Effects of contralateral noise on envelope-following responses, auditory-nerve compound action potentials, and otoacoustic emissions measured simultaneously.

This study assessed whether the effects of contralateral acoustic stimulation (CAS) are consistent with eliciting the medial olivocochlear (MOC) reflex for measurements sensitive to outer hair cell (otoacoustic emissions, OAEs), auditory-nerve (AN; compound action potential, CAP), and brainstem/cortical (envelope-following response, EFR) function. The effects of CAS were evaluated for simultaneous measurement of OAEs, CAPs, and EFRs in participants with normal hearing. Clicks were presented at 40 or 98 Hz in three ipsilateral noise conditions (no noise, 45 dB SPL, and 55 dB SPL). For the no noise condition, CAS suppressed or enhanced EFR amplitudes for 40- and 98-Hz clicks, respectively, while CAS had no significant effect on CAP amplitudes. A follow-up experiment using slower rates (4.4-22.2 Hz) assessed whether this insignificant CAS effect on CAPs was from ipsilateral MOC stimulation or AN adaptation; however, CAS effects remained insignificant despite favorable signal-to-noise ratios. CAS-related enhancements of EFR and CAP amplitudes in ipsilateral noise were not observed, contrary to the anti-masking effect of the MOC reflex. EFR and OAE suppression from CAS were not significantly correlated. Thus, the effects of CAS on EFRs may not be solely mediated by the MOC reflex and may be partially mediated by higher auditory centers.

Subcortical auditory model including efferent dynamic gain control with inputs from cochlear nucleus and inferior colliculus.

An auditory model has been developed with a time-varying, gain-control signal based on the physiology of the efferent system and subcortical neural pathways. The medial olivocochlear (MOC) efferent stage of the model receives excitatory projections from fluctuation-sensitive model neurons of the inferior colliculus (IC) and wide-dynamic-range model neurons of the cochlear nucleus. The response of the model MOC stage dynamically controls cochlear gain via simulated outer hair cells. In response to amplitude-modulated (AM) noise, firing rates of most IC neurons with band-enhanced modulation transfer functions in awake rabbits increase over a time course consistent with the dynamics of the MOC efferent feedback. These changes in the rates of IC neurons in awake rabbits were employed to adjust the parameters of the efferent stage of the proposed model. Responses of the proposed model to AM noise were able to simulate the increasing IC rate over time, whereas the model without the efferent system did not show this trend. The proposed model with efferent gain control provides a powerful tool for testing hypotheses, shedding insight on mechanisms in hearing, specifically those involving the efferent system.

Computational modeling of the human compound action potential.

The auditory nerve (AN) compound action potential (CAP) is an important tool for assessing auditory disorders and monitoring the health of the auditory periphery during surgical procedures. The CAP has been mathematically conceptualized as the convolution of a unit response (UR) waveform with the firing rate of a population of AN fibers. Here, an approach for predicting experimentally recorded CAPs in humans is proposed, which involves the use of human-based computational models to simulate AN activity. CAPs elicited by clicks, chirps, and amplitude-modulated carriers were simulated and compared with empirically recorded CAPs from human subjects. In addition, narrowband CAPs derived from noise-masked clicks and tone bursts were simulated. Many morphological, temporal, and spectral aspects of human CAPs were captured by the simulations for all stimuli tested. These findings support the use of model simulations of the human CAP to refine existing human-based models of the auditory periphery, aid in the design and analysis of auditory experiments, and predict the effects of hearing loss, synaptopathy, and other auditory disorders on the human CAP.

Middle ear muscle and medial olivocochlear activity inferred from individual human ears via cochlear potentials.

The peripheral auditory system is influenced by the medial olivocochlear (MOC) and middle ear muscle (MEM) reflexes. When elicited by contralateral acoustic stimulation (CAS), these reflexes reduce cochlear amplification (MOC reflex) and limit low-frequency transmission through the middle ear (MEM reflex). The independent roles of these reflexes on auditory physiology and perception are difficult to distinguish. The amplitude of the cochlear microphonic (CM) is expected to increase or decrease when the MOC and MEM reflexes are elicited by CAS, respectively, which could lead to a straightforward interpretation of what reflex is dominant for a given CAS level. CM and ear canal sound pressure level (SPL) were measured for a 500 Hz, 90 dB SPL probe in the presence of contralateral broadband noise (CBBN) for levels ranging from 45-75 dB SPL. In most subjects, CM amplitude increased for CBBN levels of 45 and 55 dB SPL, while no change in ear canal SPL was observed, consistent with eliciting the MOC reflex. Conversely, CM amplitude decreased, and ear canal SPL increased in the presence of 65 and 75 dB SPL CBBN, consistent with eliciting the MEM reflex. A CM-based test of the MOC reflex may facilitate detection of MEM effects and the assessment of adults with cochlear hearing loss.

Minimum Detectable Differences in Electrocochleography Measurements: Bayesian-Based Predictions.

Physiology of the cochlea and auditory nerve can be assessed with electrocochleography (ECochG), a technique that involves measuring auditory evoked potentials from an electrode placed near or within the cochlea. Research, clinical, and operating room applications of ECochG have in part centered on measuring the auditory nerve compound action potential (AP) amplitude, the summating potential (SP) amplitude, and the ratio of the two (SP/AP). Despite the common use of ECochG, the variability of repeated amplitude measurements for individuals and groups is not well understood. We analyzed ECochG measurements made with a tympanic membrane electrode in a group of younger normal-hearing participants to characterize the within-participant and group-level variability for the AP amplitude, SP amplitude, and SP/AP amplitude ratio. Results show that the measurements have substantial variability and that, especially with smaller sample sizes, significant reduction in variability can be obtained by averaging measurements across repeated electrode placements within subjects. Using a Bayesian-based model of the data, we generated simulated data to predict minimum detectable differences in AP and SP amplitudes for experiments with a given number of participants and repeated measurements. Our findings provide evidence-based recommendations for the design and sample size determination of future experiments using ECochG amplitude measurements, and the evaluation of previous publications in terms of sensitivity to detecting experimental effects on ECochG amplitude measurements. Accounting for the variability of ECochG measurements should result in more consistent results in the clinical and basic assessments of hearing and hearing loss, either hidden or overt.

Temporal Envelope Coding of the Human Auditory Nerve Inferred from Electrocochleography: Comparison with Envelope Following Responses.

Neural coding of the slow amplitude fluctuations of sound (i.e., temporal envelope) is thought to be essential for speech understanding; however, such coding by the human auditory nerve is poorly understood. Here, neural coding of the temporal envelope by the human auditory nerve is inferred from measurements of the compound action potential in response to an amplitude modulated carrier (CAPENV) for modulation frequencies ranging from 20 to 1000 Hz. The envelope following response (EFR) was measured simultaneously with CAPENV from active electrodes placed on the high forehead and tympanic membrane, respectively. Results support the hypothesis that phase locking to higher modulation frequencies (> 80 Hz) will be stronger for CAPENV, compared to EFR, consistent with the upper-frequency limits of phase locking for auditory nerve fibers compared to auditory brainstem/cortex neurons. Future work is needed to determine the extent to which (1) CAPENV is a useful tool for studying how temporal processing of the auditory nerve is affected by aging, hearing loss, and noise-induced cochlear synaptopathy and (2) CAPENV reveals the relationship between auditory nerve temporal processing and perception of the temporal envelope.

Firing Rate Adaptation of the Human Auditory Nerve Optimizes Neural Signal-to-Noise Ratios.

Several physiological mechanisms act on the response of the auditory nerve (AN) during acoustic stimulation, resulting in an adjustment in auditory gain. These mechanisms include-but are not limited to-firing rate adaptation, dynamic range adaptation, the middle ear muscle reflex, and the medial olivocochlear reflex. A potential role of these mechanisms is to improve the neural signal-to-noise ratio (SNR) at the output of the AN in real time. This study tested the hypothesis that neural SNRs, inferred from non-invasive assessment of the human AN, improve over the duration of acoustic stimulation. Cochlear potentials were measured in response to a series of six high-level clicks embedded in a series of six lower-level broadband noise bursts. This paradigm elicited a compound action potential (CAP) in response to each click and to the onset of each noise burst. The ratio of CAP amplitudes elicited by each click and noise burst pair (i.e., neural SNR) was tracked over the six click/noise bursts. The main finding was a rapid (< 24 ms) increase in neural SNR from the first to the second click/noise burst, consistent with a real-time adjustment in the response of the auditory periphery toward improving the SNR of the signal transmitted to the brainstem. Analysis of cochlear microphonic and ear canal sound pressure recordings, as well as the time course for this improvement in neural SNR, supports the conclusion that firing rate adaptation is likely the primary mechanism responsible for improving neural SNR, while dynamic range adaptation, the middle ear muscle reflex, and the medial olivocochlear reflex played a secondary role on the effects observed in this study. Real-time improvements in neural SNR are significant because they may be essential for robust encoding of speech and other relevant stimuli in the presence of background noise.

Perceived Anger in Clear and Conversational Speech: Contributions of Age and Hearing Loss.

A previous investigation demonstrated differences between younger adult normal-hearing listeners and older adult hearing-impaired listeners in the perceived emotion of clear and conversational speech. Specifically, clear speech sounded angry more often than conversational speech for both groups, but the effect was smaller for the older listeners. These listener groups differed by two confounding factors, age (younger vs. older adults) and hearing status (normal vs. impaired). The objective of the present study was to evaluate the contributions of aging and hearing loss to the reduced perception of anger in older adults with hearing loss. We investigated perceived anger in clear and conversational speech in younger adults with and without a simulated age-related hearing loss, and in older adults with normal hearing. Younger adults with simulated hearing loss performed similarly to normal-hearing peers, while normal-hearing older adults performed similarly to hearing-impaired peers, suggesting that aging was the primary contributor to the decreased anger perception seen in previous work. These findings confirm reduced anger perception for older adults compared to younger adults, though the significant speaking style effect-regardless of age and hearing status-highlights the need to identify methods of producing clear speech that is emotionally neutral or positive.

The role of the medial olivocochlear reflex in psychophysical masking and intensity resolution in humans: a review.

This review addresses the putative role of the medial olivocochlear (MOC) reflex in psychophysical masking and intensity resolution in humans. A framework for interpreting psychophysical results in terms of the expected influence of the MOC reflex is introduced. This framework is used to review the effects of a precursor or contralateral acoustic stimulation on 1) simultaneous masking of brief tones, 2) behavioral estimates of cochlear gain and frequency resolution in forward masking, 3) the buildup and decay of forward masking, and 4) measures of intensity resolution. Support, or lack thereof, for a role of the MOC reflex in psychophysical perception is discussed in terms of studies on estimates of MOC strength from otoacoustic emissions and the effects of resection of the olivocochlear bundle in patients with vestibular neurectomy. Novel, innovative approaches are needed to resolve the dissatisfying conclusion that current results are unable to definitively confirm or refute the role of the MOC reflex in masking and intensity resolution.

Masking of short tones in noise: Evidence for envelope-based, rather than energy-based detection.

The "temporal effect" in simultaneous masking may be characterized by better probe detection thresholds for a short, tonal probe presented at the temporal center of a masker compared to at the onset of a masker. Energy-based models of masking have been used to interpret the temporal effect as evidence that the gain of the auditory system decreases during acoustic stimulation. This study shows that masking from temporal-envelope fluctuations of a precursor or from a temporal gap between stimuli violates the assumptions of energy-based models and complicates the interpretation of temporal effects in terms of a reduction in gain. Detection thresholds were measured for a 6-ms, 4000-Hz probe preceded by a narrowband precursor and presented 2-, 197-, or 392-ms after the onset of a narrowband masker. The delay between the precursor offset and masker onset ranged from -2 to 250 ms. Probe thresholds were elevated in the presence of precursors with fluctuating compared to flattened temporal envelopes and when a temporal gap was inserted between the precursor and masker. The results suggest that the interpretation and design of temporal-effect studies should consider the masking effects of temporal-envelope fluctuations. These findings are consistent with speech-perception experiments that show masking from temporal-envelope fluctuations.

Techniques for Obtaining High-quality Recordings in Electrocochleography.

There are several technical challenges to obtaining high-quality recordings of cochlear potentials in human electrocochleography (ECochG). These challenges include electrical artifacts from devices such as acoustic transducers, biological artifacts from excessive myogenic and electroencephalographic potentials, and issues associated with the placement of a tympanic membrane (TM) electrode on the eardrum. This article presents approaches for dealing with these challenges for ECochG measurement using a TM electrode. Emphasis is placed on eliminating stimulus artifact, optimizing the placement of the electrode, and comparing a custom-made electrode with a commercially-available electrode. This comparison revealed that the custom-made electrode results in greater subject comfort, superior ease of placing the electrode on the eardrum, and larger compound action potential (CAP) amplitudes.

Effects of Masker Envelope Fluctuations on the Temporal Effect.

Under certain conditions, detection thresholds in simultaneous masking improve when the onset of a short sinusoidal probe is delayed from the onset of a long masker. This improvement, known as the temporal effect, is largest for broadband maskers and is smaller or absent for narrowband maskers centered on the probe frequency. This study tests the hypothesis that small or absent temporal effects for narrowband maskers are due to the inherent temporal envelope fluctuations of Gaussian noise. Temporal effects were measured for narrowband noise maskers with fluctuating ("fluctuating maskers") and flattened ("flattened maskers") temporal envelopes as a function of masker level (Exp. I) and in the presence of fluctuating and flattened precursors (Exp. II). The temporal effect was absent for fluctuating narrowband maskers and as large as ~ 7 dB for flattened narrowband maskers. The AC-coupled power of the temporal envelopes of precursors and maskers accounted for 94 % of the variance in probe detection thresholds measured with fluctuating and flattened precursors and maskers. These results suggest that masker temporal envelope fluctuations contribute to the temporal effect and should be considered in future modeling efforts.

Amplitude modulation detection with a short-duration carrier: Effects of a precursor and hearing loss.

This study tests the hypothesis that amplitude modulation (AM) detection will be better under conditions where basilar membrane (BM) response growth is expected to be linear rather than compressive. This hypothesis was tested by (1) comparing AM detection for a tonal carrier as a function of carrier level for subjects with and without cochlear hearing impairment (HI), and by (2) comparing AM detection for carriers presented with and without an ipsilateral notched-noise precursor, under the assumption that the precursor linearizes BM responses. Average AM detection thresholds were approximately 5 dB better for subjects with HI than for subjects with normal hearing (NH) at moderate-level carriers. Average AM detection for low-to-moderate level carriers was approximately 2 dB better with the precursor than without the precursor for subjects with NH, whereas precursor effects were absent or smaller for subjects with HI. Although effect sizes were small and individual differences were noted, group differences are consistent with better AM detection for conditions where BM responses are less compressive due to cochlear hearing loss or due to a reduction in cochlear gain. These findings suggest the auditory system may quickly adjust to the local soundscape to increase effective AM depth and improve signal-to-noise ratios.

Notched-noise precursors improve detection of low-frequency amplitude modulation.

Amplitude modulation (AM) detection was measured with a short (50 ms), high-frequency carrier as a function of carrier level (Experiment I) and modulation frequency (Experiment II) for conditions with or without a notched-noise precursor. A longer carrier (500 ms) was also included in Experiment I. When the carrier was preceded by silence (no precursor condition) AM detection thresholds worsened for moderate-level carriers compared to lower- or higher-level carriers, resulting in a "mid-level hump." AM detection thresholds with a precursor were better than those without a precursor, primarily for moderate-to-high level carriers, thus eliminating the mid-level hump in AM detection. When the carrier was 500 ms, AM thresholds improved by a constant (across all levels) relative to AM thresholds with a precursor, consistent with the longer carrier providing more "looks" to detect the AM signal. Experiment II revealed that improved AM detection with compared to without a precursor is limited to low-modulation frequencies (<60 Hz). These results are consistent with (1) a reduction in cochlear gain over the course of the precursor perhaps via the medial olivocochlear reflex or (2) a form of perceptual enhancement which may be mediated by adaptation of inhibition.

Effects of age and hearing loss on overshoot.

The detection of a brief, sinusoidal probe in a long broadband, simultaneous masker improves as the probe is delayed from the masker's onset. This improvement ("overshoot") may be mediated by a reduction in cochlear amplifier gain over the timecourse of the masker via the medial olivocochlear (MOC) reflex. Overshoot was measured in younger adults with normal hearing and in older adults with normal and impaired hearing to test the hypothesis that aging and cochlear hearing loss result in abnormal overshoot, consistent with changes in certain structures along the MOC pathway. Overshoot decreased with increasing quiet probe thresholds and was only minimally influenced by increasing age. Marked individual differences in overshoot were observed due to differences in masking thresholds for probes presented near the masker's onset. Model simulations support the interpretation that reduced overshoot in hearing-impaired listeners is due to limited cochlear amplifier gain and therefore less gain to adjust over the timecourse of the masker. Similar overshoot among younger and older adults with normal hearing suggests that age-related changes to mechanisms underlying overshoot do not result in significant differences in overshoot among younger and older adults with normal hearing.

PsyAcoustX: A flexible MATLAB(®) package for psychoacoustics research.

The demands of modern psychophysical studies require precise stimulus delivery and flexible platforms for experimental control. Here, we describe PsyAcoustX, a new, freely available suite of software tools written in the MATLAB(®) environment to conduct psychoacoustics research on a standard PC. PsyAcoustX provides a flexible platform to generate and present auditory stimuli in real time and record users' behavioral responses. Data are automatically logged by stimulus condition and aggregated in an exported spreadsheet for offline analysis. Detection thresholds can be measured adaptively under basic and complex auditory masking tasks and other paradigms (e.g., amplitude modulation detection) within minutes. The flexibility of the module offers experimenters access to nearly every conceivable combination of stimulus parameters (e.g., probe-masker relations). Example behavioral applications are highlighted including the measurement of audiometric thresholds, basic simultaneous and non-simultaneous (i.e., forward and backward) masking paradigms, gap detection, and amplitude modulation detection. Examples of these measurements are provided including the psychoacoustic phenomena of temporal overshoot, psychophysical tuning curves, and temporal modulation transfer functions. Importantly, the core design of PsyAcoustX is easily modifiable, allowing users the ability to adapt its basic structure and create additional modules for measuring discrimination/detection thresholds for other auditory attributes (e.g., pitch, intensity, etc.) or binaural paradigms.

Computational modeling of individual differences in behavioral estimates of cochlear nonlinearities.

Temporal masking curves (TMCs) are often used to estimate cochlear compression in individuals with normal and impaired hearing. These estimates may yield a wide range of individual differences, even among subjects with similar quiet thresholds. This study used an auditory model to assess potential sources of variance in TMCs from 51 listeners in Poling et al. [J Assoc Res Otolaryngol, 13:91-108 (2012)]. These sources included threshold elevation, the contribution of outer and inner hair cell dysfunction to threshold elevation, compression of the off-frequency linear reference, and detection efficiency. Simulations suggest that detection efficiency is a primary factor contributing to individual differences in TMCs measured in normal-hearing subjects, while threshold elevation and the contribution of outer and inner hair cell dysfunction are primary factors in hearing-impaired subjects. Approximating the most compressive growth rate of the cochlear response from TMCs was achieved only in subjects with the highest detection efficiency. Simulations included off-frequency nonlinearity in basilar membrane and inner hair cell processing; however, this nonlinearity did not improve predictions, suggesting that other sources, such as the decay of masking and the strength of the medial olivocochlear reflex, may mimic off-frequency nonlinearity. Findings from this study suggest that sources of individual differences can play a strong role in behavioral estimates of compression, and these sources should be considered when using forward masking to study cochlear function in individual listeners or across groups of listeners.

Psychophysical auditory filter estimates reveal sharper cochlear tuning in musicians.

Musicianship confers enhancements to hearing at nearly all levels of the auditory system from periphery to percept. Musicians' superior psychophysical abilities are particularly evident in spectral discrimination and noise-degraded listening tasks, achieving higher perceptual sensitivity than their nonmusician peers. Greater spectral acuity implies that musicianship may increase auditory filter selectivity. This hypothesis was directly tested by measuring both forward- and simultaneous-masked psychophysical tuning curves. Sharper filter tuning (i.e., higher Q10) was observed in musicians compared to nonmusicians. Findings suggest musicians' pervasive listening benefits may be facilitated, in part, by superior spectral processing/decomposition as early as the auditory periphery.

Evaluating the effects of olivocochlear feedback on psychophysical measures of frequency selectivity.

Frequency selectivity was evaluated under two conditions designed to assess the influence of a "precursor" stimulus on auditory filter bandwidths. The standard condition consisted of a short masker, immediately followed by a short signal. The precursor condition was identical except a 100-ms sinusoid at the signal frequency (i.e., the precursor) was presented before the masker. The standard and precursor conditions were compared for measurements of psychophysical tuning curves (PTCs), and notched noise tuning characteristics. Estimates of frequency selectivity were significantly broader in the precursor condition. In the second experiment, PTCs in the standard and precursor conditions were simulated to evaluate the influence of the precursor on PTC bandwidth. The model was designed to account for the influence of additivity of masking between the masker and precursor. Model simulations were able to qualitatively account for the perceptual data when outer hair cell gain of the model was reduced in the precursor condition. These findings suggest that the precursor may have reduced cochlear gain, in addition to producing additivity of masking. This reduction in gain may be mediated by the medial olivocochlear reflex.

Auditory filter tuning inferred with short sinusoidal and notched-noise maskers.

The physiology of the medial olivocochlear reflex suggests that a sufficiently long stimulus (>100 ms) may reduce cochlear gain and result in broadened frequency selectivity. The current study attempted to avoid gain reduction by using short maskers (20 ms) to measure psychophysical tuning curves (PTCs) and notched-noise tuning characteristics, with a 4-kHz signal. The influence of off-frequency listening on PTCs was evaluated using two types of background noise. Iso-level curves were derived using an estimate of the cochlear input/output (I/O) function, which was obtained using an off-frequency masker as a linear reference. The influence of masker duration on PTCs was assessed using a model that assumed long maskers (>20 ms) evoked gain reduction. The results suggested that the off-frequency masker was a valid linear reference when deriving I/O functions and that off-frequency listening may have occurred in auditory filters apical to the signal place. The iso-level curves from this growth-of-masking study were consistent with those from a temporal-masking-curve study by Eustaquio-Martin and Lopez-Poveda [J. Assoc. Res. Otolaryngol. 12, 281-299. (2011)], suggesting that either approach may be used to derive iso-level curves. Finally, model simulations suggested that masker duration may not influence estimates of frequency selectivity.