Psychoacoustics publications

Active mechanical amplification of sound occurs in cochlear outer hair cells (OHCs) that change their length with oscillations of their membrane potential. Such length changes are the proposed cellular source of the cochlear amplifier, and prestin is the motor protein responsible for OHC electromotility. Previous findings have shown that mice lacking prestin displayed a loss of OHC electromotility, subsequent loss of distortion-product otoacoustic emissions, and a 40-60 dB increase in hearing thresholds. In this study we were interested in studying the functional consequences of the complete loss of cochlear amplification on neural coding of frequency selectivity, tuning, and temporal processing in the auditory midbrain. We recorded near-field auditory evoked potentials and multi-unit activity from the inferior colliculus (IC) of prestin (-/-) null and prestin (+/+) wild-type control mice and determined frequency response areas (FRAs), tuning sharpness, and gap detection to tone bursts and silent gaps embedded in broadband noise. We were interested in determining if the moderate to severe sensorineural hearing loss associated with the loss of motor protein prestin would also impair auditory midbrain temporal-processing measures, or if compensatory mechanisms within the brainstem could compensate for the loss of prestin. In prestin knockout mice we observed that there are severe impairments in midbrain tuning, thresholds, excitatory drive, and gap detection suggesting that brainstem and midbrain processing could not overcome the auditory processing deficits afforded by the loss of OHC electromotility mediated by the prestin protein.

Forward masking of a sinusoidal signal is determined not only by the masker's power spectrum but also by its phase spectrum. Specifically, when the phase spectrum is such that the output of an auditory filter centred on the signal has a highly modulated ("peaked") envelope, there is less masking than when that envelope is flat. This finding has been attributed to non-linearities, such as compression, reducing the average neural response to maskers that produce more peaked auditory filter outputs (Carlyon and Datta, J Acoust Soc Am 101:3636-3647, 1997). Here we evaluate an alternative explanation proposed by Wotcjzak and Oxenham (Wojtczak and Oxenham, J Assoc Res Otolaryngol 10:595-607, 2009). They reported a masker phase effect for 6-kHz signals when the masker components were at least an octave below the signal frequency. Wotcjzak and Oxenham argued that this effect was inconsistent with cochlear compression, and, because it did not occur at lower signal frequencies, was also inconsistent with more central compression. It was instead attributed to activation of the efferent system reducing the response to the subsequent probe. Here, experiment 1 replicated their main findings. Experiment 2 showed that the phase effect on off-frequency forward masking is similar at signal frequencies of 2 and 6 kHz, provided that one equates the number of components likely to interact within an auditory filter centred on the signal, thereby roughly equating the effect of masker phase on the peakiness of that filter output. Experiment 3 showed that for some subjects, masker phase also had a strong influence on off-frequency backward masking of the signal, and that the size of this effect correlated across subjects with that observed in forward masking. We conclude that the masker phase effect is mediated mainly by cochlear non-linearities, with a possible additional effect of more central compression. The data are not consistent with a role for the efferent system.

The goal of this study was to investigate how cognitive factors influence performance in a multi-talker, "cocktail-party" like environment in musicians and non-musicians. This was achieved by relating performance in a spatial hearing task to cognitive processing abilities assessed using measures of executive function (EF) and visual attention in musicians and non-musicians. For the spatial hearing task, a speech target was presented simultaneously with two intelligible speech maskers that were either colocated with the target (0° azimuth) or were symmetrically separated from the target in azimuth (at ±15°). EF assessment included measures of cognitive flexibility, inhibition control and auditory working memory. Selective attention was assessed in the visual domain using a multiple object tracking task (MOT). For the MOT task, the observers were required to track target dots (n = 1,2,3,4,5) in the presence of interfering distractor dots. Musicians performed significantly better than non-musicians in the spatial hearing task. For the EF measures, musicians showed better performance on measures of auditory working memory compared to non-musicians. Furthermore, across all individuals, a significant correlation was observed between performance on the spatial hearing task and measures of auditory working memory. This result suggests that individual differences in performance in a cocktail party-like environment may depend in part on cognitive factors such as auditory working memory. Performance in the MOT task did not differ between groups. However, across all individuals, a significant correlation was found between performance in the MOT and spatial hearing tasks. A stepwise multiple regression analysis revealed that musicianship and performance on the MOT task significantly predicted performance on the spatial hearing task. Overall, these findings confirm the relationship between musicianship and cognitive factors including domain-general selective attention and working memory in solving the "cocktail party problem".

Are musicians better able to understand speech in noise than non-musicians? Recent findings have produced contradictory results. Here we addressed this question by asking musicians and non-musicians to understand target sentences masked by other sentences presented from different spatial locations, the classical ‘cocktail party problem’ in speech science. We found that musicians obtained a substantial benefit in this situation, with thresholds ~6 dB better than non-musicians. Large individual differences in performance were noted particularly for the non-musically trained group. Furthermore, in different conditions we manipulated the spatial location and intelligibility of the masking sentences, thus changing the amount of ‘informational masking’ (IM) while keeping the amount of ‘energetic masking’ (EM) relatively constant. When the maskers were unintelligible and spatially separated from the target (low in IM), musicians and non-musicians performed comparably. These results suggest that the characteristics of speech maskers and the amount of IM can influence the magnitude of the differences found between musicians and non-musicians in multiple-talker “cocktail party” environments. Furthermore, considering the task in terms of the EM-IM distinction provides a conceptual framework for future behavioral and neuroscientific studies which explore the underlying sensory and cognitive mechanisms contributing to enhanced “speech-in-noise” perception by musicians.

Auditory neurons that exhibit stimulus-specific adaptation (SSA) decrease their response to common tones while retaining responsiveness to rare ones. We recorded single-unit responses from the inferior colliculus (IC) where SSA is known to occur and we explored for the first time SSA in the cochlear nucleus (CN) of rats. We assessed an important functional outcome of SSA, the extent to which frequency discriminability depends on sensory context. For this purpose, pure tones were presented in an oddball sequence as standard (high probability of occurrence) or deviant (low probability of occurrence) stimuli. To study frequency discriminability under different probability contexts, we varied the probability of occurrence and the frequency separation between tones. The neuronal sensitivity was estimated in terms of spike-count probability using signal detection theory. We reproduced the finding that many neurons in the IC exhibited SSA, but we did not observe significant SSA in our CN sample. We concluded that strong SSA is not a ubiquitous phenomenon in the CN. As predicted, frequency discriminability was enhanced in IC when stimuli were presented in an oddball context, and this enhancement was correlated with the degree of SSA shown by the neurons. In contrast, frequency discrimination by CN neurons was independent of stimulus context. Our results demonstrated that SSA is not widespread along the entire auditory pathway, and suggest that SSA increases frequency discriminability of single neurons beyond that expected from their tuning curves.

Key points The cerebral cortex plays a critical role in perception and in learning‐induced plasticity. We show that reversibly silencing any of the main regions of auditory cortex impairs the ability of adult ferrets to localize sound, with the largest deficit seen after deactivating the primary fields. Although these animals had no trouble localizing longer sound bursts, their performance dropped considerably when auditory spatial cues were altered by occluding one ear with an earplug. In contrast to control ferrets, which recovered their localization abilities with intensive training, adaptation to an earplug was impaired following cortical inactivation, with the greatest disruption in plasticity observed after silencing higher‐level cortical areas. These findings imply regional differences in the processing of spatial information across the auditory cortex. Abstract The contribution of auditory cortex to spatial information processing was explored behaviourally in adult ferrets by reversibly deactivating different cortical areas by subdural placement of a polymer that released the GABA A agonist muscimol over a period of weeks. The spatial extent and time course of cortical inactivation were determined electrophysiologically. Muscimol‐Elvax was placed bilaterally over the anterior (AEG), middle (MEG) or posterior ectosylvian gyrus (PEG), so that different regions of the auditory cortex could be deactivated in different cases. Sound localization accuracy in the horizontal plane was assessed by measuring both the initial head orienting and approach‐to‐target responses made by the animals. Head orienting behaviour was unaffected by silencing any region of the auditory cortex, whereas the accuracy of approach‐to‐target responses to brief sounds (40 ms noise bursts) was reduced by muscimol‐Elvax but not by drug‐free implants. Modest but significant localization impairments were observed after deactivating the MEG, AEG or PEG, although the largest deficits were produced in animals in which the MEG, where the primary auditory fields are located, was silenced. We also examined experience‐induced spatial plasticity by reversibly plugging one ear. In control animals, localization accuracy for both approach‐to‐target and head orienting responses was initially impaired by monaural occlusion, but recovered with training over the next few days. Deactivating any part of the auditory cortex resulted in less complete recovery than in controls, with the largest deficits observed after silencing the higher‐level cortical areas in the AEG and PEG. Although suggesting that each region of auditory cortex contributes to spatial learning, differences in the localization deficits and degree of adaptation between groups imply a regional specialization in the processing of spatial information across the auditory cortex.

Elderly individuals often complain of difficulties in understanding speech, especially when heard against a background noise or when there are multiple speakers[...]

We systematically determined which spectrotemporal modulations in speech are necessary for comprehension by human listeners. Speech comprehension has been shown to be robust to spectral and temporal degradations, but the specific relevance of particular degradations is arguable due to the complexity of the joint spectral and temporal information in the speech signal. We applied a novel modulation filtering technique to recorded sentences to restrict acoustic information quantitatively and to obtain a joint spectrotemporal modulation transfer function for speech comprehension, the speech MTF. For American English, the speech MTF showed the criticality of low modulation frequencies in both time and frequency. Comprehension was significantly impaired when temporal modulations <12 Hz or spectral modulations <4 cycles/kHz were removed. More specifically, the MTF was bandpass in temporal modulations and low-pass in spectral modulations: temporal modulations from 1 to 7 Hz and spectral modulations <1 cycles/kHz were the most important. We evaluated the importance of spectrotemporal modulations for vocal gender identification and found a different region of interest: removing spectral modulations between 3 and 7 cycles/kHz significantly increases gender misidentifications of female speakers. The determination of the speech MTF furnishes an additional method for producing speech signals with reduced bandwidth but high intelligibility. Such compression could be used for audio applications such as file compression or noise removal and for clinical applications such as signal processing for cochlear implants.

Perceptual consequences of disrupted auditory nerve activity were systematically studied in 21 subjects who had been clinically diagnosed with auditory neuropathy (AN), a recently defined disorder characterized by normal outer hair cell function but disrupted auditory nerve function. Neurological and electrophysical evidence suggests that disrupted auditory nerve activity is due to desynchronized or reduced neural activity or both. Psychophysical measures showed that the disrupted neural activity has minimal effects on intensity-related perception, such as loudness discrimination, pitch discrimination at high frequencies, and sound localization using interaural level differences. In contrast, the disrupted neural activity significantly impairs timing related perception, such as pitch discrimination at low frequencies, temporal integration, gap detection, temporal modulation detection, backward and forward masking, signal detection in noise, binaural beats, and sound localization using interaural time differences. These perceptual consequences are the opposite of what is typically observed in cochlear-impaired subjects who have impaired intensity perception but relatively normal temporal processing after taking their impaired intensity perception into account. These differences in perceptual consequences between auditory neuropathy and cochlear damage suggest the use of different neural codes in auditory perception: a suboptimal spike count code for intensity processing, a synchronized spike code for temporal processing, and a duplex code for frequency processing. We also proposed two underlying physiological models based on desynchronized and reduced discharge in the auditory nerve to successfully account for the observed neurological and behavioral data. These methods and measures cannot differentiate between these two AN models, but future studies using electric stimulation of the auditory nerve via a cochlear implant might. These results not only show the unique contribution of neural synchrony to sensory perception but also provide guidance for translational research in terms of better diagnosis and management of human communication disorders.

The present study aimed to examine the effect of noise on vowel-like suprathreshold discrimination in cochlear-implant listeners. The task was to detect an increment in level at the middle harmonic (400 Hz) in the background of a seven-harmonic complex from 100 to 700 Hz in 100-Hz steps. The task was performed in the absence (control) and presence of a white noise presented over a 20-35-dB range from inaudible to loud. The present result shows that discrimination of suprathreshold harmonic stimuli was significantly enhanced, particularly at the soft signal level, with suprathreshold noise. This result suggests that tuning of the noise level is required to optimize performance of different tasks in cochlear implants.

When the source of a tone moves with respect to a listener’s ears, dichotic (or interaural) phase and amplitude modulations (PM and AM) are produced. Two experiments investigated the psychophysical characteristics of dichotic linear ramp modulations in phase and amplitude, and compared them with the psychophysics of diotic PM and AM. In experiment 1, subjects were substantially more sensitive to dichotic PM than diotic PM, but AM sensitivity was equivalent in the dichotic and diotic conditions. Thresholds for discriminating modulation direction were smaller than detection thresholds for dichotic AM, and both diotic AM and PM. Dichotic PM discrimination thresholds were similar to detection thresholds. In experiment 2, the effects of ramp duration were examined. Sensitivity to dichotic AM and PM, and diotic AM increased as duration was increased from 20 ms to 200 ms. The functions relating sensitivity to ramp duration differed across the stimuli; sensitivity to dichotic PM increased more rapidly than sensitivity to dichotic or diotic AM. This was also reflected in shorter time-constants and minimum integration times for dichotic PM detection. These findings support the hypothesis that the analysis of dichotic PM and AM rely on separate mechanisms.