Interaural Time Differences Discrimination Thresholds Research Articles

The frequency and level dependence of across-frequency binaural interference for interaural time differences

Binaural hearing enables listeners to localize sound in the horizontal plane, even when dealing with intricate speech-like signals characterized by distributed energy across a broad range of frequencies and varying sound pressure levels. Low-frequency interaural time differences (ITDs) are the most important cue for human horizontal-plane sound localization. Across-frequency binaural interference tasks (where changes in ITD discrimination thresholds of a target signal are measured in the presence of a remote interferer signal) can be used to measure how a listener weighs different frequency regions for ITD. While it is known that low-frequency (0.5 kHz) targets experience less binaural interference from higher-frequency (4 kHz) interferers than vice versa, the impact of the relative level of the two signals has remained mostly unexplored. ITD discrimination thresholds were measured in normal-hearing listeners with target/interferer intensities of 35, 55, and 75 dB-A and frequencies of 0.5, 4, and 8 kHz. Besides the expected frequency effects, data to date show binaural interference increases with increasing interferer level. These results help better understand how sound level variation and more realistic complex sounds like speech are localized.

Interaural frequency mismatch jointly modulates neural brainstem binaural interaction and behavioral interaural time difference sensitivity in humans

The binaural interaction component (BIC) of the auditory brainstem response (ABR) is the difference obtained after subtracting the sum of right and left ear ABRs from binaurally evoked ABRs. The BIC has attracted interest as a biomarker of binaural processing abilities. Best binaural processing is presumed to require spectrally-matched inputs at the two ears, but peripheral pathology and/or impacts of hearing devices can lead to mismatched inputs. Such mismatching can degrade behavioral sensitivity to interaural time difference (ITD) cues, but might be detected using the BIC. Here, we examine the effect of interaural frequency mismatch (IFM) on BIC and behavioral ITD sensitivity in audiometrically normal adult human subjects (both sexes). Binaural and monaural ABRs were recorded and BICs computed from subjects in response to narrowband tones. Left ear stimuli were fixed at 4000 Hz while right ear stimuli varied over a ∼2-octave range (re: 4000 Hz). Separately, subjects performed psychophysical lateralization tasks using the same stimuli to determine ITD discrimination thresholds jointly as a function of IFM and sound level. Results demonstrated significant effects of IFM on BIC amplitudes, with lower amplitudes in mismatched conditions than frequency-matched. Behavioral ITD discrimination thresholds were elevated at mismatched frequencies and lower sound levels, but also more sharply modulated by IFM at lower sound levels. Combinations of ITD, IFM and overall sound level that resulted in fused and lateralized percepts were bound by the empirically-measured BIC, and also by model predictions simulated using an established computational model of the brainstem circuit thought to generate the BIC.

Effects of Hearing Loss on Interaural Time Difference Sensitivity at Low and High Frequencies.

While many studies have reported a loss of sensitivity to interaural time differences (ITDs) carried in the fine structure of low-frequency signals for listeners with hearing loss, relatively few data are available on the perception of ITDs carried in the envelope of high-frequency signals in this population. The relevant studies found stronger effects of hearing loss at high frequencies than at low frequencies in most cases, but small subject numbers and several confounding effects prevented strong conclusions from being drawn. In the present study, we revisited this question while addressing some of the issues identified in previous studies. Participants were ten young adults with normal hearing (NH) and twenty adults with sensorineural hearing impairment (HI) spanning a range of ages. ITD discrimination thresholds were measured for octave-band-wide “rustle” stimuli centered at 500 Hz or 4000 Hz, which were presented at 20 or 40 dB sensation level. Broadband rustle stimuli and 500-Hz pure-tone stimuli were also tested. Thresholds were poorer on average for the HI group than the NH group. The ITD deficit, relative to the NH group, was similar at low and high frequencies for most HI participants. For a small number of participants, however, the deficit was strongly frequency-dependent. These results provide new insights into the binaural perception of complex sounds and may inform binaural models that incorporate effects of hearing loss.

The Relationship Between Interaural Insertion-Depth Differences, Scalar Location, and Interaural Time-Difference Processing in Adult Bilateral Cochlear-Implant Listeners.

Sensitivity to interaural time differences (ITDs) in acoustic hearing involves comparison of interaurally frequency-matched inputs. Bilateral cochlear-implant arrays are, however, only approximately aligned in angular insertion depth and scalar location across the cochleae. Interaural place-of-stimulation mismatch therefore has the potential to impact binaural perception. ITD left-right discrimination thresholds were examined in 23 postlingually-deafened adult bilateral cochlear-implant listeners, using low-rate constant-amplitude pulse trains presented via direct stimulation to single electrodes in each ear. Angular insertion depth and scalar location measured from computed-tomography (CT) scans were used to quantify interaural mismatch, and their association with binaural performance was assessed. Number-matched electrodes displayed a median interaural insertion-depth mismatch of 18° and generally yielded best or near-best ITD discrimination thresholds. Two listeners whose discrimination thresholds did not show this pattern were confirmed via CT to have atypical array placement. Listeners with more number-matched electrode pairs located in the scala tympani displayed better thresholds than listeners with fewer such pairs. ITD tuning curves as a function of interaural electrode separation were broad; bandwidths at twice the threshold minimum averaged 10.5 electrodes (equivalent to 5.9 mm for a Cochlear-brand pre-curved array). Larger angular insertion-depth differences were associated with wider bandwidths. Wide ITD tuning curve bandwidths appear to be a product of both monopolar stimulation and angular insertion-depth mismatch. Cases of good ITD sensitivity with very wide bandwidths suggest that precise matching of insertion depth is not critical for discrimination thresholds. Further prioritizing scala tympani location at implantation should, however, benefit ITD sensitivity.

Effects of place of stimulation on the interaural time difference sensitivity in bilateral electrical intracochlear stimulations: Neurophysiological study in a rat model.

We examined the sensitivity of the neurons in the inferior colliculus (IC) in male and female rats to the interaural time differences (ITDs) conveyed in electrical pulse trains. Using bipolar pairs of electrodes that selectively activate the auditory nerve fibers at different intracochlear locations, we assessed whether the responses to electrical stimulation with ITDs in different frequency regions were processed differently. Most well-isolated single units responded to the electrical stimulation in only one of the apical or basal cochlear regions, and they were classified as either apical or basal units. Regardless of the cochlear stimulating location, more than 70% of both apical and basal units were sensitive to ITDs of electrical stimulation. However, the pulse rate dependence of neural ITD sensitivity differed significantly depending on the location of the stimulation. Moreover, ITD discrimination thresholds and the relative incidence of ITD tuning type markedly differed between units activated by apical and basal stimulations. With apical stimulation, IC neurons had a higher incidence of peak-type ITD function, which mostly exhibited the steepest position of the tuning curve within the rat's physiological ITD range of ±160μs and, accordingly, had better ITD discrimination thresholds than those with basal stimulation. These results support the idea that ITD processing in the IC might be determined by functionally segregated frequency-specific pathways from the cochlea to the auditory midbrain.

Consequences of interaural decorrelation of speech temporal fine structure

The ability to understand a target speech signal against a background of interfering speech signals is typically improved when the interfering signals are spatially separated (spatial release from masking; SRM). Swaminathan et al. (2016) found a significant reduction in SRM when the temporal fine structure (TFS) across the left and right ears was decorrelated, suggesting that binaural TFS provides important cues that support SRM. One interpretation is that degrading binaural TFS prevents the extraction of reliable interaural time differences (ITDs), which in turn leads to a reduction in SRM. We tested this hypothesis by systematically decorrelating the binaural TFS, and measuring the effects on both ITD discrimination for speech stimuli and SRM in normal hearing listeners. We show that decorrelation leads to a systematic increase in ITD discrimination thresholds, as well as a systematic decrease in SRM. This supports the idea that binaural TFS is needed to access ITD information in speech, which is in turn required for SRM. Additionally, the relationship between these two tasks provides a framework for determining the extent to which reduced SRM in listeners with hearing loss can be attributed to reduced sensitivity to binaural TFS.

Neural ITD Sensitivity and Temporal Coding with Cochlear Implants in an Animal Model of Early-Onset Deafness.

Users of cochlear implant (CI) face challenges in everyday situations such as understanding conversations in noise, even with CIs in both ears. These challenges are related to difficulties with tasks that require fine temporal processing such as discrimination of pulse rates or interaural time differences (ITD), a major cue for sound localization. The degradation in temporal processing and ITD sensitivity are especially acute in those who lost hearing in early childhood. Here, we characterized temporal coding and ITD sensitivity of single neurons in a novel animal model of early-onset deafness. Rabbits were deafened as neonates and deprived of auditory stimulation until they reached adult age when single-unit recordings from the auditory midbrain were made chronically using an unanesthetized preparation. The results are compared to measurements from adult-deafened rabbits with normal auditory development to understand the effect of early-onset deafness on neural temporal coding and ITD sensitivity. Neurons in the inferior colliculus (IC) of early-deafened rabbits were less likely to show sustained, excitatory responses to pulse train stimulation and more likely to show suppressive responses compared to neurons in adult-deaf animals. Fewer neurons showed synchronized responses to pulse trains at any rate in the early-deaf group. In addition, fewer neurons showed significant ITD sensitivity in their overall firing rate in the early-deaf group compared to adult-deaf animals. Neural ITD discrimination thresholds in the early-deaf group were poorer than thresholds in adult-deaf group, especially at high pulse rates. The overall degradation in neural ITD sensitivity is consistent with the difficulties encountered by human CI users with early-onset hearing loss. These results lay the groundwork for investigating whether the degradations in temporal coding and ITD sensitivity observed in early-deaf animals can be reversed by appropriate CI stimulation during development.

Localization of broadband sounds carrying interaural time differences: Effects of frequency, reference location, and interaural coherence.

The auditory processes involved in the localization of sounds in rooms are still poorly understood. The present study investigated the auditory system's across-frequency processing of interaural time differences (ITDs) and the impact of the interaural coherence (IC) of the stimuli in ITD discrimination and localization. First, ITD discrimination thresholds were measured as a function of signal frequency, reference ITD, and IC using critical-band wide noises. The resulting data were fitted with a set of analytical functions and ITD weights were derived using concepts from signal detection theory. Inspired by the weighted-image model [Stern, Zeiberg, and Trahiotis. (1988). J. Acoust. Soc. Am. 84, 156-165], the derived ITD weights were then integrated in a simplified localization model using an optimal combination of ITD information across frequency. To verify this model, a series of localization experiments were conducted using broadband noise in which ITD and IC were varied across frequency. The model predictions were in good agreement with the experimental data, supporting the assumption that the auditory system performs a weighted integration of ITD information across frequency to localize a sound source. The results could be valuable for the design of new paradigms to measure localization in more complex acoustic conditions and may provide constraints for future localization models.

Binaural localization of musical pitch using interaural time differences in congenital amusia.

Naturally occurring sounds are routinely periodic. The ability to phase-lock to such periodicity facilitates pitch perception and interaural time differences (ITDs) determination in binaural localization. We examined whether deficient pitch processing in individuals with congenital amusia (tone deafness) is accompanied by impaired ability to lateralize musical pitch at auditory periphery and memorize the location of pitch at the working memory level. If common mechanisms subserve processing of temporal-fine-structure based pitch and ITDs, then deficient processing of one feature should impair performance on the other. Thus, we measured ITD discrimination thresholds using an adaptive-tracking procedure for lateralizing musical tone pairs separated by different semitone intervals. Amusic individuals exhibited normal ITD thresholds comparable to those of matched controls, which were not affected by concurrent pitch changes. For working memory tasks, the amusic group performed significantly worse than matched controls in probed pitch recall, irrespective of the complexity level of concurrent variations along the ITD dimension of the melodic sequence. Interestingly, despite normal peripheral ITD thresholds, amusic individuals performed worse than controls in recalling probed locations of tones within a sequence of musical notes originating from different ITD-simulated locations. Findings suggest that individuals with congenital amusia are unimpaired in temporal fine-structure encoding to determine the location of musical pitch based on binaural ITD information at the auditory periphery. However, working memory for a sequence of sounds’ ITD-dependent spatial location is here shown to be impaired and dissociated from the pitch feature of sounds at the working memory level.

Influence of envelope waveform on ITD sensitivity of neurons in the auditory midbrain.

Interaural time differences (ITDs) conveyed by the modulated envelopes of high-frequency sounds can serve as a cue for localizing a sound source. Klein-Hennig et al. (J Acoust Soc Am 129: 3856, 2011) demonstrated the envelope attack (the rate at which stimulus energy in the envelope increases) and the duration of the pause (the interval between successive envelope pulses) as important factors affecting sensitivity to envelope ITDs in human listeners. Modulated sounds with rapid attacks and long pauses produce the lowest ITD discrimination thresholds. The duration of the envelope's sustained component (sustain) and the rate at which stimulus energy falls at the offset of the envelope (decay) are only minor factors. We assessed the responses of 71 single neurons, recorded from the midbrains of 15 urethane-anesthetized tri-colored guinea pigs, to envelope shapes in which the four envelope components, i.e., attack, sustain, decay, and pause, were systematically varied. We confirmed the importance of the attack and pause components in generating ITD-sensitive responses. Analysis of neural firing rates demonstrated more neurons (49/71) show ITD sensitivity in response to "damped" stimuli (fast attack and slow decay) compared with "ramped" stimuli (slow attack and fast decay) (14/71). Furthermore, the lowest threshold for the damped stimulus (91 μs) was lower by a factor of 4 than that for the temporally reversed ramped envelope shape (407 μs). The data confirm the importance of fast attacks and optimal pause durations in generating sensitivity to ITDs conveyed in the modulated envelopes of high-frequency sounds and are incompatible with models of ITD processing based on the integration of sound energy over time.NEW & NOTEWORTHY Using single-neuron electrophysiology, we show that the precise shape of a sound's "energy envelope" is a critical factor in determining how well midbrain neurons are able to convey information about auditory spatial cues. Consistent with human behavioral performance, sounds with rapidly rising energy and relatively long intervals between energy bursts are best at conveying spatial information. The data suggest specific sound energy patterns that might best be applied to hearing devices to aid spatial listening.

The Relationship Between Intensity Coding and Binaural Sensitivity in Adults With Cochlear Implants.

Many bilateral cochlear implant users show sensitivity to binaural information when stimulation is provided using a pair of synchronized electrodes. However, there is large variability in binaural sensitivity between and within participants across stimulation sites in the cochlea. It was hypothesized that within-participant variability in binaural sensitivity is in part affected by limitations and characteristics of the auditory periphery which may be reflected by monaural hearing performance. The objective of this study was to examine the relationship between monaural and binaural hearing performance within participants with bilateral cochlear implants. Binaural measures included dichotic signal detection and interaural time difference discrimination thresholds. Diotic signal detection thresholds were also measured. Monaural measures included dynamic range and amplitude modulation detection. In addition, loudness growth was compared between ears. Measures were made at three stimulation sites per listener. Greater binaural sensitivity was found with larger dynamic ranges. Poorer interaural time difference discrimination was found with larger difference between comfortable levels of the two ears. In addition, poorer diotic signal detection thresholds were found with larger differences between the dynamic ranges of the two ears. No relationship was found between amplitude modulation detection thresholds or symmetry of loudness growth and the binaural measures. The results suggest that some of the variability in binaural hearing performance within listeners across stimulation sites can be explained by factors nonspecific to binaural processing. The results are consistent with the idea that dynamic range and comfortable levels relate to peripheral neural survival and the width of the excitation pattern which could affect the fidelity with which central binaural nuclei process bilateral inputs.

A method to enhance the use of interaural time differences for cochlear implants in reverberant environments

The ability of normal-hearing (NH) listeners to exploit interaural time difference (ITD) cues conveyed in the modulated envelopes of high-frequency sounds is poor compared to ITD cues transmitted in the temporal fine structure at low frequencies. Sensitivity to envelope ITDs is further degraded when envelopes become less steep, when modulation depth is reduced, and when envelopes become less similar between the ears, common factors when listening in reverberant environments. The vulnerability of envelope ITDs is particularly problematic for cochlear implant (CI) users, as they rely on information conveyed by slowly varying amplitude envelopes. Here, an approach to improve access to envelope ITDs for CIs is described in which, rather than attempting to reduce reverberation, the perceptual saliency of cues relating to the source is increased by selectively sharpening peaks in the amplitude envelope judged to contain reliable ITDs. Performance of the algorithm with room reverberation was assessed through simulating listening with bilateral CIs in headphone experiments with NH listeners. Relative to simulated standard CI processing, stimuli processed with the algorithm generated lower ITD discrimination thresholds and increased extents of laterality. Depending on parameterization, intelligibility was unchanged or somewhat reduced. The algorithm has the potential to improve spatial listening with CIs.

Nonuniform temporal weighting of interaural time differences in 500 Hz tones.

The discrimination and lateralization of interaural time differences (ITD) in rapidly modulated high-frequency sounds is dominated by cues present in the initial portion of the sound (i.e., at sound onset). The importance of initial ITD at low frequencies is, however, less clear. Here, ITD discrimination thresholds were measured in 500 Hz pure tones with diotic envelopes and static or dynamic fine-structure ITD. Static-ITD thresholds improved as tone duration increased from 40 to 640 ms but by an amount less than expected from uniform temporal weighting of binaural information. Dynamic conditions eliminated ITD from either the beginning or end of the sound by presenting slightly different frequencies to the two ears. While overall thresholds were lower when ITD was available at sound onset than when it was not, listeners differed appreciably in that regard. The results demonstrate that weighting of ITD is not temporally uniform. Instead, for many listeners, ITD discrimination at 500 Hz appears dominated by ITD cues present in the initial part of the sound. To a variable degree, other listeners rely more equally on ITD cues occurring near sound onsets and offsets, although no listeners appear to utilize such cues uniformly throughout the sound's duration.

Factors affecting the use of envelope interaural time differences in reverberation

At high frequencies, interaural time differences (ITDs) are conveyed by the sound envelope. Sensitivity to envelope ITDs depends crucially on the envelope shape. Reverberation degrades the envelope shape, reducing the modulation depth of the envelope and the slope of its flanks. Reverberation also reduces the envelope interaural coherence (i.e., the similarity of the envelopes at two ears). The current study investigates the extent to which these changes affect sensitivity to envelope ITDs. The first experiment measured ITD discrimination thresholds at low and high frequencies in a simulated room. The stimulus was either a low-frequency narrowband noise or the same noise transposed to a higher frequency. The results suggest that the effect of reverberation on ITD thresholds was multiplicative. Given that the threshold without reverberation was larger for the transposed than for the low-frequency stimulus, this meant that, in absolute terms, the thresholds for the transposed stimulus showed a much greater increase due to reverberation than those for the low-frequency stimulus. Three further experiments indicated that the effect of reverberation on the envelope ITD thresholds was due to the combined effect of the reduction in the envelope modulation depth and slopes, as well as the decrease in the envelope interaural coherence.

Naive performance on interaural time difference discrimination with high-frequency amplitude-modulated tones

Humans are sensitive to changes in interaural time differences (ITDs) in high-frequency sounds with fluctuating amplitude envelopes. However, despite noted large individual differences in high-frequency ITD discrimination ability, to date, most investigations of this performance have employed only a few, typically experienced, listeners. Thus, it is not clear whether the data thus obtained are representative of the general, naive, population. Here, ITD discrimination thresholds with high-frequency amplitude-modulated tones were examined in nineteen listeners, all naive to psychoacoustic testing. The large range of performance across these listeners accommodated the collective data previously reported, but was not fully illustrated by any single previous investigation with a small sample size. For these naive listeners, ITD discrimination performance at high frequencies, 1) was much worse than that at low frequencies, 2) was better with a slower (0.15 kHz) than a faster (0.3 kHz) modulation rate with a 4-kHz carrier, and 3) did not differ between a lower (4 kHz) and higher (6 kHz) carrier frequency with a 0.3-kHz modulation rate. These data help establish high-frequency ITD discrimination performance in the general population, and may aid in the interpretation of performance measured in situations where testing time is limited. [Work supported by NIH.]

Contributions of Intrinsic Neural and Stimulus Variance to Binaural Sensitivity

The discrimination of a change in a stimulus is determined both by the magnitude of that change and by the variability in the neural response to the stimulus. When the stimulus is itself noisy, then the relative contributions of the neural (intrinsic) and stimulus induced variability becomes a critical question. We measured the contribution of intrinsic neural noise and interstimulus variability to the discrimination of interaural time differences (ITDs) and interaural correlation (IAC). We measured discharge rate versus characteristic frequency (CF) tone ITD functions, and CF-centered narrowband noise ITD and IAC functions in interleaved blocks in the same units in the inferior colliculus of urethane-anesthetized guinea pigs. Ten "frozen" tokens of noise were synthesized and the responses to each token were separately analyzed to allow the relative contributions of intrinsic and stimulus variability to be assessed. ITD and IAC discrimination thresholds were determined for a simulated two-interval forced-choice experiment, based on the firing rate distributions, using receiver operating characteristic analysis. On average, between stimulus variability contributed 19% (range, 1.5-30%) of the variance in noise ITD discrimination and 27% (range, 3-50%) in IAC discrimination. Noise ITD thresholds were slightly higher than tone ITD thresholds. Taking the mean of the thresholds for individual noise tokens gave a similar result to pooling across all noise tokens. This implies that although the stimulus induced variability is measurable, it is insignificant in relation to the intrinsic noise in ITD and IAC discrimination.

Sound lateralization test in adolescent blind individuals

Blind individuals require to compensate for the lack of visual information by other sensory inputs. In particular, auditory inputs are crucial to such individuals. To investigate whether blind individuals localize sound in space better than sighted individuals, we tested the auditory ability of adolescent blind individuals using a sound lateralization method. The interaural time difference discrimination thresholds of blind individuals were statistically significantly shorter than those of blind individuals with residual vision and controls. These findings suggest that blind individuals have better auditory spatial ability than individuals with visual cues; therefore, some perceptual compensation occurred in the former.

Effects of different amounts of brief training and rest on the generalization of learning from interaural-level-difference to interaural-time-difference discrimination

Training-induced improvements on perceptual skills can be enhanced by increasing the amount of training and by resting between training and testing. However, how these two factors affect the generalization of learning from a briefly trained condition to an untrained one is unknown. Here, listeners were trained on an interaural-level-difference (ILD) discrimination condition (4-kHz tones), then were tested for generalization to an interaural-time-difference (ITD) discrimination condition (0.5-kHz tones). The amount of training and the time between training and testing differed across four groups. Listeners tested 10 hours after training had significantly lower ITD discrimination thresholds than naive listeners (n=94), regardless of whether training lasted for 20 min (n=14) or 2 hs (n=11). Thus, when there was a long time between training and testing, learning generalized from ILD to ITD discrimination regardless of the amount of training. In contrast, listeners tested immediately after training on ILD discrimination showed generalization to ITD discrimination with 20 min (n=14), but not 2 h (n=14), of training, suggesting that without rest between training and testing, longer training disrupted generalization. Hence, increased training interfered with generalization from ILD to ITD discrimination for short, but not long, delays between training and testing. [Work supported by NIH.]

The basic hearing abilities of absolute pitch possessors.

Generally, people tend to think that absolute pitch possessors or musical experts are superior in all of their auditory and musical abilities. However, not much data has been shown to prove whether any differences exist in the basic hearing abilities of absolute pitch possessors or musical experts for general sounds outside the context of music. In this study, we conducted four experiments to investigate if absolute pitch possessors or music experts are superior in their basic hearing abilities with respect to frequency resolution, temporal resolution, and spatial resolution. In Experiments 1 and 2, we measured frequency discrimination thresholds and thresholds to detect a tone in notched noise to examine the characteristics of frequency resolution. In Experiment 3, we conducted a gap detection task to measure the temporal resolution. Finally, in Experiment 4, we measured interaural time difference discrimination thresholds to examine the spatial resolution abilities. The overall results show that there were no significant differences in frequency, temporal and spatial resolutions among groups with different absolute pitch capability. This indicates that absolute-pitch possessors do not have particularly ‘good ears’ in terms of resolution.