowel "eh" in cat auditory nerve fibers. , 1997, The Journal of the Acoustical Society of America.

[110]  M. Ruggero,et al.  Response to noise of auditory nerve fibers in the squirrel monkey. , 1973, Journal of neurophysiology.

[111]  S Buus,et al.  Psychometric functions for level discrimination. , 1987, The Journal of the Acoustical Society of America.

[112]  M. Liberman,et al.  Afferent and efferent innervation of the cat cochlea: Quantitative analysis with light and electron microscopy , 1990, The Journal of comparative neurology.

[113]  B C Moore,et al.  Auditory filter shapes in subjects with unilateral and bilateral cochlear impairments. , 1986, The Journal of the Acoustical Society of America.

[114]  M. Sachs,et al.  Rate versus level functions for auditory-nerve fibers in cats: tone-burst stimuli. , 1974, The Journal of the Acoustical Society of America.

[115]  Harold F. Schuknecht,et al.  Auditory and Cytocochlear Correlates of Inner Ear Disorders , 1994, Otolaryngology--head and neck surgery : official journal of American Academy of Otolaryngology-Head and Neck Surgery.

[116]  N. Viemeister,et al.  Auditory intensity discrimination at high frequencies in the presence of noise. , 1983, Science.

[117]  G. K. Yates,et al.  Outer hair cell receptor current and sensorineural hearing loss , 1989, Hearing Research.

[118]  H S Colburn,et al.  Theory of binaural interaction based on auditory-nerve data. II. Detection of tones in noise. , 1977, The Journal of the Acoustical Society of America.

[119]  E F Evans,et al.  The sharpening of cochlear frequency selectivity in the normal and abnormal cochlea. , 1975, Audiology : official organ of the International Society of Audiology.

[120]  Julius L. Goldstein,et al.  Relations among compression, suppression, and combination tones in mechanical responses of the basilar membrane: data and MBPNL model , 1995, Hearing Research.

[121]  M. Liberman,et al.  Single-neuron labeling and chronic cochlear pathology. IV. Stereocilia damage and alterations in rate- and phase-level functions , 1984, Hearing Research.

[122]  C W Turner,et al.  Discrimination of spectral-peak amplitude by normal and hearing-impaired subjects. , 1987, The Journal of the Acoustical Society of America.

[123]  M G Heinz,et al.  Detection of time- and bandlimited increments and decrements in a random-level noise. , 1999, The Journal of the Acoustical Society of America.

[124]  R L Jenison,et al.  Correlated cortical populations can enhance sound localization performance. , 2000, The Journal of the Acoustical Society of America.

[125]  R D Hienz,et al.  Vowel discrimination in cats: acquisition, effects of stimulus level, and performance in noise. , 1996, The Journal of the Acoustical Society of America.

[126]  M. Ruggero,et al.  Furosemide alters organ of corti mechanics: evidence for feedback of outer hair cells upon the basilar membrane , 1991, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[127]  C. Daniel Geisler,et al.  Responses of primary auditory fibers to combined noise and tonal stimuli , 1980, Hearing Research.

[128]  W. M. Rabinowitz,et al.  Intensity perception. VI. Summary of recent data on deviations from Weber's law for 1000-Hz tone pulses. , 1976, The Journal of the Acoustical Society of America.

[129]  Brian C. J. Moore,et al.  Mechanisms underlying the frequency discrimination of pulsed tones and the detection of frequency modulation , 1989 .

[130]  So,et al.  An excitation‐pattern model for intensity discrimination , 1981 .

[131]  Brian C. J. Moore,et al.  Overshoot and the ‘‘severe departure’’ from Weber’s law , 1995 .

[132]  B C Moore,et al.  Comparisons of frequency selectivity in simultaneous and forward masking for subjects with unilateral cochlear impairments. , 1986, The Journal of the Acoustical Society of America.

[133]  W. J. McGill,et al.  A study of the near-miss involving Weber’s law and pure-tone intensity discrimination , 1968 .

[134]  Emanuel Parzen,et al.  Stochastic Processes , 1962 .

[135]  B. Moore Psychophysical tuning curves measured in simultaneous and forward masking. , 1978, The Journal of the Acoustical Society of America.

[136]  E. Owens,et al.  An Introduction to the Psychology of Hearing , 1997 .

[137]  S Buus,et al.  Temporal gap detection in sensorineural and simulated hearing impairments. , 1984, Journal of speech and hearing research.

[138]  N I Durlach,et al.  Intensity perception. I. Preliminary theory of intensity resolution. , 1969, The Journal of the Acoustical Society of America.

[139]  Laurel H. Carney,et al.  Spatiotemporal encoding of sound level: Models for normal encoding and recruitment of loudness , 1994, Hearing Research.

[140]  A. Parker,et al.  Sense and the single neuron: probing the physiology of perception. , 1998, Annual review of neuroscience.

[141]  C M Reed,et al.  Intensity perception. XIV. Intensity discrimination in listeners with sensorineural hearing loss. , 1993, The Journal of the Acoustical Society of America.

[142]  L. Carney,et al.  A phenomenological model for the responses of auditory-nerve fibers: I. Nonlinear tuning with compression and suppression. , 2001, The Journal of the Acoustical Society of America.

[143]  B C Moore,et al.  Auditory filter shapes derived in simultaneous and forward masking. , 1981, The Journal of the Acoustical Society of America.

[144]  E. Keithley,et al.  Frequency map of the spiral ganglion in the cat. , 1987, The Journal of the Acoustical Society of America.

[145]  L. A. Westerman,et al.  A diffusion model of the transient response of the cochlear inner hair cell synapse. , 1988, The Journal of the Acoustical Society of America.

[146]  Laurel H. Carney,et al.  Evaluating Auditory Performance Limits: II. One-Parameter Discrimination with Random-Level Variation , 2001, Neural Computation.

[147]  E. de Boer,et al.  Intensity Discrimination of Fluctuating Signals , 1966 .

[148]  M B Sachs,et al.  Strategies for the representation of a tone in background noise in the temporal aspects of the discharge patterns of auditory-nerve fibers. , 1987, The Journal of the Acoustical Society of America.

[149]  L. Carney Sensitivities of cells in anteroventral cochlear nucleus of cat to spatiotemporal discharge patterns across primary afferents. , 1990, Journal of neurophysiology.

[150]  Robert Patuzzi,et al.  Cochlear Micromechanics and Macromechanics , 1996 .

[151]  Richard J. Baker,et al.  Characterising auditory filter nonlinearity , 1994, Hearing Research.

[152]  S Buus Level discrimination of frozen and random noise. , 1990, The Journal of the Acoustical Society of America.

[153]  B. Delgutte,et al.  Neural correlates of the pitch of complex tones. I. Pitch and pitch salience. , 1996, Journal of neurophysiology.

[154]  Bertrand Delgutte,et al.  Two-tone rate suppression in auditory-nerve fibers: Dependence on suppressor frequency and level , 1990, Hearing Research.

[155]  B. Moore,et al.  Benefits of linear amplification and multichannel compression for speech comprehension in backgrounds with spectral and temporal dips. , 1999, The Journal of the Acoustical Society of America.

[156]  A Kohlrausch,et al.  Effect of masker level on overshoot in running- and frozen-noise maskers. , 1994, The Journal of the Acoustical Society of America.

[157]  A. T. Rasmussen Studies of the viiith cranial nerve of man , 1940 .

[158]  Thomas Gold,et al.  Hearing. II. The Physical Basis of the Action of the Cochlea , 1948, Proceedings of the Royal Society of London. Series B - Biological Sciences.

[159]  B. Moore Frequency difference limens for short-duration tones. , 1973, The Journal of the Acoustical Society of America.

[160]  R Probst,et al.  A review of otoacoustic emissions. , 1991, The Journal of the Acoustical Society of America.

[161]  William F. Sewell,et al.  The effects of furosemide on the endocochlear potential and auditory-nerve fiber tuning curves in cats , 1984, Hearing Research.

[162]  M F Dorman,et al.  Minimum spectral contrast for vowel identification by normal-hearing and hearing-impaired listeners. , 1987, The Journal of the Acoustical Society of America.

[163]  C Kaernbach,et al.  Psychophysical evidence against the autocorrelation theory of auditory temporal processing. , 1998, The Journal of the Acoustical Society of America.

[164]  M. Sachs,et al.  Effects of nonlinearities on speech encoding in the auditory nerve. , 1979, The Journal of the Acoustical Society of America.

[165]  N. Viemeister,et al.  Intensity discrimination of noise in the presence of band-reject noise. , 1974, The Journal of the Acoustical Society of America.

[166]  C D Geisler,et al.  The phases of basilar-membrane vibrations. , 1982, The Journal of the Acoustical Society of America.

[167]  Julius L. Goldstein Modeling the nonlinear cochlear mechanical basis of psychophysical tuning. , 1991 .

[168]  N. Viemeister,et al.  Intensity discrimination, increment detection, and magnitude estimation for 1-kHz tones. , 1988, The Journal of the Acoustical Society of America.

[169]  R A Levine,et al.  Auditory-nerve activity in cats with normal and abnormal cochleas. In: Sensorineural hearing loss. , 1970, Ciba Foundation symposium.

[170]  J. W. Horst,et al.  Coding of spectral fine structure in the auditory nerve. II: Level-dependent nonlinear responses. , 1990, The Journal of the Acoustical Society of America.

[171]  D Robertson,et al.  Tuning in the mammalian cochlea. , 1988, Physiological reviews.

[172]  B C Moore,et al.  Auditory filter shapes at 8 and 10 kHz. , 1990, The Journal of the Acoustical Society of America.

[173]  P Dallos,et al.  The level dependence of response phase: observations from cochlear hair cells. , 1998, The Journal of the Acoustical Society of America.

[174]  A R Palmer,et al.  Intensity coding in low-frequency auditory-nerve fibers of the guinea pig. , 1991, The Journal of the Acoustical Society of America.

[175]  Raimond L. Winslow,et al.  Some Aspects of Rate Coding in the Auditory Nerve , 1986 .

[176]  M. Sachs,et al.  Vowel representations in the ventral cochlear nucleus of the cat: effects of level, background noise, and behavioral state. , 1998, Journal of neurophysiology.

[177]  E D Young,et al.  Rate responses of auditory nerve fibers to tones in noise near masked threshold. , 1986, The Journal of the Acoustical Society of America.

[178]  David J. Anderson,et al.  Temporal Position of Discharges in Single Auditory Nerve Fibers within the Cycle of a Sine‐Wave Stimulus: Frequency and Intensity Effects , 1971 .

[179]  M. Florentine,et al.  Level discrimination as a function of level for tones from 0.25 to 16 kHz. , 1987, The Journal of the Acoustical Society of America.

[180]  Bertrand Delgutte,et al.  Peripheral Auditory Processing of Speech Information: Implications from a Physiological Study of Intensity Discrimination , 1987 .

[181]  H. Cramér Mathematical methods of statistics , 1947 .

[182]  I. Russell,et al.  The effect of efferent stimulation on basilar membrane displacement in the basal turn of the guinea pig cochlea , 1996, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[183]  D H Johnson,et al.  Analysis of discharges recorded simultaneously from pairs of auditory nerve fibers. , 1976, Biophysical journal.

[184]  M. Ruggero,et al.  Basilar-membrane responses to clicks at the base of the chinchilla cochlea. , 1998, The Journal of the Acoustical Society of America.

[185]  T. Dau,et al.  A quantitative model of the "effective" signal processing in the auditory system. II. Simulations and measurements. , 1996, The Journal of the Acoustical Society of America.

[186]  D. D. Greenwood A cochlear frequency-position function for several species--29 years later. , 1990, The Journal of the Acoustical Society of America.

[187]  Julius L. Goldstein,et al.  Modeling rapid waveform compression on the basilar membrane as multiple-bandpass-nonlinearity filtering , 1990, Hearing Research.

[188]  B C Moore,et al.  Intensity discrimination: a severe departure from Weber's law. , 1984, The Journal of the Acoustical Society of America.

[189]  D. M. Green,et al.  Frequency discrimination as a function of frequency and sensation level. , 1977, The Journal of the Acoustical Society of America.

[190]  L. Braida,et al.  Towards a model for discrimination of broadband signals. , 1986, The Journal of the Acoustical Society of America.

[191]  A. Oxenham,et al.  Suppression and the upward spread of masking. , 1998, The Journal of the Acoustical Society of America.

[192]  M Florentine,et al.  Level discrimination of tones as a function of duration. , 1986, The Journal of the Acoustical Society of America.

[193]  R A Lutfi,et al.  On the growth of masking asymmetry with stimulus intensity. , 1984, The Journal of the Acoustical Society of America.

[194]  N. Y. S. Kiang,et al.  Discharge Rates of Single Auditory‐Nerve Fibers as Functions of Tone Level , 1969 .

[195]  N. Kiang,et al.  Curious oddments of auditory-nerve studies , 1990, Hearing Research.

[196]  Wolfgang Ellermeier,et al.  A reexamination of the frequency discrimination of random‐amplitude tones, and a test of Henning’s modified energy‐detector model , 1989 .

[197]  N Suga,et al.  Properties of ‘two‐tone inhibition’ in primary auditory neurones , 1971, The Journal of physiology.

[198]  M. Liberman,et al.  Single-neuron labeling and chronic cochlear pathology. III. Stereocilia damage and alterations of threshold tuning curves , 1984, Hearing Research.

[199]  E. F. Evans,et al.  The Dynamic Range Problem: Place and Time Coding at the Level of Cochlear Nerve and Nucleus , 1981 .

[200]  Laurel H. Carney,et al.  Evaluating Auditory Performance Limits: I. One-Parameter Discrimination Using a Computational Model for the Auditory Nerve , 2001, Neural Computation.

[201]  S. Parker,et al.  Intensity discrimination and loudness for tones in notched noise , 1987, Perception & psychophysics.

[202]  T. Houtgast Auditory-filter characteristics derived from direct-masking data and pulsation-threshold data with a rippled-noise masker. , 1977, The Journal of the Acoustical Society of America.

[203]  D M Green,et al.  Two procedures for estimating internal noise. , 1981, The Journal of the Acoustical Society of America.

[204]  M. Sachs,et al.  The representations of the steady-state vowel sound /e/ in the discharge patterns of cat anteroventral cochlear nucleus neurons. , 1990, Journal of neurophysiology.

[205]  Torsten Daub Modeling auditory processing of amplitude modulation II. Spectral and temporal integration , 1997 .

[206]  G. Henning,et al.  Frequency discrimination of random-amplitude tones. , 1965, The Journal of the Acoustical Society of America.

[207]  Brian C. J. Moore,et al.  Formulae describing frequency selectivity as a function of frequency and level, and their use in calculating excitation patterns , 1987, Hearing Research.

[208]  A. Oxenham,et al.  Basilar-membrane nonlinearity and the growth of forward masking. , 1996, The Journal of the Acoustical Society of America.

[209]  S Rosen,et al.  Auditory filter nonlinearity at 2 kHz in normal hearing listeners. , 1998, The Journal of the Acoustical Society of America.

[210]  S. Buus,et al.  Possible Relation of Auditory Nerve Adaptation to Slow Improvement in Level Discrimination with Increasing Duration , 1992 .

[211]  H S Colburn,et al.  Binaural processing of noisy stimuli: internal/external noise ratios for diotic and dichotic stimuli. , 1989, The Journal of the Acoustical Society of America.

[212]  M R Leek,et al.  Reduced frequency selectivity and the preservation of spectral contrast in noise. , 1996, The Journal of the Acoustical Society of America.

[213]  R Plomp,et al.  Auditory handicap of hearing impairment and the limited benefit of hearing aids. , 1978, The Journal of the Acoustical Society of America.

[214]  G. Wilkinson The Theory of Hearing , 1925, Nature.

[215]  Ian C. Bruce,et al.  Improvements of an Auditory-periphery Model to Describe Auditory Nerve Response to Speech Stimuli , 2000 .

[216]  Jont B. Allen,et al.  Acoustics 1992: Otoacoustic emissions , 1993 .

[217]  L. Carney,et al.  A model for the responses of low-frequency auditory-nerve fibers in cat. , 1993, The Journal of the Acoustical Society of America.

[218]  Lionel Collet,et al.  On the role of the olivocochlear bundle in hearing: A case study , 1994, Hearing Research.

[219]  S P Bacon,et al.  Growth of simultaneous masking for fm < fs: effects of overall frequency and level. , 1999, The Journal of the Acoustical Society of America.

[220]  J. Guinan Physiology of Olivocochlear Efferents , 1996 .

[221]  T Dau,et al.  On the role of envelope fluctuation processing in spectral masking. , 2000, The Journal of the Acoustical Society of America.

[222]  A Erell Rate coding model for discrimination of simple tones in the presence of noise. , 1988, The Journal of the Acoustical Society of America.

[223]  B C Moore,et al.  The role of spread excitation and suppression in simultaneous masking. , 1997, The Journal of the Acoustical Society of America.

[224]  Evans Ef,et al.  Cochlear fibre responses in guinea pigs with well defined cochlear lesions. , 1979 .