Time scales of adaptation to context in horizontal sound localization.

Psychophysical experiments explored how the repeated presentation of a context, consisting of an adaptor and a target, induces plasticity in the localization of an identical target presented alone on interleaved trials. The plasticity, and its time course, was examined both in a classroom and in an anechoic chamber. Adaptors and targets were 2 ms noise clicks and listeners were tasked with localizing the targets while ignoring the adaptors (when present). The context was either simple, consisting of a single-click adaptor and a target, or complex, containing either a single-click or an eight-click adaptor that varied from trial to trial. The adaptor was presented either from a frontal or a lateral location, fixed within a run. The presence of context caused responses to the isolated targets to be displaced up to 14° away from the adaptor location. This effect was stronger and slower if the context was complex, growing over the 5 min duration of the runs. Additionally, the simple context buildup had a slower onset in the classroom. Overall, the results illustrate that sound localization is subject to slow adaptive processes that depend on the spatial and temporal structure of the context and on the level of reverberation in the environment.

Auditory-visual interactions in egocentric distance perception: Ventriloquism effect and aftereffect.

This study describes data on auditory-visual integration and visually-guided adaptation of auditory distance perception using the ventriloquism effect (VE) and ventriloquism aftereffect (VAE). In an experiment, participants judged egocentric distance of interleaved auditory or auditory-visual stimuli with the auditory component located from 0.7 to 2.04 m in front of listeners in a real reverberant environment. The visual component of auditory-visual stimuli was displaced 30% closer (V-closer), 30% farther (V-farther), or aligned (V-aligned) with respect to the auditory component. The VE and VAE were measured in auditory and auditory-visual trials, respectively. Both effects were approximately independent of target distance when expressed in logarithmic units. The VE strength, defined as a difference of V-misaligned and V-aligned response bias, was approximately 72% of the auditory-visual disparity regardless of the visual-displacement direction, while the VAE was stronger in the V-farther (44%) than the V-closer (31%) condition. The VAE persisted to post-adaptation auditory-only blocks of trials, although it was diminished. The rates of build-up/break-down of the VAE were asymmetrical, with slower adaptation in the V-closer condition. These results suggest that auditory-visual distance integration is independent of the direction of induced shift, while the re-calibration is stronger and faster when evoked by more distant visual stimuli.

Cortical auditory distance representation based on direct-to-reverberant energy ratio.

Auditory distance perception and its neuronal mechanisms are poorly understood, mainly because 1) it is difficult to separate distance processing from intensity processing, 2) multiple intensity-independent distance cues are often available, and 3) the cues are combined in a context-dependent way. A recent fMRI study identified human auditory cortical area representing intensity-independent distance for sources presented along the interaural axis (Kopco et al. PNAS, 109, 11019-11024). For these sources, two intensity-independent cues are available, interaural level difference (ILD) and direct-to-reverberant energy ratio (DRR). Thus, the observed activations may have been contributed by not only distance-related, but also direction-encoding neuron populations sensitive to ILD. Here, the paradigm from the previous study was used to examine DRR-based distance representation for sounds originating in front of the listener, where ILD is not available. In a virtual environment, we performed behavioral and fMRI experiments, combined with computational analyses to identify the neural representation of distance based on DRR. The stimuli varied in distance (15-100 â€‹cm) while their received intensity was varied randomly and independently of distance. Behavioral performance showed that intensity-independent distance discrimination is accurate for frontal stimuli, even though it is worse than for lateral stimuli. fMRI activations for sounds varying in frontal distance, as compared to varying only in intensity, increased bilaterally in the posterior banks of Heschl's gyri, the planum temporale, and posterior superior temporal gyrus regions. Taken together, these results suggest that posterior human auditory cortex areas contain neuron populations that are sensitive to distance independent of intensity and of binaural cues relevant for directional hearing.

Hemisphere-specific properties of the ventriloquism aftereffect.

Visual calibration of auditory space requires re-alignment of representations differing in (1) format (auditory hemispheric channels vs visual maps) and (2) reference frames (head-centered vs eye-centered). Here, a ventriloquism paradigm from Kopco, Lin, Shinn-Cunningham, and Groh [J. Neurosci. 29, 13809-13814 (2009)] was used to examine these processes in humans for ventriloquism induced within one spatial hemifield. Results show that (1) the auditory representation can be adapted even by aligned audio-visual stimuli, and (2) the spatial reference frame is primarily head-centered, with a weak eye-centered modulation. These results support the view that the ventriloquism aftereffect is driven by multiple spatially non-uniform, hemisphere-specific processes.

Temporal characteristics of contextual effects in sound localization.

Two experiments examined plasticity induced by context in a simple target localization task. The context was represented by interleaved localization trials with the target preceded by a distractor. In a previous study, the context induced large response shifts when the target and distractor stimuli were identical 2-ms-noise clicks [Kopco, Best, and Shinn-Cunningham (2007). J. Acoust. Soc. Am. 121, 420-432]. Here, the temporal characteristics of the contextual effect were examined for the same stimuli. Experiment 1 manipulated the context presentation rate and the distractor-target inter-stimulus interval (ISI). Experiment 2 manipulated the temporal structure of the context stimulus, replacing the one-click distractor either by a distractor consisting of eight sequentially presented clicks or by a noise burst with total energy and duration identical to the eight-click distractor. In experiment 1, the contextual shift size increased with increasing context rate while being largely independent of ISI. In experiment 2, the eight-click-distractor induced a stronger shift than the one-click-distractor context, while the noise-distractor context induced a very small shift. These results suggest that contextual plasticity is an adaptation driven both by low-level factors like spatiotemporal context distribution and higher-level factors like perceptual similarity between the stimuli, possibly related to precedence buildup.

Streaming and sound localization with a preceding distractor.

Localization of a 2-ms click target was previously shown to be influenced by a preceding identical distractor for inter-click-intervals up to 400 ms [Kopco, Best, and Shinn-Cunningham (2007). J. Acoust. Soc. Am. 121, 420-432]. Here, two experiments examined whether perceptual organization plays a role in this effect. In the experiments, the distractor was designed either to be grouped with the target (a single-click distractor) or to be processed in a separate stream (an 8-click train). The two distractors affected performance differently, both in terms of bias and variance, suggesting that grouping and streaming play a role in localization in multisource environments.

Intracortical depth analyses of frequency-sensitive regions of human auditory cortex using 7TfMRI.

Despite recent advances in auditory neuroscience, the exact functional organization of human auditory cortex (AC) has been difficult to investigate. Here, using reversals of tonotopic gradients as the test case, we examined whether human ACs can be more precisely mapped by avoiding signals caused by large draining vessels near the pial surface, which bias blood-oxygen level dependent (BOLD) signals away from the actual sites of neuronal activity. Using ultra-high field (7T) fMRI and cortical depth analysis techniques previously applied in visual cortices, we sampled 1mm isotropic voxels from different depths of AC during narrow-band sound stimulation with biologically relevant temporal patterns. At the group level, analyses that considered voxels from all cortical depths, but excluded those intersecting the pial surface, showed (a) the greatest statistical sensitivity in contrasts between activations to high vs. low frequency sounds and (b) the highest inter-subject consistency of phase-encoded continuous tonotopy mapping. Analyses based solely on voxels intersecting the pial surface produced the least consistent group results, even when compared to analyses based solely on voxels intersecting the white-matter surface where both signal strength and within-subject statistical power are weakest. However, no evidence was found for reduced within-subject reliability in analyses considering the pial voxels only. Our group results could, thus, reflect improved inter-subject correspondence of high and low frequency gradients after the signals from voxels near the pial surface are excluded. Using tonotopy analyses as the test case, our results demonstrate that when the major physiological and anatomical biases imparted by the vasculature are controlled, functional mapping of human ACs becomes more consistent from subject to subject than previously thought.

Modeling of speech localization in a multi-talker mixture using periodicity and energy-based auditory features.

A recent study showed that human listeners are able to localize a short speech target simultaneously masked by four speech tokens in reverberation [Kopco, Best, and Carlile (2010). J. Acoust. Soc. Am. 127, 1450-1457]. Here, an auditory model for solving this task is introduced. The model has three processing stages: (1) extraction of the instantaneous interaural time difference (ITD) information, (2) selection of target-related ITD information ("glimpses") using a template-matching procedure based on periodicity, spectral energy, or both, and (3) target location estimation. The model performance was compared to the human data, and to the performance of a modified model using an ideal binary mask (IBM) at stage (2). The IBM-based model performed similarly to the subjects, indicating that the binaural model is able to accurately estimate source locations. Template matching using spectral energy and using a combination of spectral energy and periodicity achieved good results, while using periodicity alone led to poor results. Particularly, the glimpses extracted from the initial portion of the signal were critical for good performance. Simulation data show that the auditory features investigated here are sufficient to explain human performance in this challenging listening condition and thus may be used in models of auditory scene analysis.

Neuronal representations of distance in human auditory cortex.

Neuronal mechanisms of auditory distance perception are poorly understood, largely because contributions of intensity and distance processing are difficult to differentiate. Typically, the received intensity increases when sound sources approach us. However, we can also distinguish between soft-but-nearby and loud-but-distant sounds, indicating that distance processing can also be based on intensity-independent cues. Here, we combined behavioral experiments, fMRI measurements, and computational analyses to identify the neural representation of distance independent of intensity. In a virtual reverberant environment, we simulated sound sources at varying distances (15-100 cm) along the right-side interaural axis. Our acoustic analysis suggested that, of the individual intensity-independent depth cues available for these stimuli, direct-to-reverberant ratio (D/R) is more reliable and robust than interaural level difference (ILD). However, on the basis of our behavioral results, subjects' discrimination performance was more consistent with complex intensity-independent distance representations, combining both available cues, than with representations on the basis of either D/R or ILD individually. fMRI activations to sounds varying in distance (containing all cues, including intensity), compared with activations to sounds varying in intensity only, were significantly increased in the planum temporale and posterior superior temporal gyrus contralateral to the direction of stimulation. This fMRI result suggests that neurons in posterior nonprimary auditory cortices, in or near the areas processing other auditory spatial features, are sensitive to intensity-independent sound properties relevant for auditory distance perception.

Contextual plasticity, top-down, and non-auditory factors in sound localization with a distractor.

Localization of a 2-ms-click target was previously shown to be influenced by interleaved localization trials in which the target was preceded by an identical distractor [Kopco, Best, and Shinn-Cunningham (2007). J. Acoust. Soc. Am. 121, 420-432]. Here, two experiments were conducted to explore this contextual effect. Results show that context-related bias is not eliminated (1) when the response method is changed so that vision is available or that no hand-pointing is required; or (2) when the distractor-target order is reversed. Additionally, a keyboard-based localization response method is introduced and shown to be more accurate than traditional pointer-based methods.

Toward a Unified Theory of the Reference Frame of the Ventriloquism Aftereffect.

The ventriloquism aftereffect (VAE), observed as a shift in the perceived locations of sounds after audio-visual stimulation, requires reference frame (RF) alignment since hearing and vision encode space in different RFs (head-centered vs. eye-centered). Previous experimental studies reported inconsistent results, observing either a mixture of head-centered and eye-centered frames, or a predominantly head-centered frame. Here, a computational model is introduced, examining the neural mechanisms underlying these effects. The basic model version assumes that the auditory spatial map is head-centered and the visual signals are converted to head-centered frame prior to inducing the adaptation. Two mechanisms are considered as extended model versions to describe the mixed-frame experimental data: (1) additional presence of visual signals in eye-centered frame and (2) eye-gaze direction-dependent attenuation in VAE when eyes shift away from the training fixation. Simulation results show that the mixed-frame results are mainly due to the second mechanism, suggesting that the RF of VAE is mainly head-centered. Additionally, a mechanism is proposed to explain a new ventriloquism-aftereffect-like phenomenon in which adaptation is induced by aligned audio-visual signals when saccades are used for responding to auditory targets. A version of the model extended to consider such response-method-related biases accurately predicts the new phenomenon. When attempting to model all the experimentally observed phenomena simultaneously, the model predictions are qualitatively similar but less accurate, suggesting that the proposed neural mechanisms interact in a more complex way than assumed in the model.

Object continuity enhances selective auditory attention.

In complex scenes, the identity of an auditory object can build up across seconds. Given that attention operates on perceptual objects, this perceptual buildup may alter the efficacy of selective auditory attention over time. Here, we measured identification of a sequence of spoken target digits presented with distracter digits from other directions to investigate the dynamics of selective attention. Performance was better when the target location was fixed rather than changing between digits, even when listeners were cued as much as 1 s in advance about the position of each subsequent digit. Spatial continuity not only avoided well known costs associated with switching the focus of spatial attention, but also produced refinements in the spatial selectivity of attention across time. Continuity of target voice further enhanced this buildup of selective attention. Results suggest that when attention is sustained on one auditory object within a complex scene, attentional selectivity improves over time. Similar effects may come into play when attention is sustained on an object in a complex visual scene, especially in cases where visual object formation requires sustained attention.

Effect of stimulus spectrum on distance perception for nearby sources.

The effects of stimulus frequency and bandwidth on distance perception were examined for nearby sources in simulated reverberant space. Sources to the side [containing reverberation-related cues and interaural level difference (ILD) cues] and to the front (without ILDs) were simulated. Listeners judged the distance of noise bursts presented at a randomly roving level from simulated distances ranging from 0.15 to 1.7 m. Six stimuli were tested, varying in center frequency (300-5700 Hz) and bandwidth (200-5400 Hz). Performance, measured as the correlation between simulated and response distances, was worse for frontal than for lateral sources. For both simulated directions, performance was inversely proportional to the low-frequency stimulus cutoff, independent of stimulus bandwidth. The dependence of performance on frequency was stronger for frontal sources. These correlation results were well summarized by considering how mean response, as opposed to response variance, changed with stimulus direction and spectrum: (1) little bias was observed for lateral sources, but listeners consistently overestimated distance for frontal nearby sources; (2) for both directions, increasing the low-frequency cut-off reduced the range of responses. These results are consistent with the hypothesis that listeners used a direction-independent but frequency-dependent mapping of a reverberation-related cue, not the ILD cue, to judge source distance.

Localization in speech mixtures by listeners with hearing loss.

The ability of listeners with bilateral sensorineural hearing loss to localize a speech source in a multitalker mixture was measured. Five simultaneous words spoken by different talkers were presented over loudspeakers in a small room, and listeners localized one target word. Errors were significantly larger in this group compared to a control group with normal hearing. Localization of the target presented alone was not different between groups. The results suggest that hearing loss does not impair spatial hearing per se, but degrades the spatial representation of multiple simultaneous sounds.

Reweighting of Binaural Localization Cues Induced by Lateralization Training.

Normal-hearing listeners adapt to alterations in sound localization cues. This adaptation can result from the establishment of a new spatial map of the altered cues or from a stronger relative weighting of unaltered compared to altered cues. Such reweighting has been shown for monaural vs. binaural cues. However, studies attempting to reweight the two binaural cues, interaural differences in time (ITD) and level (ILD), yielded inconclusive results. This study investigated whether binaural-cue reweighting can be induced by lateralization training in a virtual audio-visual environment. Twenty normal-hearing participants, divided into two groups, completed the experiment consisting of 7 days of lateralization training, preceded and followed by a test measuring the binaural-cue weights. Participants' task was to lateralize 500-ms bandpass-filtered (2-4 kHz) noise bursts containing various combinations of spatially consistent and inconsistent binaural cues. During training, additional visual cues reinforced the azimuth corresponding to ITDs in one group and ILDs in the other group and the azimuthal ranges of the binaural cues were manipulated group-specifically. Both groups showed a significant increase of the reinforced-cue weight from pre- to posttest, suggesting that participants reweighted the binaural cues in the expected direction. This reweighting occurred within the first training session. The results are relevant as binaural-cue reweighting likely occurs when normal-hearing listeners adapt to new acoustic environments. Reweighting might also be a factor underlying the low contribution of ITDs to sound localization of cochlear-implant listeners as they typically do not experience reliable ITD cues with clinical devices.

Calibration of Consonant Perception to Room Reverberation.

Purpose We examined how consonant perception is affected by a preceding speech carrier simulated in the same or a different room, for different classes of consonants. Carrier room, carrier length, and carrier length/target room uncertainty were manipulated. A phonetic feature analysis tested which phonetic categories are influenced by the manipulations in the acoustic context of the carrier. Method Two experiments were performed, each with nine participants. Targets consisted of 10 or 16 vowel-consonant (VC) syllables presented in one of two strongly reverberant rooms, preceded by a multiple-VC carrier presented in either the same room, a different reverberant room, or an anechoic room. In Experiment 1, the carrier length and the target room randomly varied from trial to trial, whereas in Experiment 2, they were fixed within a block of trials. Results Overall, a consistent carrier provided an advantage for consonant perception compared to inconsistent carriers, whether in anechoic or differently reverberant rooms. Phonetic analysis showed that carrier inconsistency significantly degraded identification of the manner of articulation, especially for stop consonants and, in one of the rooms, also of voicing. Carrier length and carrier/target uncertainty did not affect adaptation to reverberation for individual phonetic features. The detrimental effects of anechoic and different reverberant carriers on target perception were similar. Conclusions The strength of calibration varies across different phonetic features, as well as across rooms with different levels of reverberation. Even though place of articulation is the feature that is affected by reverberation the most, it is the manner of articulation and, partially, voicing for which room adaptation is observed.

Reference frame of the ventriloquism aftereffect.

Seeing the image of a newscaster on a television set causes us to think that the sound coming from the loudspeaker is actually coming from the screen. How images capture sounds is mysterious because the brain uses different methods for determining the locations of visual versus auditory stimuli. The retina senses the locations of visual objects with respect to the eyes, whereas differences in sound characteristics across the ears indicate the locations of sound sources referenced to the head. Here, we tested which reference frame (RF) is used when vision recalibrates perceived sound locations. Visually guided biases in sound localization were induced in seven humans and two monkeys who made eye movements to auditory or audiovisual stimuli. On audiovisual (training) trials, the visual component of the targets was displaced laterally by 5-6 degrees. Interleaved auditory-only (probe) trials served to evaluate the effect of experience with mismatched visual stimuli on auditory localization. We found that the displaced visual stimuli induced ventriloquism aftereffect in both humans (approximately 50% of the displacement size) and monkeys (approximately 25%), but only for locations around the trained spatial region, showing that audiovisual recalibration can be spatially specific. We tested the reference frame in which the recalibration occurs. On probe trials, we varied eye position relative to the head to dissociate head- from eye-centered RFs. Results indicate that both humans and monkeys use a mixture of the two RFs, suggesting that the neural mechanisms involved in ventriloquism occur in brain region(s) using a hybrid RF for encoding spatial information.

Speech localization in a multitalker mixture.

An experiment was performed that measured, for the frontal audio-visual horizon, how accurately listeners could localize a female-voice target amidst four spatially distributed male-voice maskers. To examine whether listeners can make use of a priori knowledge about the configuration of the sources, performance was examined in two conditions: either the masker locations were fixed (in one of five known patterns) or the locations varied from trial to trial. The presence of maskers disrupted speech localization, even after accounting for reduced target detectability. Averaged across all target locations, the rms error in responses decreased by 20% when a priori knowledge about masker locations was available. The effect was even stronger for the target locations that did not coincide with the maskers (error reduction of 36%), while no change in errors was observed for targets coinciding with maskers. The benefits were reduced when the target-to-masker intensity ratio was increased or when the maskers were in a pattern that made it difficult to make use of the a priori information. The results confirm that localization in speech mixtures is modified by the listener's expectations about the spatial arrangement of the sources.

Exploring the benefit of auditory spatial continuity.

Continuity of spatial location was recently shown to improve the ability to identify and recall a sequence of target digits presented in a mixture of confusable maskers [Best et al. (2008). Proc. Natl. Acad. Sci. U.S.A. 105, 13174-13178]. Three follow-up experiments were conducted to explore the basis of this improvement. The results suggest that the benefits of spatial continuity cannot be attributed to (a) the ability to plan where to direct attention in advance; (b) freedom from having to redirect attention across large distances; or (c) the challenge of filtering out signals that are confusable with the target.

Sound Externalization: A Review of Recent Research.

Sound externalization, or the perception that a sound source is outside of the head, is an intriguing phenomenon that has long interested psychoacousticians. While previous reviews are available, the past few decades have produced a substantial amount of new data.In this review, we aim to synthesize those data and to summarize advances in our understanding of the phenomenon. We also discuss issues related to the definition and measurement of sound externalization and describe quantitative approaches that have been taken to predict the outcomes of externalization experiments. Last, sound externalization is of practical importance for many kinds of hearing technologies. Here, we touch on two examples, discussing the role of sound externalization in augmented/virtual reality systems and bringing attention to the somewhat overlooked issue of sound externalization in wearers of hearing aids.