Unraveling the Relation between EEG Correlates of Attentional Orienting and Sound Localization Performance: A Diffusion Model Approach.

Understanding the contribution of cognitive processes and their underlying neurophysiological signals to behavioral phenomena has been a key objective in recent neuroscience research. Using a diffusion model framework, we investigated to what extent well-established correlates of spatial attention in the electroencephalogram contribute to behavioral performance in an auditory free-field sound localization task. Younger and older participants were instructed to indicate the horizontal position of a predefined target among three simultaneously presented distractors. The central question of interest was whether posterior alpha lateralization and amplitudes of the anterior contralateral N2 subcomponent (N2ac) predict sound localization performance (accuracy, mean RT) and/or diffusion model parameters (drift rate, boundary separation, non-decision time). Two age groups were compared to explore whether, in older adults (who struggle with multispeaker environments), the brain-behavior relationship would differ from younger adults. Regression analyses revealed that N2ac amplitudes predicted drift rate and accuracy, whereas alpha lateralization was not related to behavioral or diffusion modeling parameters. This was true irrespective of age. The results indicate that a more efficient attentional filtering and selection of information within an auditory scene, reflected by increased N2ac amplitudes, was associated with a higher speed of information uptake (drift rate) and better localization performance (accuracy), while the underlying response criteria (threshold separation), mean RTs, and non-decisional processes remained unaffected. The lack of a behavioral correlate of poststimulus alpha power lateralization constrasts with the well-established notion that prestimulus alpha power reflects a functionally relevant attentional mechanism. This highlights the importance of distinguishing anticipatory from poststimulus alpha power modulations.

Bihemispheric anodal transcranial direct-current stimulation over temporal cortex enhances auditory selective spatial attention.

The capacity to selectively focus on a particular speaker of interest in a complex acoustic environment with multiple persons speaking simultaneously-a so-called "cocktail-party" situation-is of decisive importance for human verbal communication. Here, the efficacy of single-dose transcranial direct-current stimulation (tDCS) in improving this ability was tested in young healthy adults (n = 24), using a spatial task that required the localization of a target word in a simulated "cocktail-party" situation. In a sham-controlled crossover design, offline bihemispheric double-monopolar anodal tDCS was applied for 30 min at 1 mA over auditory regions of temporal lobe, and the participant's performance was assessed prior to tDCS, immediately after tDCS, and 1 h after tDCS. A significant increase in the amount of correct localizations by on average 3.7 percentage points (d = 1.04) was found after active, relative to sham, tDCS, with only insignificant reduction of the effect within 1 h after tDCS offset. Thus, the method of bihemispheric tDCS could be a promising tool for enhancement of human auditory attentional functions that are relevant for spatial orientation and communication in everyday life.

Testing the dual-pathway model for auditory processing in human cortex.

Analogous to the visual system, auditory information has been proposed to be processed in two largely segregated streams: an anteroventral ("what") pathway mainly subserving sound identification and a posterodorsal ("where") stream mainly subserving sound localization. Despite the popularity of this assumption, the degree of separation of spatial and non-spatial auditory information processing in cortex is still under discussion. In the present study, a statistical approach was implemented to investigate potential behavioral dissociations for spatial and non-spatial auditory processing in stroke patients, and voxel-wise lesion analyses were used to uncover their neural correlates. The results generally provided support for anatomically and functionally segregated auditory networks. However, some degree of anatomo-functional overlap between "what" and "where" aspects of processing was found in the superior pars opercularis of right inferior frontal gyrus (Brodmann area 44), suggesting the potential existence of a shared target area of both auditory streams in this region. Moreover, beyond the typically defined posterodorsal stream (i.e., posterior superior temporal gyrus, inferior parietal lobule, and superior frontal sulcus), occipital lesions were found to be associated with sound localization deficits. These results, indicating anatomically and functionally complex cortical networks for spatial and non-spatial auditory processing, are roughly consistent with the dual-pathway model of auditory processing in its original form, but argue for the need to refine and extend this widely accepted hypothesis.

Absence of direction-specific cross-modal visual-auditory adaptation in motion-onset event-related potentials.

Adaptation to visual or auditory motion affects within-modality motion processing as reflected by visual or auditory free-field motion-onset evoked potentials (VEPs, AEPs). Here, a visual-auditory motion adaptation paradigm was used to investigate the effect of visual motion adaptation on VEPs and AEPs to leftward motion-onset test stimuli. Effects of visual adaptation to (i) scattered light flashes, and motion in the (ii) same or in the (iii) opposite direction of the test stimulus were compared. For the motion-onset VEPs, i.e. the intra-modal adaptation conditions, direction-specific adaptation was observed--the change-N2 (cN2) and change-P2 (cP2) amplitudes were significantly smaller after motion adaptation in the same than in the opposite direction. For the motion-onset AEPs, i.e. the cross-modal adaptation condition, there was an effect of motion history only in the change-P1 (cP1), and this effect was not direction-specific--cP1 was smaller after scatter than after motion adaptation to either direction. No effects were found for later components of motion-onset AEPs. While the VEP results provided clear evidence for the existence of a direction-specific effect of motion adaptation within the visual modality, the AEP findings suggested merely a motion-related, but not a direction-specific effect. In conclusion, the adaptation of veridical auditory motion detectors by visual motion is not reflected by the AEPs of the present study.

The effect of brain lesions on sound localization in complex acoustic environments.

Localizing sound sources of interest in cluttered acoustic environments--as in the 'cocktail-party' situation--is one of the most demanding challenges to the human auditory system in everyday life. In this study, stroke patients' ability to localize acoustic targets in a single-source and in a multi-source setup in the free sound field were directly compared. Subsequent voxel-based lesion-behaviour mapping analyses were computed to uncover the brain areas associated with a deficit in localization in the presence of multiple distracter sound sources rather than localization of individually presented sound sources. Analyses revealed a fundamental role of the right planum temporale in this task. The results from the left hemisphere were less straightforward, but suggested an involvement of inferior frontal and pre- and postcentral areas. These areas appear to be particularly involved in the spectrotemporal analyses crucial for effective segregation of multiple sound streams from various locations, beyond the currently known network for localization of isolated sound sources in otherwise silent surroundings.

Ventral and dorsal visual pathways support auditory motion processing in the blind: evidence from electrical neuroimaging.

Blindness induces processes of neural plasticity, resulting in recruitment of the deafferentated visual areas for non-visual sensory functions. These processes are related to superior abilities of blind compared with sighted individuals for specific auditory and tactile tasks. Recently, an exceptional performance of the blind has been demonstrated for auditory motion perception, with a minimum audible movement angle that was half that of sighted controls (J. Lewald (2013) Neuropsychologia, 51, 181-186). The present study revealed an electrophysiological correlate of this finding by analysing the so-called motion-onset response, a prominent auditory-evoked potential to the onset of motion. The cN1 component of this response, appearing about 170 ms after motion onset, was two times higher in amplitude for blind compared with matched sighted control subjects. At the time of the cN1, electrical neuroimaging using sLORETA revealed stronger activation in blind than sighted subjects primarily in ventral visual areas (V1v, V2v, VP, V4v) of the right occipital lobe. Activation was also obtained in middle temporal area V5. These findings suggest that blindness results in stronger involvement of both non-motion areas of the ventral visual stream and motion areas of the dorsal visual stream in processing of auditory motion at the same point in time after motion onset. This argues against the view that visual motion areas, such as area V5, are preferentially recruited for auditory motion analysis in the blind. Rather, cross-modal reorganization of cortical areas induced by blindness seems to be largely independent of the specific visual functions of the same areas in sighted persons.

Effect of attention on cortical processing of sound motion: an EEG study.

The onset of motion in an otherwise continuous sound elicits a prominent auditory evoked potential, the so-called motion onset response (MOR). The MOR has recently been shown to be modulated by stimulus-dependent factors, such as velocity, while the possible role of task-dependent factors has remained unclear. Here, the effect of spatial attention on the MOR was investigated in 19 listeners. In each trial, the subject initially heard a free-field sound, consisting of a stationary period and a subsequent period of motion. Then, two successive stationary test tones were presented that differed in location and pitch. Subjects either judged whether or not the starting and final positions of the preceded motion matched the positions of the two test tones ('motion-focused condition'), or whether or not the test tones were identical in pitch, irrespective of the preceded motion stimulus ('baseline condition'). These two tasks were presented in separate experimental blocks. The performance level in both tasks was similar. However, especially later portions of the MOR were significantly increased in amplitude when auditory motion was task-relevant. Cortical source localization indicated that this extra activation originated in dorsofrontal areas that have been proposed to be part of the dorsal auditory processing stream. These results support the assumption that auditory motion processing is based on a complex interaction of both stimulus-specific and attentional processes.

Short-Term Audiovisual Spatial Training Enhances Electrophysiological Correlates of Auditory Selective Spatial Attention.

Audiovisual cross-modal training has been proposed as a tool to improve human spatial hearing. Here, we investigated training-induced modulations of event-related potential (ERP) components that have been associated with processes of auditory selective spatial attention when a speaker of interest has to be localized in a multiple speaker ("cocktail-party") scenario. Forty-five healthy participants were tested, including younger (19-29 years; n = 21) and older (66-76 years; n = 24) age groups. Three conditions of short-term training (duration 15 min) were compared, requiring localization of non-speech targets under "cocktail-party" conditions with either (1) synchronous presentation of co-localized auditory-target and visual stimuli (audiovisual-congruency training) or (2) immediate visual feedback on correct or incorrect localization responses (visual-feedback training), or (3) presentation of spatially incongruent auditory-target and visual stimuli presented at random positions with synchronous onset (control condition). Prior to and after training, participants were tested in an auditory spatial attention task (15 min), requiring localization of a predefined spoken word out of three distractor words, which were presented with synchronous stimulus onset from different positions. Peaks of ERP components were analyzed with a specific focus on the N2, which is known to be a correlate of auditory selective spatial attention. N2 amplitudes were significantly larger after audiovisual-congruency training compared with the remaining training conditions for younger, but not older, participants. Also, at the time of the N2, distributed source analysis revealed an enhancement of neural activity induced by audiovisual-congruency training in dorsolateral prefrontal cortex (Brodmann area 9) for the younger group. These findings suggest that cross-modal processes induced by audiovisual-congruency training under "cocktail-party" conditions at a short time scale resulted in an enhancement of correlates of auditory selective spatial attention.

Shared cortical systems for processing of horizontal and vertical sound motion.

Cortical processing of horizontal and vertical sound motion in free-field space was investigated using high-density electroencephalography in combination with standardized low-resolution brain electromagnetic tomography (sLORETA). Eighteen subjects heard sound stimuli that, after an initial stationary phase in a central position, started to move centrifugally, either to the left, to the right, upward, or downward. The delayed onset of both horizontal and vertical motion elicited a specific motion-onset response (MOR), resulting in widely distributed activations, with prominent maxima in primary and nonprimary auditory cortices, insula, and parietal lobe. The comparison of MORs to horizontal and vertical motion orientations did not indicate any significant differences in latency or topography. Contrasting the sLORETA solutions for the two motion orientations revealed only marginal activation in postcentral gyrus. These data are consistent with the notion that azimuth and elevation components of dynamic auditory spatial information are processed in common, rather than separate, cortical substrates. Furthermore, the findings support the assumption that the MOR originates at a stage of auditory analysis after the different spatial cues (interaural and monaural spectral cues) have been integrated into a unified space code.

Auditory space perception in left- and right-handers.

Several studies have shown that handedness has an impact on visual spatial abilities. Here we investigated the effect of laterality on auditory space perception. Participants (33 right-handers, 20 left-handers) completed two tasks of sound localization. In a dark, anechoic, and sound-proof room, sound stimuli (broadband noise) were presented via 21 loudspeakers mounted horizontally (from 80 degrees on the left to 80 degrees on the right). Participants had to localize the target either by using a swivel hand-pointer or by head-pointing. Individual lateral preferences of eye, ear, hand, and foot were obtained using a questionnaire. With both pointing methods, participants showed a bias in sound localization that was to the side contralateral to the preferred hand, an effect that was unrelated to their overall precision. This partially parallels findings in the visual modality as left-handers typically have a more rightward bias in visual line bisection compared with right-handers. Despite the differences in neural processing of auditory and visual spatial information these findings show similar effects of lateral preference on auditory and visual spatial perception. This suggests that supramodal neural processes are involved in the mechanisms generating laterality in space perception.

Distortion of auditory space in hemianopia.

Sound localization was investigated in patients with homonymous hemianopia, a visual field defect characterized by a loss of vision in one hemifield that is caused by unilateral brain lesions involving the visual cortex or its afferents. The primary aim was to clarify whether or not the known distortion of visual space in hemianopia results in processes of long-term cross-modal spatial adaptation, thus eventually inducing related alterations in auditory space perception. For this purpose, patients were tested by using tasks of either head pointing or manual pointing to acoustic targets in the azimuthal plane, under anechoic conditions in total darkness. The results obtained with both tasks consistently indicated slight, but significant, systematic errors compared with normal controls. In particular, the errors found can be interpreted by both rotation and compression of auditory space toward the anopic side. These findings can be explained by a visual miscalibration of the auditory space, as has been analogously demonstrated in studies on normal-sighted subjects after exposure to consistent auditory-visual disparity, for example by wearing prism lenses. The precision in sound localization of hemianopic patients was generally reduced across both hemispaces. Taken together, one may conclude that processes of cross-modal spatial adaptation, but not those of compensatory plasticity, occurred in patients with hemianopia.

Perception of stationary and moving sound following unilateral cortectomy.

The perception of motion is an essential prerequisite to responding adequately to the dynamic aspects of sensory information in the environment. The neural substrates of auditory motion processing are, at present, still a matter of debate. It has been hypothesized that motion information is, as in the visual system, processed separately from other aspects of auditory information, such as stationary location. Here we report data on auditory perception of stationary and motion stimuli from a subject with right-sided resection of the anterior temporal-lobe region including medial aspects of Heschl's gyrus, and from three subjects with unilateral (right-sided or left-sided) hemispherectomy. All these subjects had undergone cortectomy decades earlier. The subjects with hemispherectomy were completely unable to perceive auditory motion, but showed slight to moderate deficits in judging stationary location. The subject with temporal lobectomy exhibited quite similar stationary auditory deficits as found in the subjects with hemispherectomy, but was completely normal in judging auditory motion. Thus, there was a clear dissociation of the effects of unilateral temporal lobectomy and hemispherectomy on auditory motion perception. Collectively, these findings suggest that the unilateral anterior temporal-lobe region plays a significant role in the analysis of stationary, but not moving, sound. One may assume that the cortical "motion network" is distinct from the "stationary network", and is located either in the most posterior aspects of temporal lobe, or in non-temporal, most likely parietal, areas.

Constancy of target velocity as a critical factor in the emergence of auditory and visual representational momentum.

Representational momentum refers to the tendency to displace the judged final position of a moving auditory or visual target as being too far forward along the path of motion. This phenomenon was investigated here by comparing apparent displacements in final position with constant or with irregularly varying target velocities. Final positions of auditory or visual targets, moving along the horizontal plane, were indicated by manual pointing. In both modalities, we found a significantly smaller displacement magnitude with varying velocities compared to constant velocity. The reduction in displacement occurred irrespective of whether or not the participants pursued the visual targets with their eyes. These findings indicate that the emergence of representational momentum critically depends on the constancy of target velocity. The results are compatible with a model in which changes in the motion signal can override the extrapolation mechanism that usually causes the forward displacement of representational momentum.

Transcranial direct current stimulation of posterior temporal cortex modulates electrophysiological correlates of auditory selective spatial attention in posterior parietal cortex.

Speech perception in "cocktail-party" situations, in which a sound source of interest has to be extracted out of multiple irrelevant sounds, poses a remarkable challenge to the human auditory system. Studies on structural and electrophysiological correlates of auditory selective spatial attention revealed critical roles of the posterior temporal cortex and the N2 event-related potential (ERP) component in the underlying processes. Here, we explored effects of transcranial direct current stimulation (tDCS) to posterior temporal cortex on neurophysiological correlates of auditory selective spatial attention, with a specific focus on the N2. In a single-blind, sham-controlled crossover design with baseline and follow-up measurements, monopolar anodal and cathodal tDCS was applied for 16 min to the right posterior superior temporal cortex. Two age groups of human subjects, a younger (n = 20; age 18-30 yrs) and an older group (n = 19; age 66-77 yrs), completed an auditory free-field multiple-speakers localization task while ERPs were recorded. The ERP data showed an offline effect of anodal, but not cathodal, tDCS immediately after DC offset for targets contralateral, but not ipsilateral, to the hemisphere of tDCS, without differences between groups. This effect mainly consisted in a substantial increase of the N2 amplitude by 0.9 µV (SE 0.4 µV; d = 0.40) compared with sham tDCS. At the same point in time, cortical source localization revealed a reduction of activity in ipsilateral (right) posterior parietal cortex. Also, localization error was improved after anodal, but not cathodal, tDCS. Given that both the N2 and the posterior parietal cortex are involved in processes of auditory selective spatial attention, these results suggest that anodal tDCS specifically enhanced inhibitory attentional brain processes underlying the focusing onto a target sound source, possibly by improved suppression of irrelevant distracters.

Language lateralisation measured across linguistic and national boundaries.

The visual half-field technique has been shown to be a reliable and valid neuropsychological measurement of language lateralisation, typically showing higher accuracy and faster correct responses for linguistic stimuli presented in the right visual field (RVF) than left visual field (LVF). The RVF advantage corresponds to the well-known dominance of the left hemisphere (LH) in processing language(s). However, clinical and experimental neuroscientists around the globe use different variations of the visual half-field paradigm, making direct comparisons difficult. The current study used a word/non-word visual half-field paradigm with translingual stimuli. In total, 496 participants from seven European countries were investigated: Belgium (64), England (49), Germany (85), Italy (34), The Netherlands (87), Norway (51), and Switzerland (126), covering six international languages (Dutch, English, French, German, Italian, Norwegian). All language groups revealed a significant RVF/LH advantage in accuracy and reaction times that accounted for up to 26.1% of the total variance in performance. We found some variation in the degree of the RVF/LH advantage across language groups, accounting for a maximum of 3.7% of the total variance in performance. The RVF/LH advantage did not differ between subsamples speaking English, French or German as first or second languages or between monolingual and early/late bi/multilinguals. The findings suggest that the translingual lexical decision task (TLDT) is a simple but reliable measurement of language lateralisation that can be applied clinically and experimentally across linguistic and national boundaries.

Processing of sound location in human cortex.

This functional magnetic resonance imaging study was focused on the neural substrates underlying human auditory space perception. In order to present natural-like sound locations to the subjects, acoustic stimuli convolved with individual head-related transfer functions were used. Activation foci, as revealed by analyses of contrasts and interactions between sound locations, formed a complex network, including anterior and posterior regions of temporal lobe, posterior parietal cortex, dorsolateral prefrontal cortex and inferior frontal cortex. The distinct topography of this network was the result of different patterns of activation and deactivation, depending on sound location, in the respective voxels. These patterns suggested different levels of complexity in processing of auditory spatial information, starting with simple left/right discrimination in the regions surrounding the primary auditory cortex, while the integration of information on hemispace and eccentricity of sound may take place at later stages. Activations were identified as being located in regions assigned to both the dorsal and ventral auditory cortical streams, that are assumed to be preferably concerned with analysis of spatial and non-spatial sound features, respectively. The finding of activations also in the ventral stream could, on the one hand, reflect the well-known functional duality of auditory spectral analysis, that is, the concurrent extraction of information based on location (due to the spectrotemporal distortions caused by head and pinnae) and spectral characteristics of a sound source. On the other hand, this result may suggest the existence of shared neural networks, performing analyses of auditory 'higher-order' cues for both localization and identification of sound sources.

Cortical processing of location changes in a "cocktail-party" situation: Spatial oddball effects on electrophysiological correlates of auditory selective attention.

Neural mechanisms of selectively attending to a sound source of interest in a simulated "cocktail-party" situation, composed of multiple competing sources, were investigated using event-related potentials in combination with a spatial oddball design. Subjects either detected rare spatial deviants in a series of standard sounds or passively listened. Targets either appeared in isolation or in the presence of two distractor sound sources at different locations ("cocktail-party" condition). Deviant-minus-standard difference potentials revealed mismatch negativity, P3a, and P3b. However, mainly the P3b was modulated by spatial conditions of stimulation, with lower amplitude for "cocktail-party", than single, sounds. In the active condition, cortical source localization revealed two distinct foci of maximum differences in electrical activity for the contrast of single vs. "cocktail-party" sounds: the right inferior frontal junction and the right anterior superior parietal lobule. These areas may be specifically involved in processes associated with selective attention in a "cocktail-party" situation.

More accurate sound localization induced by short-term light deprivation.

Crossmodal reorganization processes in the brain are mainly associated with early blindness, on the assumption that recruitment of genuine visual areas, such as primary visual cortex, for non-visual functions results in superior auditory and tactile performance of blind, compared to sighted, humans. This study shows that in sighted subjects the accuracy of sound localization, measured by a task of head pointing to acoustic targets, is reversibly increased after short-term light deprivation of 90 min. However, only the systematic deviations from target positions (constant error) were reduced after light deprivation, while the general precision of head pointing remained unchanged. Return to pre-deprivation values was observed after 180 min of re-exposure to light. The post-deprivation change was similar, though less in magnitude, to the effect of blindness that was demonstrated previously. Generally, these findings indicate that auditory-visual crossmodal plasticity can be quite rapidly initiated by deprivation of the visual cortex from visual input. It seems possible that visual deprivation has an influence on neuronal circuits, that are involved in processing of auditory information in visual brain areas of normal sighted humans. Since exclusively the constant error in sound localization, not general performance, was changed, the present effect of visual deprivation may, however, not be attributable to reorganization processes in the sense of a compensation for the absence of vision. It is more likely that the observed change in accuracy was specifically induced by the absence of visual calibration of the neural representation of auditory space during light deprivation.

Numerical value biases sound localization.

Speech recognition starts with representations of basic acoustic perceptual features and ends by categorizing the sound based on long-term memory for word meaning. However, little is known about whether the reverse pattern of lexical influences on basic perception can occur. We tested for a lexical influence on auditory spatial perception by having subjects make spatial judgments of number stimuli. Four experiments used pointing or left/right 2-alternative forced choice tasks to examine perceptual judgments of sound location as a function of digit magnitude (1-9). The main finding was that for stimuli presented near the median plane there was a linear left-to-right bias for localizing smaller-to-larger numbers. At lateral locations there was a central-eccentric location bias in the pointing task, and either a bias restricted to the smaller numbers (left side) or no significant number bias (right side). Prior number location also biased subsequent number judgments towards the opposite side. Findings support a lexical influence on auditory spatial perception, with a linear mapping near midline and more complex relations at lateral locations. Results may reflect coding of dedicated spatial channels, with two representing lateral positions in each hemispace, and the midline area represented by either their overlap or a separate third channel.