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A John Van Opstal

Publications and source records attributed to A John Van Opstal.

12 recordsLinked to original sources

Sound localization under perturbed binaural hearing.

This paper reports on the acute effects of a monaural plug on directional hearing in the horizontal (azimuth) and vertical (elevation) planes of human listeners. Sound localization behavior was tested with rapid head-orienting responses toward brief high-pass filtered (>3 kHz; HP) and broadband (0.5-20 kHz; BB) noises, with sound levels between 30 and 60 dB, A-weighted (dBA). To deny listeners any consistent azimuth-related head-shadow cues, stimuli were randomly interleaved. A plug immediately degraded azimuth performance, as evidenced by a sound level-dependent shift ("bias") of responses contralateral to the plug, and a level-dependent change in the slope of the stimulus-response relation ("gain"). Although the azimuth bias and gain were highly correlated, they could not be predicted from the plug's acoustic attenuation. Interestingly, listeners performed best for low-intensity stimuli at their normal-hearing side. These data demonstrate that listeners rely on monaural spectral cues for sound-source azimuth localization as soon as the binaural difference cues break down. Also the elevation response components were affected by the plug: elevation gain depended on both stimulus azimuth and on sound level and, as for azimuth, localization was best for low-intensity stimuli at the hearing side. Our results show that the neural computation of elevation incorporates a binaural weighting process that relies on the perceived, rather than the actual, sound-source azimuth. It is our conjecture that sound localization ensues from a weighting of all acoustic cues for both azimuth and elevation, in which the weights may be partially determined, and rapidly updated, by the reliability of the particular cue.

Acoustic Stimulation↗

Using double-magnetic induction to measure head-unrestrained gaze shifts. I. Theory and validation.

So far, the double-magnetic induction (DMI) method has been successfully applied to record eye movements from head-restrained humans, monkeys and cats. An advantage of the DMI method, compared to the more widely used scleral search coil technique, is the absence of vulnerable lead wires on the eye. A disadvantage, however, is that the relationship between the eye-in-head orientation and the secondary induction signal is highly non-linear and non-monotonic. This limits the effective measuring range to maximum eye orientations of about +/-30 degrees . Here, we analyze and test two extensions required to record the full eye-head orienting range, well exceeding 90 degrees from straight-ahead in all directions. (1) The use of mutually perpendicular magnetic fields allows for the disambiguation of the non-monotonic signal from the ring. (2) The application of an artificial neural network for offline calibration of the signals. The theoretical predictions are tested for horizontal rotations with a gimbal system. Our results show that the method is a promising alternative to the search coil technique.

Animals↗

Gaze orienting in dynamic visual double steps.

Visual stimuli are initially represented in a retinotopic reference frame. To maintain spatial accuracy of gaze (i.e., eye in space) despite intervening eye and head movements, the visual input could be combined with dynamic feedback about ongoing gaze shifts. Alternatively, target coordinates could be updated in advance by using the preprogrammed gaze-motor command ("predictive remapping"). So far, previous experiments have not dissociated these possibilities. Here we study whether the visuomotor system accounts for saccadic eye-head movements that occur during target presentation. In this case, the system has to deal with fast dynamic changes of the retinal input and with highly variable changes in relative eye and head movements that cannot be preprogrammed by the gaze control system. We performed visual-visual double-step experiments in which a brief (50-ms) stimulus was presented during a saccadic eye-head gaze shift toward a previously flashed visual target. Our results show that gaze shifts remain accurate under these dynamic conditions, even for stimuli presented near saccade onset, and that eyes and head are driven in oculocentric and craniocentric coordinates, respectively. These results cannot be explained by a predictive remapping scheme. We propose that the visuomotor system adequately processes dynamic changes in visual input that result from self-initiated gaze shifts, to construct a stable representation of visual targets in an absolute, supraretinal (e.g., world) reference frame. Predictive remapping may subserve transsaccadic integration, thus enabling perception of a stable visual scene despite eye movements, whereas dynamic feedback ensures accurate actions (e.g., eye-head orienting) to a selected goal.

Adult↗

Relearning sound localization with a new ear.

Human sound localization results primarily from the processing of binaural differences in sound level and arrival time for locations in the horizontal plane (azimuth) and of spectral shape cues generated by the head and pinnae for positions in the vertical plane (elevation). The latter mechanism incorporates two processing stages: a spectral-to-spatial mapping stage and a binaural weighting stage that determines the contribution of each ear to perceived elevation as function of sound azimuth. We demonstrated recently that binaural pinna molds virtually abolish the ability to localize sound-source elevation, but, after several weeks, subjects regained normal localization performance. It is not clear which processing stage underlies this remarkable plasticity, because the auditory system could have learned the new spectral cues separately for each ear (spatial-mapping adaptation) or for one ear only, while extending its contribution into the contralateral hemifield (binaural-weighting adaptation). To dissociate these possibilities, we applied a long-term monaural spectral perturbation in 13 subjects. Our results show that, in eight experiments, listeners learned to localize accurately with new spectral cues that differed substantially from those provided by their own ears. Interestingly, five subjects, whose spectral cues were not sufficiently perturbed, never yielded stable localization performance. Our findings indicate that the analysis of spectral cues may involve a correlation process between the sensory input and a stored spectral representation of the subject's ears and that learning acts predominantly at a spectral-to-spatial mapping level rather than at the level of binaural weighting.

Acoustic Stimulation↗

Crossmodal integration in the primate superior colliculus underlying the preparation and initiation of saccadic eye movements.

Saccades to combined audiovisual stimuli often have reduced saccadic reaction times (SRTs) compared with those to unimodal stimuli. Neurons in the intermediate/deep layers of the superior colliculus (dSC) are capable of integrating converging sensory inputs to influence the time to saccade initiation. To identify how neural processing in the dSC contributes to reducing SRTs to audiovisual stimuli, we recorded activity from dSC neurons while monkeys generated saccades to visual or audiovisual stimuli. To evoke crossmodal interactions of varying strength, we used auditory and visual stimuli of different intensities, presented either in spatial alignment or to opposite hemifields. Spatially aligned audiovisual stimuli evoked the shortest SRTs. In the case of low-intensity stimuli, the response to the auditory component of the aligned audiovisual target increased the activity preceding the response to the visual component, accelerating the onset of the visual response and facilitating the generation of shorter-latency saccades. In the case of high-intensity stimuli, the auditory and visual responses occurred much closer together in time and so there was little opportunity for the auditory stimulus to influence previsual activity. Instead, the reduction in SRT for high-intensity, aligned audiovisual stimuli was correlated with increased premotor activity (activity after visual burst but preceding saccade-aligned burst). These data provide a link between changes in neural activity related to stimulus modality with changes in behavior. They further demonstrate how crossmodal interactions are not limited to the initial sensory activity but can also influence premotor activity in the SC.

Acoustic Stimulation↗

Dynamic sound localization during rapid eye-head gaze shifts.

Human sound localization relies on implicit head-centered acoustic cues. However, to create a stable and accurate representation of sounds despite intervening head movements, the acoustic input should be continuously combined with feedback signals about changes in head orientation. Alternatively, the auditory target coordinates could be updated in advance by using either the preprogrammed gaze-motor command or the sensory target coordinates to which the intervening gaze shift is made ("predictive remapping"). So far, previous experiments cannot dissociate these alternatives. Here, we study whether the auditory system compensates for ongoing saccadic eye and head movements in two dimensions that occur during target presentation. In this case, the system has to deal with dynamic changes of the acoustic cues as well as with rapid changes in relative eye and head orientation that cannot be preprogrammed by the audiomotor system. We performed visual-auditory double-step experiments in two dimensions in which a brief sound burst was presented while subjects made a saccadic eye-head gaze shift toward a previously flashed visual target. Our results show that localization responses under these dynamic conditions remain accurate. Multiple linear regression analysis revealed that the intervening eye and head movements are fully accounted for. Moreover, elevation response components were more accurate for longer-duration sounds (50 msec) than for extremely brief sounds (3 msec), for all localization conditions. Taken together, these results cannot be explained by a predictive remapping scheme. Rather, we conclude that the human auditory system adequately processes dynamically varying acoustic cues that result from self-initiated rapid head movements to construct a stable representation of the target in world coordinates. This signal is subsequently used to program accurate eye and head localization responses.

Acoustic Stimulation↗

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.

Acoustic Stimulation↗

Involvement of monkey inferior colliculus in spatial hearing.

The midbrain inferior colliculus (IC) is implicated in coding sound location, but evidence from behaving primates is scarce. Here we report single-unit responses to broadband sounds that were systematically varied within the two-dimensional (2D) frontal hemifield, as well as in sound level, while monkeys fixated a central visual target. Results show that IC neurons are broadly tuned to both sound-source azimuth and level in a way that can be approximated by multiplicative, planar modulation of the firing rate of the cell. In addition, a fraction of neurons also responded to elevation. This tuning, however, was more varied: some neurons were sensitive to a specific elevation; others responded to elevation in a monotonic way. Multiple-linear regression parameters varied from cell to cell, but the only topography encountered was a dorsoventral tonotopy. In a second experiment, we presented sounds from straight ahead while monkeys fixated visual targets at different positions. We found that auditory responses in a fraction of IC cells were weakly, but systematically, modulated by 2D eye position. This modulation was absent in the spontaneous firing rates, again suggesting a multiplicative interaction of acoustic and eye-position inputs. Tuning parameters to sound frequency, location, intensity, and eye position were uncorrelated. On the basis of simulations with a simple neural network model, we suggest that the population of IC cells could encode the head-centered 2D sound location and enable a direct transformation of this signal into the eye-centered topographic motor map of the superior colliculus. Both signals are required to generate rapid eye-head orienting movements toward sounds.

Acoustic Stimulation↗

The influence of duration and level on human sound localization.

The localization of sounds in the vertical plane (elevation) deteriorates for short-duration wideband sounds at moderate to high intensities. The effect is described by a systematic decrease of the elevation gain (slope of stimulus-response relation) at short sound durations. Two hypotheses have been proposed to explain this finding. Either the sound localization system integrates over a time window that is too short to accurately extract the spectral localization cues (neural integration hypothesis), or the effect results from cochlear saturation at high intensities (adaptation hypothesis). While the neural integration model predicts that elevation gain is independent of sound level, the adaptation hypothesis holds that low elevation gains for short-duration sounds are only obtained at high intensities. Here, these predictions are tested over a larger range of stimulus parameters than has been done so far. Subjects responded with rapid head movements to noise bursts in the two-dimensional frontal space. Stimulus durations ranged from 3 to 100 ms; sound levels from 26 to 73 dB SPL. Results show that the elevation gain decreases for short noise bursts at all sound levels, a finding that supports the integration model. On the other hand, the short-duration gain also decreases at high sound levels, which is in line with the adaptation hypothesis. The finding that elevation gain was a nonmonotonic function of sound level for all sound durations, however, is predicted by neither model. It is concluded that both mechanisms underlie the elevation gain effect and a conceptual model is proposed to reconcile these findings.

Adult↗

Plasticity in human sound localization induced by compressed spatial vision.

Auditory and visual target locations are encoded differently in the brain, but must be co-calibrated to maintain cross-sensory concordance. Mechanisms that adjust spatial calibration across modalities have been described (for example, prism adaptation in owls), though rudimentarily in humans. We quantified the adaptation of human sound localization in response to spatially compressed vision (0.5x lenses for 2-3 days). This induced a corresponding compression of auditory localization that was most pronounced for azimuth (minimal for elevation) and was restricted to the visual field of the lenses. Sound localization was also affected outside the field of visual-auditory interaction (shifted centrally, not compressed). These results suggest that spatially modified vision induces adaptive changes in adult human sound localization, including novel mechanisms that account for spatial compression. Findings are consistent with a model in which the central processing of sound location is encoded by recruitment rather than by a place code.

Acoustic Stimulation↗

A method to induce swapped binaural hearing.

This paper describes the application of a small hearing aid that precisely fits into a subject's ear canal (complete-in-canal, or CIC). The bandwidth of the device is about 7 kHz. The system allows for selective manipulation of the different acoustic cues used for sound localization. The potential of the system is illustrated by robustly interchanging the input of the left and right ear, and consequently changing the sign of the binaural difference cues (both interaural phase and intensity) that are used for horizontal sound localization. As a result, left-right perception is reversed, while high-frequency pinna cues are sufficiently preserved to maintain up-down localization. As the hearing condition is well-defined, the auditory system could in principle remap these cues into a new representation of sound azimuth by relating the modified cues to veridical sound locations. The hearing aids were applied in four human subjects. Swapped binaural hearing was tested in two of the subjects. Swapped localization experiments for an extended period indicated stable performance of both subjects. Interestingly, an adaptive response to the reversed interaural cues was not observed. The current system may prove useful for psychophysical studies that concern the independent processing of sound localization cues, as well as in long-term developmental and plasticity studies with animals.

Acoustic Stimulation↗

Bayesian reconstruction of sound localization cues from responses to random spectra.

The directionally sensitive acoustics of the pinnae enable humans to perceive the up-down and front-back direction of sound. This mechanism complements another, independent mechanism that derives sound-source azimuth from interaural difference cues. The pinnae effectively add direction-dependent spectral notches and peaks to the incoming sound, and it has been shown that such features are used to code sound direction in the median plane. However, it is still unclear which of the pinna-induced features play a role in sound localization. The present study presents a method for the reconstruction of the spatially relevant features in the spectral domain. Broadband sounds with random spectral shapes were presented in rapid succession as subjects made saccadic eye movements toward the perceived stimulus locations. The analysis, which is based on Bayesian statistics, indicates that specific spectral features could be associated with perceived spatial locations. Spectral features that were determined by this psychophysical method resemble the main characteristics of the pinna transfer functions obtained from acoustic measurements in the ear canal. Despite current experimental limitations, the approach may prove useful in the study of perceptually relevant spectral cues underlying human sound localization.

Acoustic Stimulation↗