Andrew J. Oxenham

Revised estimates of human cochlear tuning from otoacoustic and behavioral measurements

We develop an objective, noninvasive method for determining the frequency selectivity of cochlear tuning at low and moderate sound levels. Applicable in humans at frequencies of 1 kHz and above, the method is based on the measurement of stimulus-frequency otoacoustic emissions and, unlike previous noninvasive physiological methods, does not depend on the frequency selectivity of masking or suppression. The otoacoustic measurements indicate that at low sound levels human cochlear tuning is more than twice as sharp as implied by standard behavioral studies and has a different dependence on frequency. New behavioral measurements designed to minimize the influence of nonlinear effects such as suppression agree with the emission-based values. A comparison of cochlear tuning in cat, guinea pig, and human indicates that, contrary to common belief, tuning in the human cochlea is considerably sharper than that found in the other mammals. The sharper tuning may facilitate human speech communication.

Effects of simulated cochlear-implant processing on speech reception in fluctuating maskers

This study investigated the effects of simulated cochlear-implant processing on speech reception in a variety of complex masking situations. Speech recognition was measured as a function of target-to-masker ratio, processing condition (4, 8, 24 channels, and unprocessed) and masker type (speech-shaped noise, amplitude-modulated speech-shaped noise, single male talker, and single female talker). The results showed that simulated implant processing was more detrimental to speech reception in fluctuating interference than in steady-state noise. Performance in the 24-channel processing condition was substantially poorer than in the unprocessed condition, despite the comparable representation of the spectral envelope. The detrimental effects of simulated implant processing in fluctuating maskers, even with large numbers of channels, may be due to the reduction in the pitch cues used in sound source segregation, which are normally carried by the peripherally resolved low-frequency harmonics and the temporal fine structure. The results suggest that using steady-state noise to test speech intelligibility may underestimate the difficulties experienced by cochlear-implant users in fluctuating acoustic backgrounds.

A behavioral measure of basilar-membrane nonlinearity in listeners with normal and impaired hearing

This paper examines the possibility of estimating basilar-membrane (BM) nonlinearity using a psychophysical technique. The level of a forward masker required to mask a brief signal was measured for conditions where the masker was either at, or one octave below, the signal frequency. The level of the forward masker at masked threshold provided an indirect measure of the BM response to the signal, as follows. Consistent with physiological studies, it was assumed that the BM responds linearly to frequencies well below the characteristic frequency (CF). Thus the ratio of the slopes of the masking functions between a masker at the signal frequency and a masker well below the signal frequency should provide an estimate of BM compression at CF. Results obtained from normally hearing listeners were in quantitative agreement with physiological estimates of BM compression. Furthermore, differences between normally hearing listeners and listeners with cochlear hearing impairment were consistent with the physiological effects of damage to the cochlea. The results support the hypothesis that BM nonlinearity governs the nonlinear growth of the upward spread of masking, and suggest that this technique provides a straightforward method for estimating BM nonlinearity in humans.

A Neural Representation of Pitch Salience in Nonprimary Human Auditory Cortex Revealed with Functional Magnetic Resonance Imaging

Pitch, one of the primary auditory percepts, is related to the temporal regularity or periodicity of a sound. Previous functional brain imaging work in humans has shown that the level of population neural activity in centers throughout the auditory system is related to the temporal regularity of a sound, suggesting a possible relationship to pitch. In the current study, functional magnetic resonance imaging was used to measure activation in response to harmonic tone complexes whose temporal regularity was identical, but whose pitch salience (or perceptual pitch strength) differed, across conditions. Cochlear nucleus, inferior colliculus, and primary auditory cortex did not show significant differences in activation level between conditions. Instead, a correlate of pitch salience was found in the neural activity levels of a small, spatially localized region of nonprimary auditory cortex, overlapping the anterolateral end of Heschls gyrus. The present data contribute to converging evidence that anterior areas of nonprimary auditory cortex play an important role in processing pitch.

Basilar-membrane nonlinearity and the growth of forward masking

Forward masking growth functions were measured for pure-tone maskers and signals at 2 and 6 kHz as a function of the silent interval between the masker and signal. The inclusion of conditions involving short signals and short masker-signal intervals ensured that a wide range of signal thresholds were recorded. A consistent pattern was seen across all the results. When the signal level was below about 35 dB SPL the growth of masking was shallow, so that signal threshold increased at a much slower rate than masker level. When the signal level exceeded this value, the masking function steepened, approaching unity (linear growth) at the highest masker and signal levels. The results are inconsistent with an explanation for forward-masking growth in terms of saturating neural adaptation. Instead the data are well described by a model incorporating a simulation of the basilar-membrane response at characteristic frequency (which is almost linear at low levels and compressive at higher levels) followed by a sliding intensity integrator or temporal window. Taken together with previous results, the findings suggest that the principle nonlinearity in temporal masking may be the basilar membrane response function, and that subsequent to this the auditory system behaves as if it were linear in the intensity domain.

Pitch discrimination of diotic and dichotic tone complexes: Harmonic resolvability or harmonic number?

Three experiments investigated the relationship between harmonic number, harmonic resolvability, and the perception of harmonic complexes. Complexes with successive equal-amplitude sine- or random-phase harmonic components of a 100- or 200-Hz fundamental frequency (f0) were presented dichotically, with even and odd components to opposite ears, or diotically, with all harmonics presented to both ears. Experiment 1 measured performance in discriminating a 3.5%-5% frequency difference between a component of a harmonic complex and a pure tone in isolation. Listeners achieved at least 75% correct for approximately the first 10 and 20 individual harmonics in the diotic and dichotic conditions, respectively, verifying that only processes before the binaural combination of information limit frequency selectivity. Experiment 2 measured fundamental frequency difference limens (f0 DLs) as a function of the average lowest harmonic number. Similar results at both f0s provide further evidence that harmonic number, not absolute frequency, underlies the order-of-magnitude increase observed in f0 DLs when only harmonics above about the 10th are presented. Similar results under diotic and dichotic conditions indicate that the auditory system, in performing f0 discrimination, is unable to utilize the additional peripherally resolved harmonics in the dichotic case. In experiment 3, dichotic complexes containing harmonics below the 12th, or only above the 15th, elicited pitches of the f0 and twice the f0, respectively. Together, experiments 2 and 3 suggest that harmonic number, regardless of peripheral resolvability, governs the transition between two different pitch percepts, one based on the frequencies of individual resolved harmonics and the other based on the periodicity of the temporal envelope.

Estimates of Human Cochlear Tuning at Low Levels Using Forward and Simultaneous Masking

Auditory filter shapes were derived from psychophysical measurements in eight normal-hearing listeners using a variant of the notched-noise method for brief signals in forward and simultaneous masking. Signal frequencies of 1, 2, 4, 6, and 8 kHz were tested. The signal level was fixed at 10 dB above absolute threshold in the forward-masking conditions and fixed at either 10 or 35 dB above absolute threshold in the simultaneous-masking conditions. The results show that filter equivalent rectangular bandwidths (ERBs) are substantially narrower in forward masking than has been found in previous studies using simultaneous masking. Furthermore, in contrast to earlier studies, the sharpness of tuning doubles over the range of frequencies tested, giving QERB values of about 10 and 20 at signal frequencies of 1 and 8 kHz, respectively. It is argued that the new estimates of auditory filter bandwidth provide a more accurate estimate of human cochlear tuning at low levels than earlier estimates using simultaneous masking at higher levels, and that they are therefore more suitable for comparison to cochlear tuning data from other species. The data may also prove helpful in defining the parameters for nonlinear models of human cochlear processing.

Otoacoustic Estimation of Cochlear Tuning: Validation in the Chinchilla

We analyze published auditory-nerve and otoacoustic measurements in chinchilla to test a network of hypothesized relationships between cochlear tuning, cochlear traveling-wave delay, and stimulus-frequency otoacoustic emissions (SFOAEs). We find that the physiological data generally corroborate the network of relationships, including predictions from filter theory and the coherent-reflection model of OAE generation, at locations throughout the cochlea. The results support the use of otoacoustic emissions as noninvasive probes of cochlear tuning. Developing this application, we find that tuning ratios—defined as the ratio of tuning sharpness to SFOAE phase-gradient delay in periods—have a nearly species-invariant form in cat, guinea pig, and chinchilla. Analysis of the tuning ratios identifies a species-dependent parameter that locates a transition between “apical-like” and “basal-like” behavior involving multiple aspects of cochlear physiology. Approximate invariance of the tuning ratio allows determination of cochlear tuning from SFOAE delays. We quantify the procedure and show that otoacoustic estimates of chinchilla cochlear tuning match direct measures obtained from the auditory nerve. By assuming that invariance of the tuning ratio extends to humans, we derive new otoacoustic estimates of human cochlear tuning that remain mutually consistent with independent behavioral measurements obtained using different rationales, methodologies, and analysis procedures. The results confirm that at any given characteristic frequency (CF) human cochlear tuning appears sharper than that in the other animals studied, but varies similarly with CF. We show, however, that the exceptionality of human tuning can be exaggerated by the ways in which species are conventionally compared, which take no account of evident differences between the base and apex of the cochlea. Finally, our estimates of human tuning suggest that the spatial spread of excitation of a pure tone along the human basilar membrane is comparable to that in other common laboratory animals.

Forward masking: Adaptation or integration?

The aim of this study was to attempt to distinguish between neural adaptation and persistence (or temporal integration) as possible explanations of forward masking. Thresholds were measured for a sinusoidal signal as a function of signal duration for conditions where the delay between the masker offset and the signal offset (the offset-offset interval) was fixed. The masker was a 200-ms broadband noise, presented at a spectrum level of 40 dB (re: 20 microPa), and the signal was a 4-kHz sinusoid, gated with 2-ms ramps. The offset-offset interval was fixed at various durations between 4 and 102 ms and signal thresholds were measured for a range of signal durations at each interval. A substantial decrease in thresholds was observed with increasing duration for signal durations up to about 20 ms. At short offset-offset intervals, the amount of temporal integration exceeded that normally found in quiet. The results were simulated using models of temporal integration (the temporal-window model) and adaptation. For both models, the inclusion of a peripheral nonlinearity, similar to that observed physiologically in studies of the basilar membrane, was essential in producing a good fit to the data. Both models were about equally successful in accounting for the present data. However, the temporal-window model provided a somewhat better account of similar data from a simultaneous-masking experiment, using the same parameters. This suggests that the linear, time-invariant properties of the temporal-window approach are appropriate for modeling forward masking. Overall the results confirm that forward masking can be described in terms of peripheral nonlinearity followed by linear temporal integration at higher levels in the auditory system. However, the difference in predictions between the adaptation and integration models is relatively small, meaning that influence of adaptation cannot be ruled out.

Cochlear compression: perceptual measures and implications for normal and impaired hearing.

This article provides a review of recent developments in our understanding of how cochlear nonlinearity affects sound perception and how a loss of the nonlinearity associated with cochlear hearing impairment changes the way sounds are perceived. The response of the healthy mammalian basilar membrane (BM) to sound is sharply tuned, highly nonlinear, and compressive. Damage to the outer hair cells (OHCs) results in changes to all three attributes: in the case of total OHC loss, the response of the BM becomes broadly tuned and linear. Many of the differences in auditory perception and performance between normal-hearing and hearing-impaired listeners can be explained in terms of these changes in BM response. Effects that can be accounted for in this way include poorer audiometric thresholds, loudness recruitment, reduced frequency selectivity, and changes in apparent temporal processing. All these effects can influence the ability of hearing-impaired listeners to perceive speech, especially in complex acoustic backgrounds. A number of behavioral methods have been proposed to estimate cochlear nonlinearity in individual listeners. By separating the effects of cochlear nonlinearity from other aspects of hearing impairment, such methods may contribute towards identifying the different physiological mechanisms responsible for hearing loss in individual patients. This in turn may lead to more accurate diagnoses and more effective hearing-aid fitting for individual patients. A remaining challenge is to devise a behavioral measure that is sufficiently accurate and efficient to be used in a clinical setting.