Cochlear filtering results in earlier responses to high than to low frequencies. This study examined potential perceptual correlates of cochlear delays by measuring the perception of relative timing between tones of different frequencies. A brief 250-Hz tone was combined with a brief 1-, 2-, 4-, or 6-kHz tone. Two experiments were performed, one involving subjective judgments of perceived synchrony, the other involving asynchrony detection and discrimination. The functions relating the proportion of "synchronous" responses to the delay between the tones were similar for all tone pairs. Perceived synchrony was maximal when the tones in a pair were gated synchronously. The perceived-synchrony function slopes were asymmetric, being steeper on the low-frequency-leading side. In the second experiment, asynchrony-detection thresholds were lower for low-frequency rather than for high-frequency leading pairs. In contrast with previous studies, but consistent with the first experiment, thresholds did not depend on frequency separation between the tones, perhaps because of the elimination of within-channel cues. The results of the two experiments were related quantitatively using a decision-theoretic model, and were found to be highly correlated. Overall the results suggest that frequency-dependent cochlear group delays are compensated for at higher processing stages, resulting in veridical perception of timing relationships across frequency.

Chronic tinnitus is a prevalent hearing disorder, and yet no successful treatments or objective diagnostic tests are currently available. The aim of this study was to investigate the relationship between the presence of tinnitus and the strength of the middle-ear-muscle reflex (MEMR) in humans with normal and near-normal hearing. Clicks were used as test stimuli to obtain a wideband measure of the effect of reflex activation on ear-canal sound pressure. The reflex was elicited using a contralateral broadband noise. The results show that the reflex strength is significantly reduced in individuals with noise-induced continuous tinnitus and normal or near-normal audiometric thresholds compared with no-tinnitus controls. Due to a shallower growth of the reflex strength in the tinnitus group, the difference between the two groups increased with increasing elicitor level. No significant difference in the effect of tinnitus on the strength of the middle-ear muscle reflex was found between males and females. The weaker reflex could not be accounted for by differences in audiometric hearing thresholds between the tinnitus and control groups. Similarity between our findings in humans and the findings of a reduced middle-ear muscle reflex in noise-exposed animals suggests that noise-induced tinnitus in individuals with clinically normal hearing may be a consequence of cochlear synaptopathy, a loss of synaptic connections between inner hair cells (IHCs) in the cochlea and auditory-nerve (AN) fibers that has been termed hidden hearing loss.

Attention to a target stimulus within a complex scene often results in enhanced cortical representations of the target relative to the background. It remains unclear where along the auditory pathways attentional effects can first be measured. Anatomy suggests that attentional modulation could occur through corticofugal connections extending as far as the cochlea itself. Earlier attempts to investigate the effects of attention on human cochlear processing have revealed small and inconsistent effects. In this study, stimulus-frequency otoacoustic emissions were recorded from a total of 30 human participants as they performed tasks that required sustained selective attention to auditory or visual stimuli. In the first sample of 15 participants, emission magnitudes were significantly weaker when participants attended to the visual stimuli than when they attended to the auditory stimuli, by an average of 5.4 dB. However, no such effect was found in the second sample of 15 participants. When the data were pooled across samples, the average attentional effect was significant, but small (2.48 dB), with 12 of 30 listeners showing a significant effect, based on bootstrap analysis of the individual data. The results highlight the need for considering sources of individual differences and using large sample sizes in future investigations. V

Several studies have postulated that psychoacoustic measures of auditory perception are influenced by efferent-induced changes in cochlear responses, but these postulations have generally remained untested. This study measured the effect of stimulus phase curvature and temporal envelope modulation on the medial olivocochlear reflex (MOCR) and on the middle-ear muscle reflex (MEMR). The role of the MOCR was tested by measuring changes in the ear-canal pressure at 6 kHz in the presence and absence of a band-limited harmonic complex tone with various phase curvatures, centered either at (on-frequency) or well below (off-frequency) the 6-kHz probe frequency. The influence of possible MEMR effects was examined by measuring phase-gradient functions for the elicitor effects and by measuring changes in the ear-canal pressure with a continuous suppressor of the 6-kHz probe. Both on- and off-frequency complex tone elicitors produced significant changes in ear canal sound pressure. However, the pattern of results was not consistent with the earlier hypotheses postulating that efferent effects produce the psychoacoustic dependence of forward-masked thresholds on masker phase curvature. The results also reveal unexpectedly long time constants associated with some efferent effects, the source of which remains unknown.

Auditory enhancement refers to the perceptual phenomenon that a target sound is heard out more readily from a background sound if the background is presented alone first. Here we used stimulus-frequency otoacoustic emissions (SFOAEs) to test the hypothesis that activation of the medial olivocochlear efferent system contributes to auditory enhancement effects. The SFOAEs were used as a tool to measure changes in cochlear responses to a target component and the neighboring components of a multitone background between conditions producing enhancement and conditions producing no enhancement. In the "enhancement" condition, the target and multitone background were preceded by a precursor stimulus with a spectral notch around the signal frequency; in the control (no-enhancement) condition, the target and multitone background were presented without the precursor. In an experiment using a wideband multitone stimulus known to produce significant psychophysical enhancement effects, SFOAEs showed no changes consistent with enhancement, but some aspects of the results indicated possible contamination of the SFOAE magnitudes by the activation of the middle-ear-muscle reflex. The same SFOAE measurements performed using narrower-band stimuli at lower sound levels also showed no SFOAE changes consistent with either absolute or relative enhancement despite robust psychophysical enhancement effects observed in the same listeners with the same stimuli. The results suggest that cochlear efferent control does not play a significant role in auditory enhancement effects.

The role of temporal stimulus parameters in the perception of across-frequency synchrony and asynchrony was investigated using pairs of 500-ms tones consisting of a 250-Hz tone and a tone with a higher frequency of 1, 2, 4, or 6 kHz. Subjective judgments suggested veridical perception of across-frequency synchrony but with greater sensitivity to changes in asynchrony for pairs in which the lower-frequency tone was leading than for pairs in which it was lagging. Consistent with the subjective judgments, thresholds for the detection of asynchrony measured in a threealternative forced-choice task were lower when the signal interval contained a pair with the lowfrequency tone leading than a pair with a high-frequency tone leading. A similar asymmetry was observed for asynchrony discrimination when the standard asynchrony was relatively small ( 20 ms) but not for larger standard asynchronies. Independent manipulation of onset and offset ramp durations indicated a dominant role of onsets in the perception of across-frequency asynchrony. A physiologically inspired model, involving broadly tuned monaural coincidence detectors that receive inputs from frequency-selective onset detectors, was able to accurately reproduce the asymmetric distributions of synchrony judgments. The model provides testable predictions for future physiological investigations of responses to broadband stimuli with across-frequency delays.

The effects of selectively attending to a target stimulus in a background containing distractors can be observed in cortical representations of sound as an attenuation of the representation of distractor stimuli. The locus in the auditory system at which attentional modulations first arise is unknown, but anatomical evidence suggests that cortically driven modulation of neural activity could extend as peripherally as the cochlea itself. Previous studies of selective attention have used otoacoustic emissions to probe cochlear function under varying conditions of attention with mixed results. In the current study, two experiments combined visual and auditory tasks to maximize sustained attention, perceptual load, and cochlear dynamic range in an attempt to improve the likelihood of observing selective attention effects on cochlear responses. Across a total of 45 listeners in the two experiments, no systematic effects of attention or perceptual load were observed on stimulus-frequency otoacoustic emissions. The results revealed significant between-subject variability in the otoacoustic-emission measure of cochlear function that does not depend on listener performance in the behavioral tasks and is not related to movement-generated noise. The findings suggest that attentional modulation of auditory information in humans arises at stages of processing beyond the cochlea.

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