Marc M. Van Wanrooij

Contribution of Head Shadow and Pinna Cues to Chronic Monaural Sound Localization

Monaurally deaf people lack the binaural acoustic difference cues in sound level and timing that are needed to encode sound location in the horizontal plane (azimuth). It has been proposed that these people therefore rely on spectral pinna cues of their normal ear to localize sounds. However, the acoustic head-shadow effect (HSE) might also serve as an azimuth cue, despite its ambiguity when absolute sound levels are unknown. Here, we assess the contribution of either cue in the monaural deaf to two-dimensional (2D) sound localization. In a localization test with randomly interleaved sound levels, we show that all monaurally deaf listeners relied heavily on the HSE, whereas binaural control listeners ignore this cue. However, some monaural listeners responded partly to actual sound-source azimuth, regardless of sound level. We show that these listeners extracted azimuth information from their pinna cues. The better monaural listeners were able to localize azimuth on the basis of spectral cues, the better their ability to also localize sound-source elevation. In a subsequent localization experiment with one fixed sound level, monaural listeners rapidly adopted a strategy on the basis of the HSE. We conclude that monaural spectral cues are not sufficient for adequate 2D sound localization under unfamiliar acoustic conditions. Thus, monaural listeners strongly rely on the ambiguous HSE, which may help them to cope with familiar acoustic environments.Monaurally deaf people lack the binaural acoustic difference cues in sound level and timing that are needed to encode sound location in the horizontal plane (azimuth). It has been proposed that these people therefore rely on spectral pinna cues of their normal ear to localize sounds. However, the acoustic head-shadow effect (HSE) might also serve as an azimuth cue, despite its ambiguity when absolute sound levels are unknown. Here, we assess the contribution of either cue in the monaural deaf to two-dimensional (2D) sound localization. In a localization test with randomly interleaved sound levels, we show that all monaurally deaf listeners relied heavily on the HSE, whereas binaural control listeners ignore this cue. However, some monaural listeners responded partly to actual sound-source azimuth, regardless of sound level. We show that these listeners extracted azimuth information from their pinna cues. The better monaural listeners were able to localize azimuth on the basis of spectral cues, the better their ability to also localize sound-source elevation. In a subsequent localization experiment with one fixed sound level, monaural listeners rapidly adopted a strategy on the basis of the HSE. We conclude that monaural spectral cues are not sufficient for adequate 2D sound localization under unfamiliar acoustic conditions. Thus, monaural listeners strongly rely on the ambiguous HSE, which may help them to cope with familiar acoustic environments.

Improved horizontal directional hearing in bone conduction device users with acquired unilateral conductive hearing loss

We examined horizontal directional hearing in patients with acquired severe unilateral conductive hearing loss (UCHL). All patients (n = 12) had been fitted with a bone conduction device (BCD) to restore bilateral hearing. The patients were tested in the unaided (monaural) and aided (binaural) hearing condition. Five listeners without hearing loss were tested as a control group while listening with a monaural plug and earmuff, or with both ears (binaural). We randomly varied stimulus presentation levels to assess whether listeners relied on the acoustic head-shadow effect (HSE) for horizontal (azimuth) localization. Moreover, to prevent sound localization on the basis of monaural spectral shape cues from head and pinna, subjects were exposed to narrow band (1/3 octave) noises. We demonstrate that the BCD significantly improved sound localization in 8/12 of the UCHL patients. Interestingly, under monaural hearing (BCD off), we observed fairly good unaided azimuth localization performance in 4/12 of the patients. Our multiple regression analysis shows that all patients relied on the ambiguous HSE for localization. In contrast, acutely plugged control listeners did not employ the HSE. Our data confirm and further extend results of recent studies on the use of sound localization cues in chronic and acute monaural listening.

Acquired prior knowledge modulates audiovisual integration

Orienting responses to audiovisual events in the environment can benefit markedly by the integration of visual and auditory spatial information. However, logically, audiovisual integration would only be considered successful for stimuli that are spatially and temporally aligned, as these would be emitted by a single object in space–time. As humans do not have prior knowledge about whether novel auditory and visual events do indeed emanate from the same object, such information needs to be extracted from a variety of sources. For example, expectation about alignment or misalignment could modulate the strength of multisensory integration. If evidence from previous trials would repeatedly favour aligned audiovisual inputs, the internal state might also assume alignment for the next trial, and hence react to a new audiovisual event as if it were aligned. To test for such a strategy, subjects oriented a head‐fixed pointer as fast as possible to a visual flash that was consistently paired, though not always spatially aligned, with a co‐occurring broadband sound. We varied the probability of audiovisual alignment between experiments. Reaction times were consistently lower in blocks containing only aligned audiovisual stimuli than in blocks also containing pseudorandomly presented spatially disparate stimuli. Results demonstrate dynamic updating of the subject’s prior expectation of audiovisual congruency. We discuss a model of prior probability estimation to explain the results.

Pinna Cues Determine Orienting Response Modes to Synchronous Sounds in Elevation

To program a goal-directed orienting response toward a sound source embedded in an acoustic scene, the audiomotor system should detect and select the target against a background. Here, we focus on whether the system can segregate synchronous sounds in the midsagittal plane (elevation), a task requiring the auditory system to dissociate the pinna-induced spectral localization cues. Human listeners made rapid head-orienting responses toward either a single sound source (broadband buzzer or Gaussian noise) or toward two simultaneously presented sounds (buzzer and noise) at a wide variety of locations in the midsagittal plane. In the latter case, listeners had to orient to the buzzer (target) and ignore the noise (nontarget). In the single-sound condition, localization was accurate. However, in the double-sound condition, response endpoints depended on relative sound level and spatial disparity. The loudest sound dominated the responses, regardless of whether it was the target or the nontarget. When the sounds had about equal intensities and their spatial disparity was sufficiently small, endpoint distributions were well described by weighted averaging. However, when spatial disparities exceeded ∼45°, response endpoint distributions became bimodal. Similar response behavior has been reported for visuomotor experiments, for which averaging and bimodal endpoint distributions are thought to arise from neural interactions within retinotopically organized visuomotor maps. We show, however, that the auditory-evoked responses can be well explained by the idiosyncratic acoustics of the pinnae. Hence basic principles of target representation and selection for audition and vision appear to differ profoundly.

Single-sided deafness and directional hearing: contribution of spectral cues and high-frequency hearing loss in the hearing ear

Direction-specific interactions of sound waves with the head, torso, and pinna provide unique spectral-shape cues that are used for the localization of sounds in the vertical plane, whereas horizontal sound localization is based primarily on the processing of binaural acoustic differences in arrival time (interaural time differences, or ITDs) and sound level (interaural level differences, or ILDs). Because the binaural sound-localization cues are absent in listeners with total single-sided deafness (SSD), their ability to localize sound is heavily impaired. However, some studies have reported that SSD listeners are able, to some extent, to localize sound sources in azimuth, although the underlying mechanisms used for localization are unclear. To investigate whether SSD listeners rely on monaural pinna-induced spectral-shape cues of their hearing ear for directional hearing, we investigated localization performance for low-pass filtered (LP, <1.5 kHz), high-pass filtered (HP, >3kHz), and broadband (BB, 0.5–20 kHz) noises in the two-dimensional frontal hemifield. We tested whether localization performance of SSD listeners further deteriorated when the pinna cavities of their hearing ear were filled with a mold that disrupted their spectral-shape cues. To remove the potential use of perceived sound level as an invalid azimuth cue, we randomly varied stimulus presentation levels over a broad range (45–65 dB SPL). Several listeners with SSD could localize HP and BB sound sources in the horizontal plane, but inter-subject variability was considerable. Localization performance of these listeners strongly reduced after diminishing of their spectral pinna-cues. We further show that inter-subject variability of SSD can be explained to a large extent by the severity of high-frequency hearing loss in their hearing ear.

Spectrotemporal Response Properties of Core Auditory Cortex Neurons in Awake Monkey

So far, most studies of core auditory cortex (AC) have characterized the spectral and temporal tuning properties of cells in non-awake, anesthetized preparations. As experiments in awake animals are scarce, we here used dynamic spectral-temporal broadband ripples to study the properties of the spectrotemporal receptive fields (STRFs) of AC cells in awake monkeys. We show that AC neurons were typically most sensitive to low ripple densities (spectral) and low velocities (temporal), and that most cells were not selective for a particular spectrotemporal sweep direction. A substantial proportion of neurons preferred amplitude-modulated sounds (at zero ripple density) to dynamic ripples (at non-zero densities). The vast majority (>93%) of modulation transfer functions were separable with respect to spectral and temporal modulations, indicating that time and spectrum are independently processed in AC neurons. We also analyzed the linear predictability of AC responses to natural vocalizations on the basis of the STRF. We discuss our findings in the light of results obtained from the monkey midbrain inferior colliculus by comparing the spectrotemporal tuning properties and linear predictability of these two important auditory stages.

Stable bottom-up processing during dynamic top-down modulations in monkey auditory cortex

It is unclear whether top‐down processing in the auditory cortex (AC) interferes with its bottom‐up analysis of sound. Recent studies indicated non‐acoustic modulations of AC responses, and that attention changes a neurons spectrotemporal tuning. As a result, the AC would seem ill‐suited to represent a stable acoustic environment, which is deemed crucial for auditory perception. To assess whether top‐down signals influence acoustic tuning in tasks without directed attention, we compared monkey single‐unit AC responses to dynamic spectrotemporal sounds under different behavioral conditions. Recordings were mostly made from neurons located in primary fields (primary AC and area R of the AC) that were well tuned to pure tones, with short onset latencies. We demonstrated that responses in the AC were substantially modulated during an auditory detection task and that these modulations were systematically related to top‐down processes. Importantly, despite these significant modulations, the spectrotemporal receptive fields of all neurons remained remarkably stable. Our results suggest multiplexed encoding of bottom‐up acoustic and top‐down task‐related signals at single AC neurons. This mechanism preserves a stable representation of the acoustic environment despite strong non‐acoustic modulations.

Temporal Cortex Activation to Audiovisual Speech in Normal-Hearing and Cochlear Implant Users Measured with Functional Near-Infrared Spectroscopy

Background: Speech understanding may rely not only on auditory, but also on visual information. Non-invasive functional neuroimaging techniques can expose the neural processes underlying the integration of multisensory processes required for speech understanding in humans. Nevertheless, noise (from functional MRI, fMRI) limits the usefulness in auditory experiments, and electromagnetic artifacts caused by electronic implants worn by subjects can severely distort the scans (EEG, fMRI). Therefore, we assessed audio-visual activation of temporal cortex with a silent, optical neuroimaging technique: functional near-infrared spectroscopy (fNIRS). Methods: We studied temporal cortical activation as represented by concentration changes of oxy- and deoxy-hemoglobin in four, easy-to-apply fNIRS optical channels of 33 normal-hearing adult subjects and five post-lingually deaf cochlear implant (CI) users in response to supra-threshold unisensory auditory and visual, as well as to congruent auditory-visual speech stimuli. Results: Activation effects were not visible from single fNIRS channels. However, by discounting physiological noise through reference channel subtraction (RCS), auditory, visual and audiovisual (AV) speech stimuli evoked concentration changes for all sensory modalities in both cohorts (p < 0.001). Auditory stimulation evoked larger concentration changes than visual stimuli (p < 0.001). A saturation effect was observed for the AV condition. Conclusions: Physiological, systemic noise can be removed from fNIRS signals by RCS. The observed multisensory enhancement of an auditory cortical channel can be plausibly described by a simple addition of the auditory and visual signals with saturation.

The influence of static eye and head position on the ventriloquist effect

Orienting responses to audiovisual events have shorter reaction times and better accuracy and precision when images and sounds in the environment are aligned in space and time. How the brain constructs an integrated audiovisual percept is a computational puzzle because the auditory and visual senses are represented in different reference frames: the retina encodes visual locations with respect to the eyes; whereas the sound localisation cues are referenced to the head. In the well‐known ventriloquist effect, the auditory spatial percept of the ventriloquists voice is attracted toward the synchronous visual image of the dummy, but does this visual bias on sound localisation operate in a common reference frame by correctly taking into account eye and head position? Here we studied this question by independently varying initial eye and head orientations, and the amount of audiovisual spatial mismatch. Human subjects pointed head and/or gaze to auditory targets in elevation, and were instructed to ignore co‐occurring visual distracters. Results demonstrate that different initial head and eye orientations are accurately and appropriately incorporated into an audiovisual response. Effectively, sounds and images are perceptually fused according to their physical locations in space independent of an observers point of view. Implications for neurophysiological findings and modelling efforts that aim to reconcile sensory and motor signals for goal‐directed behaviour are discussed.Orienting responses to audiovisual events have shorter reaction times and better accuracy and precision when images and sounds in the environment are aligned in space and time. How the brain constructs an integrated audiovisual percept is a computational puzzle because the auditory and visual senses are represented in different reference frames: the retina encodes visual locations with respect to the eyes; whereas the sound localisation cues are referenced to the head. In the well-known ventriloquist effect, the auditory spatial percept of the ventriloquists voice is attracted toward the synchronous visual image of the dummy, but does this visual bias on sound localisation operate in a common reference frame by correctly taking into account eye and head position? Here we studied this question by independently varying initial eye and head orientations, and the amount of audiovisual spatial mismatch. Human subjects pointed head and/or gaze to auditory targets in elevation, and were instructed to ignore co-occurring visual distracters. Results demonstrate that different initial head and eye orientations are accurately and appropriately incorporated into an audiovisual response. Effectively, sounds and images are perceptually fused according to their physical locations in space independent of an observers point of view. Implications for neurophysiological findings and modelling efforts that aim to reconcile sensory and motor signals for goal-directed behaviour are discussed.

Task-related preparatory modulations multiply with acoustic processing in monkey auditory cortex

We characterised task‐related top‐down signals in monkey auditory cortex cells by comparing single‐unit activity during passive sound exposure with neuronal activity during a predictable and unpredictable reaction‐time task for a variety of spectral‐temporally modulated broadband sounds. Although animals were not trained to attend to particular spectral or temporal sound modulations, their reaction times demonstrated clear acoustic spectral‐temporal sensitivity for unpredictable modulation onsets. Interestingly, this sensitivity was absent for predictable trials with fast manual responses, but re‐emerged for the slower reactions in these trials. Our analysis of neural activity patterns revealed a task‐related dynamic modulation of auditory cortex neurons that was locked to the animals reaction time, but invariant to the spectral and temporal acoustic modulations. This finding suggests dissociation between acoustic and behavioral signals at the single‐unit level. We further demonstrated that single‐unit activity during task execution can be described by a multiplicative gain modulation of acoustic‐evoked activity and a task‐related top‐down signal, rather than by linear summation of these signals.