Psychoacoustics publications

Background: The characteristic voice quality of a speaker conveys important linguistic, paralinguistic, and vocal health-related information. Pitch strength refers to the salience of pitch sensation in a sound and was recently reported to be strongly correlated with the magnitude of perceived breathiness based on a small number of voice stimuli. Objective: The current study examined the relationship between perceptual judgments of breathiness and computational estimates of pitch strength based on the Aud-SWIPE (P-NP) algorithm for a large number of voice stimuli (330 synthetic and 57 natural). Methods and results: Similar to the earlier study, the current results confirm a strong relationship between estimated pitch strength and listener judgments of breathiness such that low pitch-strength values are associated with voices that have high perceived breathiness. Based on this result, a model was developed for the perception of breathy voice quality using a pitch-strength estimator. Regression functions derived between the pitch-strength estimates and perceptual judgments of breathiness obtained from matching task revealed a linear relationship for a subset of the natural stimuli. We then used this function to obtain predicted breathiness values for the synthetic and the remaining natural stimuli. Conclusions: Predicted breathiness values from our model were highly correlated with the perceptual data for both types of stimuli. Systematic differences between the breathiness of natural and synthetic stimuli are discussed.

The study measured listener sensitivity to increments in the duration of one or two target silent intervals embedded within unaccented tone sequences and sequences that featured a single accented component. The baseline unaccented sequences consisted of six 1000-Hz 40-ms tone bursts that were separated equally by silent intervals to establish a slower tone sequence rate, with tonal inter-onset intervals (IOIs) set to 200 ms, or a faster rate with tonal IOIs set to 100 ms. Stimulus accent was created by doubling the baseline duration of a single sequence component, either the second tone burst (tonal accent), or the second tonal IOI (interval accent). Duration difference limens for increments of the target interval(s) were measured adaptively by varying a single inter-tone silent interval or co-varying two successive inter-tone silent intervals; target intervals occurred at the third, or third and fourth, sequence locations. Listeners included younger normal-hearing adults and groups of older listeners with and without hearing loss. Discrimination for the two older groups was equivalent and poorer than that of the younger listeners, especially for the faster accented sequences. Discrimination was best for stimuli with two successive target intervals, indicating that target repetition within accented sequences acts to improve listener temporal sensitivity.

Physical features of sensory stimuli are fixed, but sensory perception is context dependent. The precise mechanisms that govern contextual modulation remain unknown. Here, we trained mice to switch between two contexts: passively listening to pure tones and performing a recognition task for the same stimuli. Two-photon imaging showed that many excitatory neurons in auditory cortex were suppressed during behavior, while some cells became more active. Whole-cell recordings showed that excitatory inputs were affected only modestly by context, but inhibition was more sensitive, with PV+, SOM+, and VIP+ interneurons balancing inhibition and disinhibition within the network. Cholinergic modulation was involved in context switching, with cholinergic axons increasing activity during behavior and directly depolarizing inhibitory cells. Network modeling captured these findings, but only when modulation coincidently drove all three interneuron subtypes, ruling out either inhibition or disinhibition alone as sole mechanism for active engagement. Parallel processing of cholinergic modulation by cortical interneurons therefore enables context-dependent behavior.

The detection of a sensory stimulus arises from a significant change in neural activity, but a sensory neuron's response is rarely identical to successive presentations of the same stimulus. Large trial-to-trial variability would limit the central nervous system's ability to reliably detect a stimulus, presumably affecting perceptual performance. However, if response variability were to decrease while firing rate remained constant, then neural sensitivity could improve. Here, we asked whether engagement in an auditory detection task can modulate response variability, thereby increasing neural sensitivity. We recorded telemetrically from the core auditory cortex of gerbils, both while they engaged in an amplitude-modulation detection task and while they sat quietly listening to the identical stimuli. Using a signal detection theory framework, we found that neural sensitivity was improved during task performance, and this improvement was closely associated with a decrease in response variability. Moreover, units with the greatest change in response variability had absolute neural thresholds most closely aligned with simultaneously measured perceptual thresholds. Our findings suggest that the limitations imposed by response variability diminish during task performance, thereby improving the sensitivity of neural encoding and potentially leading to better perceptual sensitivity. SIGNIFICANCE STATEMENT The detection of a sensory stimulus arises from a significant change in neural activity. However, trial-to-trial variability of the neural response may limit perceptual performance. If the neural response to a stimulus is quite variable, then the response on a given trial could be confused with the pattern of neural activity generated when the stimulus is absent. Therefore, a neural mechanism that served to reduce response variability would allow for better stimulus detection. By recording from the cortex of freely moving animals engaged in an auditory detection task, we found that variability of the neural response becomes smaller during task performance, thereby improving neural detection thresholds.

Listeners with normal hearing thresholds (NHTs) differ in their ability to steer attention to whatever sound source is important. This ability depends on top-down executive control, which modulates the sensory representation of sound in the cortex. Yet, this sensory representation also depends on the coding fidelity of the peripheral auditory system. Both of these factors may thus contribute to the individual differences in performance. We designed a selective auditory attention paradigm in which we could simultaneously measure envelope following responses (EFRs, reflecting peripheral coding), onset event-related potentials (ERPs) from the scalp (reflecting cortical responses to sound) and behavioral scores. We performed two experiments that varied stimulus conditions to alter the degree to which performance might be limited due to fine stimulus details vs. due to control of attentional focus. Consistent with past work, in both experiments we find that attention strongly modulates cortical ERPs. Importantly, in Experiment I, where coding fidelity limits the task, individual behavioral performance correlates with subcortical coding strength (derived by computing how the EFR is degraded for fully masked tones compared to partially masked tones); however, in this experiment, the effects of attention on cortical ERPs were unrelated to individual subject performance. In contrast, in Experiment II, where sensory cues for segregation are robust (and thus less of a limiting factor on task performance), inter-subject behavioral differences correlate with subcortical coding strength. In addition, after factoring out the influence of subcortical coding strength, behavioral differences are also correlated with the strength of attentional modulation of ERPs. These results support the hypothesis that behavioral abilities amongst listeners with NHTs can arise due to both subcortical coding differences and differences in attentional control, depending on stimulus characteristics and task demands.

Spatial judgments are often dominated by low-frequency binaural cues and onset cues when binaural cues vary across the spectrum and duration, respectively, of a brief sound. This study combined these dimensions to assess the spectrotemporal weighting of binaural information. Listeners discriminated target interaural time difference (ITD) and interaural level difference (ILD) carried by the onset, offset, or full duration of a 4-kHz Gabor click train with a 2-ms period in the presence or absence of a diotic 500-Hz interferer tone. ITD and ILD thresholds were significantly elevated by the interferer in all conditions and by a similar amount to previous reports for static cues. Binaural interference was dramatically greater for ITD targets lacking onset cues compared to onset and full-duration conditions. Binaural interference for ILD targets was similar across dynamic-cue conditions. These effects mirror the baseline discriminability of dynamic ITD and ILD cues [Stecker and Brown. (2010). J. Acoust. Soc. Am. 127, 3092–3103], consistent with stronger interference for less-robust/higher-variance cues. The results support the view that binaural cue integration occurs simultaneously across multiple variance-weighted dimensions, including time and frequency.

Background A link between musical expertise and auditory temporal processing abilities was examined. Material and Methods Trained musicians (n=13) and non-musicians (n=12) were tested on speech tasks (phonetic identification, speech recognition in noise) and non-speech tasks (temporal gap detection). Results Results indicated musicians had shorter between-channel gap detection thresholds and sharper phonetic identification functions, suggesting that perceptual reorganization following musical training assists basic temporal auditory processes. Conclusions In general, our results provide a conceptual advance in understanding how musical training influences speech processing, an ability which, when impaired, can affect speech and reading competency.

Difficulties with second-language vowel perception may be related to the significant challenges in using acoustic-phonetic cues. This study investigated the effects of perception training with duration-equalized vowels on native Chinese listeners' English vowel perception and their use of acoustic-phonetic cues. Seventeen native Chinese listeners were perceptually trained with duration-equalized English vowels, and another 17 native Chinese listeners watched English videos as a control group. Both groups were tested with English vowel identification and vowel formant discrimination before training, immediately after training, and three months later. The results showed that the training effect was greater for the vowel training group than for the control group, while both groups improved their English vowel identification and vowel formant discrimination after training. Moreover, duration-equalized vowel perception training significantly reduced listeners' reliance on duration cues and improved their use of spectral cues in identifying English vowels, but video-watching did not help. The results suggest that duration-equalized English vowel perception training may improve non-native listeners' English vowel perception by changing their perceptual weights of acoustic-phonetic cues.

Previous research demonstrated that while selectively attending to relevant aspects of the external world, the brain extracts pertinent information by aligning its neuronal oscillations to key time points of stimuli or their sampling by sensory organs. This alignment mechanism is termed oscillatory entrainment. We investigated the global, long-timescale dynamics of this mechanism in the primary auditory cortex of nonhuman primates, and hypothesized that lapses of entrainment would correspond to lapses of attention. By examining electrophysiological and behavioral measures, we observed that besides the lack of entrainment by external stimuli, attentional lapses were also characterized by high-amplitude alpha oscillations, with alpha frequency structuring of neuronal ensemble and single-unit operations. Entrainment and alpha-oscillation-dominated periods were strongly anticorrelated and fluctuated rhythmically at an ultra-slow rate. Our results indicate that these two distinct brain states represent externally versus internally oriented computational resources engaged by large-scale task-positive and task-negative functional networks.

The high prevalence of noise-induced and age-related hearing loss in the general population has warranted the use of animal models to study the etiology of these pathologies. Quick and accurate auditory threshold determination is a prerequisite for experimental manipulations targeting hearing loss in animal models. The standard auditory brainstem response (ABR) measurement is fairly quick and translational across species, but is limited by the need for anesthesia and a lack of perceptual assessment. The goal of this study was to develop a new method of hearing assessment utilizing prepulse inhibition (PPI) of the acoustic startle reflex, a commonly used tool that measures detection thresholds in awake animals, and can be performed on multiple animals simultaneously. We found that in control mice PPI audiometric functions are similar to both ABR and traditional operant conditioning audiograms. The hearing thresholds assessed with PPI audiometry in sound exposed mice were also similar to those detected by ABR thresholds one day after exposure. However, three months after exposure PPI threshold shifts were still evident at and near the frequency of exposure whereas ABR thresholds recovered to the pre-exposed level. In contrast, PPI audiometry and ABR wave one amplitudes detected similar losses. PPI audiometry provides a high throughput automated behavioral screening tool of hearing in awake animals. Overall, PPI audiometry and ABR assessments of the auditory system are robust techniques with distinct advantages and limitations, which when combined, can provide ample information about the functionality of the auditory system.

Objective: This study examined the effect of microphone placement on the interaural level differences (ILDs) available to bilateral cochlear implant (BiCI) users, and the subsequent effects on horizontal-plane sound localization. Design: Virtual acoustic stimuli for sound localization testing were created individually for eight BiCI users by making acoustic transfer function measurements for microphones placed in the ear (ITE), behind the ear (BTE), and on the shoulders (SHD). The ILDs across source locations were calculated for each placement to analyze their effect on sound localization performance. Sound localization was tested using a repeated-measures, within-participant design for the three microphone placements. Results: The ITE microphone placement provided significantly larger ILDs compared to BTE and SHD placements, which correlated with overall localization errors. However, differences in localization errors across the microphone conditions were small. Conclusions: The BTE microphones worn by many BiCI users in everyday life do not capture the full range of acoustic ILDs available, and also reduce the change in cue magnitudes for sound sources across the horizontal plane. Acute testing with an ITE placement reduced sound localization errors along the horizontal plane compared to the other placements in some patients. Larger improvements may be observed if patients had more experience with the new ILD cues provided by an ITE placement.

Mice often produce ultrasonic vocalizations (USVs) that sweep upwards in frequency from around 60 to around 80 kHz and downwards in frequency from 80 to 60 kHz. Whether or not these USVs are used for communication purposes is still unknown. Here, mice were trained and tested using operant conditioning procedures and positive reinforcement to discriminate between synthetic upsweeps and downsweeps. The stimuli varied in bandwidth, duration, and direction of sweep. The mice performed significantly worse when discriminating between background and test stimuli when the stimuli all occupied the same bandwidths. Further, the mice's discrimination performance became much worse for stimuli that had durations similar to those natural vocalizations of the mice. Sweeps composed of different frequency ranges and longer durations had improved discrimination. These results collected using artificial stimuli created to mimic natural USVs indicate that the bandwidth of the vocalizations may be much more important for communication than the frequency contours of the vocalizations.

The Speech Intelligibility Index (SII) assumes additivity of the importance of acoustically independent bands of speech. To further evaluate this assumption, open-set speech recognition was measured for words and sentences, in quiet and in noise, when the speech stimuli were presented to the listener in selected frequency bands. The filter passbands were constructed from various combinations of 20 bands having equivalent (0.05) importance in the SII framework. This permitted the construction of a variety of equal-SII band patterns that were then evaluated by nine different groups of young adults with normal hearing. For monosyllabic words, a similar dependence on band pattern was observed for SII values of 0.4, 0.5, and 0.6 in both quiet and noise conditions. Specifically, band patterns concentrated toward the lower and upper frequency range tended to yield significantly lower scores than those more evenly sampling a broader frequency range. For all stimuli and test conditions, equal SII values did not yield equal performance. Because the spectral distortions of speech evaluated here may not commonly occur in everyday listening conditions, this finding does not necessarily represent a serious deficit for the application of the SII. These findings, however, challenge the band-independence assumption of the theory underlying the SII.

Just-noticeable differences (JNDs) in interaural time delay (ITD), interaural level difference (ILD), and interaural cross-correlation (ICC) were measured with low- and high-frequency noise bands over multiple sessions for 10 normal-hearing (NH) and 11 hearing-impaired (HI) listeners. Individual subject thresholds tended to improve with training then stabilize. Measured JNDs varied over these experienced listeners, for both subject groups and all tasks. Group JNDs were seldom predictable from hearing level. Individual listeners' JNDs were highly correlated across frequency for each task and group, except for ICC in the HI listeners. Further, ITD JNDs almost always significantly correlated with ILD JNDs within a group. Finally, although the ICC JNDs always significantly correlated with the ITD or ILD JNDs for the NH listeners, they often did not for the HI listeners. These findings suggest that little information about binaural sensitivity is added for NH listeners with multiple ITD, ILD, and ICC measures. For HI listeners, however, while ITD and ILD measures are well correlated, information is added with ICC measures. In general, the results suggest that less information is added with JND measures for NH listeners (15 significant correlations) than for HI listeners (six significant correlations).

Gap detection or gap pre-pulse inhibition of the acoustic startle (GPIAS) has been successfully used in rat and guinea pig models of tinnitus, yet this system has been proven to have low efficacy in CBA mice, with low basal GPIAS and subtle tinnitus-like effects. Here, we tested five mouse strains (CBA, BalbC, CD-1, C57BL/6 and 129sv) for pre-pulse inhibition (PPI) and gap detection with varying interstimulus intervals (ISI) and found that mice from a CBA genetic background had the poorest capacities of suppressing the startle response in the presence of a pre-pulse or a gap. CD-1 mice displayed variable responses throughout all ISI. Interestingly, C57BL/6, 129sv and BalbC showed efficient suppression with either pre-pulses or gaps with shorter ISI. The glutamate aspartate transporter (GLAST) is expressed in support cells from the cochlea and buffers the excess of glutamate. We hypothesized that loss of GLAST function could sensitize the ear to tinnitus-inducing agents, such as salicylate. Using shorter ISI to obtain a greater dynamic range to assess tinnitus-like effects, we found that disruption of gap detection by salicylate was exacerbated across various intensities of a 32-kHz narrow band noise gap carrier in GLAST knockout (KO) mice when compared to their wild-type (WT) littermates. Auditory brainstem responses (ABR) and distortion-product otoacoustic emission (DPOAE) were performed to evaluate the effects on hearing functions. Salicylate caused greater auditory threshold shifts (near 15 dB) in GLAST KO mice than in WT mice across all tested frequencies, despite similarly reduced DPOAE. Despite these changes, inhibition using broad-band gap carriers and 32 kHz pre-pulses were not affected. Our study suggests that GLAST deficiency could become a useful experimental model to decipher the mechanisms underlying drug-induced tinnitus. Future studies addressing the neurological correlates of tinnitus in this model could provide additional insights into the mechanisms of tinnitus.

Objects and events in the sensory environment are generally predictable, making most of the energy impinging upon sensory transducers redundant. Given this fact, efficient sensory systems should detect, extract, and exploit predictability in order to optimize sensitivity to less predictable inputs that are, by definition, more informative. Not only are perceptual systems sensitive to changes in physical stimulus properties, but growing evidence reveals sensitivity both to relative predictability of stimuli and to co-occurrence of stimulus attributes within stimuli. Recent results revealed that auditory perception rapidly reorganizes to efficiently capture covariance among stimulus attributes. Acoustic properties per se were perceptually abandoned, and sounds were instead processed relative to patterns of co-occurrence. Here, we show that listeners' ability to distinguish sounds from one another is driven primarily by the extent to which they are consistent or inconsistent with patterns of covariation among stimulus attributes and, to a lesser extent, whether they are heard frequently or infrequently. When sounds were heard frequently and deviated minimally from the prevailing pattern of covariance among attributes, they were poorly discriminated from one another. In stark contrast, when sounds were heard rarely and markedly violated the pattern of covariance, they became hyperdiscriminable with discrimination performance beyond apparent limits of the auditory system. Plausible cortical candidates underlying these dramatic changes in perceptual organization are discussed. These findings support efficient coding of stimulus statistical structure as a model for both perceptual and neural organization.

Unlabelled: While conversing in a crowded social setting, a listener is often required to follow a target speech signal amid multiple competing speech signals (the so-called "cocktail party" problem). In such situations, separation of the target speech signal in azimuth from the interfering masker signals can lead to an improvement in target intelligibility, an effect known as spatial release from masking (SRM). This study assessed the contributions of two stimulus properties that vary with separation of sound sources, binaural envelope (ENV) and temporal fine structure (TFS), to SRM in normal-hearing (NH) human listeners. Target speech was presented from the front and speech maskers were either colocated with or symmetrically separated from the target in azimuth. The target and maskers were presented either as natural speech or as "noise-vocoded" speech in which the intelligibility was conveyed only by the speech ENVs from several frequency bands; the speech TFS within each band was replaced with noise carriers. The experiments were designed to preserve the spatial cues in the speech ENVs while retaining/eliminating them from the TFS. This was achieved by using the same/different noise carriers in the two ears. A phenomenological auditory-nerve model was used to verify that the interaural correlations in TFS differed across conditions, whereas the ENVs retained a high degree of correlation, as intended. Overall, the results from this study revealed that binaural TFS cues, especially for frequency regions below 1500 Hz, are critical for achieving SRM in NH listeners. Potential implications for studying SRM in hearing-impaired listeners are discussed. Significance statement: Acoustic signals received by the auditory system pass first through an array of physiologically based band-pass filters. Conceptually, at the output of each filter, there are two principal forms of temporal information: slowly varying fluctuations in the envelope (ENV) and rapidly varying fluctuations in the temporal fine structure (TFS). The importance of these two types of information in everyday listening (e.g., conversing in a noisy social situation; the "cocktail-party" problem) has not been established. This study assessed the contributions of binaural ENV and TFS cues for understanding speech in multiple-talker situations. Results suggest that, whereas the ENV cues are important for speech intelligibility, binaural TFS cues are critical for perceptually segregating the different talkers and thus for solving the cocktail party problem.

The role of source properties in across-formant integration was explored using three-formant (F1+F2+F3) analogues of natural sentences (targets). In experiment 1, F1+F3 were harmonic analogues (H1+H3) generated using a monotonous buzz source and second-order resonators; in experiment 2, F1+F3 were tonal analogues (T1+T3). F2 could take either form (H2 or T2). Target formants were always presented monaurally; the receiving ear was assigned randomly on each trial. In some conditions, only the target was present; in others, a competitor for F2 (F2C) was presented contralaterally. Buzz-excited or tonal competitors were created using the time-reversed frequency and amplitude contours of F2. Listeners must reject F2C to optimize keyword recognition. Whether or not a competitor was present, there was no effect of source mismatch between F1+F3 and F2. The impact of adding F2C was modest when it was tonal but large when it was harmonic, irrespective of whether F2C matched F1+F3. This pattern was maintained when harmonic and tonal counterparts were loudness-matched (experiment 3). Source type and competition, rather than acoustic similarity, governed the phonetic contribution of a formant. Contrary to earlier research using dichotic targets, requiring across-ear integration to optimize intelligibility, H2C was an equally effective informational masker for H2 as for T2.

Cochlear implants are neuroprosthetic devices that provide hearing to deaf patients, although outcomes are highly variable even with prolonged training and use. The central auditory system must process cochlear implant signals, but it is unclear how neural circuits adapt—or fail to adapt—to such inputs. The knowledge of these mechanisms is required for development of next-generation neuroprosthetics that interface with existing neural circuits and enable synaptic plasticity to improve perceptual outcomes. Here, we describe a new system for cochlear implant insertion, stimulation, and behavioral training in rats. Animals were first ensured to have significant hearing loss via physiological and behavioral criteria. We developed a surgical approach for multichannel (2- or 8-channel) array insertion, comparable with implantation procedures and depth in humans. Peripheral and cortical responses to stimulation were used to program the implant objectively. Animals fitted with implants learned to use them for an auditory-dependent task that assesses frequency detection and recognition in a background of environmentally and self-generated noise and ceased responding appropriately to sounds when the implant was temporarily inactivated. This physiologically calibrated and behaviorally validated system provides a powerful opportunity to study the neural basis of neuroprosthetic device use and plasticity.

Target speaker identification is essential for speech enhancement algorithms in assistive devices aimed toward helping the hearing impaired. Several recent studies have reported that target speaker identification is possible through electroencephalography (EEG) recordings. If the EEG system could be reduced to acceptable size while retaining the signal quality, hearing aids could benefit from the integration with concealed EEG. To compare the performance of a multichannel around-the-ear EEG system with high-density cap EEG recordings an envelope tracking algorithm was applied in a competitive speaker paradigm. The data from 20 normal hearing listeners were concurrently collected from the traditional state-of-the-art laboratory wired EEG system and a wireless mobile EEG system with two bilaterally-placed around-the-ear electrode arrays (cEEGrids). The results show that the cEEGrid ear-EEG technology captured neural signals that allowed the identification of the attended speaker above chance-level, with 69.3% accuracy, while cap-EEG signals resulted in the accuracy of 84.8%. Further analyses investigated the influence of ear-EEG signal quality and revealed that the envelope tracking procedure was unaffected by variability in channel impedances. We conclude that the quality of concealed ear-EEG recordings as acquired with the cEEGrid array has potential to be used in the brain-computer interface steering of hearing aids.