Selective spatial attention in lateralized multi-talker speech perception: EEG correlates and the role of age.

Speech comprehension under dynamic cocktail party conditions requires auditory search for relevant speech content and focusing spatial attention on the target talker. Here, we investigated the development of these cognitive processes in a population of 329 participants aged 20-70 years. We used a multi-talker speech detection and perception task in which pairs of words (each consisting of a cue and a target word) were simultaneously presented from lateralized positions. Participants attended to predefined cue words and responded to the corresponding target. Task difficulty was varied by presenting cue and target stimuli at different intensity levels. Decline in performance was observed only in the oldest group (age range 53-70 years) and only in the most difficult condition. The EEG analysis of neurocognitive correlates of lateralized auditory attention and stimulus evaluation (N2ac, LPCpc, alpha power lateralization) revealed age-associated changes in focussing on and processing of task-relevant information, while no such deficits were found on early auditory search and target segregation. Irrespective of age, more challenging listening conditions were associated with an increased allocation of attentional resources.

Cross-modal interactions at the audiovisual cocktail-party revealed by behavior, ERPs, and neural oscillations.

Theories of attention argue that objects are the units of attentional selection. In real-word environments such objects can contain visual and auditory features. To understand how mechanisms of selective attention operate in multisensory environments, in this pre-registered study, we created an audiovisual cocktail-party situation, in which two speakers (left and right of fixation) simultaneously articulated brief numerals. In three separate blocks, informative auditory speech was presented (a) alone or paired with (b) congruent or (c) uninformative visual speech. In all blocks, subjects localized a pre-defined numeral. While audiovisual-congruent and uninformative speech improved response times and speed of information uptake according to diffusion modeling, an ERP analysis revealed that this did not coincide with enhanced attentional engagement. Yet, consistent with object-based attentional selection, the deployment of auditory spatial attention (N2ac) was accompanied by visuo-spatial attentional orienting (N2pc) irrespective of the informational content of visual speech. Notably, an N2pc component was absent in the auditory-only condition, demonstrating that a sound-induced shift of visuo-spatial attention relies on the availability of audio-visual features evolving coherently in time. Additional exploratory analyses revealed cross-modal interactions in working memory and modulations of cognitive control.

Attentional modulations of alpha power are sensitive to the task-relevance of auditory spatial information.

The topographical distribution of oscillatory power in the alpha band is known to vary depending on the current focus of spatial attention. Here, we investigated to what extend univariate and multivariate measures of post-stimulus alpha power are sensitive to the required spatial specificity of a task. To this end, we varied the perceptual load and the spatial demand in an auditory search paradigm. A centrally presented sound at the beginning of each trial indicated the to-be-localized target sound. This spatially unspecific pre-cue was followed by a sound array, containing either two (low perceptual load) or four (high perceptual load) simultaneously presented lateralized sound stimuli. In separate task blocks, participants were instructed either to report whether the target was located on the left or the right side of the sound array (low spatial demand) or to indicate the exact target location (high spatial demand). Univariate alpha lateralization magnitude was neither affected by perceptual load nor by spatial demand. However, an analysis of onset latencies revealed that alpha lateralization emerged earlier in low (vs high) perceptual load trials as well as in low (vs high) spatial demand trials. Finally, we trained a classifier to decode the specific target location based on the multivariate alpha power scalp topography. A comparison of decoding accuracy in the low and high spatial demand conditions suggests that the amount of spatial information present in the scalp distribution of alpha-band power increases as the task demands a higher degree of spatial specificity. Altogether, the results offer new insights into how the dynamic adaption of alpha-band oscillations in response to changing task demands is associated with post-stimulus attentional processing.

When long appears short: Effects of auditory distraction on event-related potential correlates of time perception.

Attentional models of time perception assume that the perceived duration of a stimulus depends on the extent to which attentional resources are allocated to its temporal information. Here, we studied the effects of auditory distraction on time perception, using a combined attentional-distraction duration-discrimination paradigm. Participants were confronted with a random sequence of long and short tone stimuli, most of which having a uniform (standard) pitch and only a few a different (deviant) pitch. As observed in previous studies, pitch-deviant tones impaired the discrimination of tone duration and triggered a sequence of event-related potentials (ERPs) reflecting a cycle of deviance detection, involuntary attentional distraction and reorientation (MMN, P3a, RON). Contrasting ERPs of short and long tone durations revealed that long tones elicited a more pronounced fronto-central contingent negative variation (CNV) in the time interval after the expected offset of the short tone as well as a more prominent centro-parietal late positive complex (LPC). Relative to standard-pitch tones, deviant-pitch tones especially impaired the correct discrimination of long tones, which was associated with a reduction of the CNV and LPC. These results are interpreted within the theoretical framework of resource-based models of time perception, in which involuntary distraction due to a deviant event led to a withdrawal of attentional resources from the processing of time information.

Do congruent lip movements facilitate speech processing in a dynamic audiovisual multi-talker scenario? An ERP study with older and younger adults.

In natural conversations, visible mouth and lip movements play an important role in speech comprehension. There is evidence that visual speech information improves speech comprehension, especially for older adults and under difficult listening conditions. However, the neurocognitive basis is still poorly understood. The present EEG experiment investigated the benefits of audiovisual speech in a dynamic cocktail-party scenario with 22 (aged 20-34 years) younger and 20 (aged 55-74 years) older participants. We presented three simultaneously talking faces with a varying amount of visual speech input (still faces, visually unspecific and audiovisually congruent). In a two-alternative forced-choice task, participants had to discriminate target words ("yes" or "no") among two distractors (one-digit number words). In half of the experimental blocks, the target was always presented from a central position, in the other half, occasional switches to a lateral position could occur. We investigated behavioral and electrophysiological modulations due to age, location switches and the content of visual information, analyzing response times and accuracy as well as the P1, N1, P2, N2 event-related potentials (ERPs) and the contingent negative variation (CNV) in the EEG. We found that audiovisually congruent speech information improved performance and modulated ERP amplitudes in both age groups, suggesting enhanced preparation and integration of the subsequent auditory input. In the older group, larger amplitude measures were found in early phases of processing (P1-N1). Here, amplitude measures were reduced in response to audiovisually congruent stimuli. In later processing phases (P2-N2) we found decreased amplitude measures in the older group, while an amplitude reduction for audiovisually congruent compared to visually unspecific stimuli was still observable. However, these benefits were only observed as long as no location switches occurred, leading to enhanced amplitude measures in later processing phases (P2-N2). To conclude, meaningful visual information in a multi-talker setting, when presented from the expected location, is shown to be beneficial for both younger and older adults.

EEG correlates of spatial shifts of attention in a dynamic multi-talker speech perception scenario in younger and older adults.

Speech perception under "cocktail-party" conditions critically depends on the focusing of attention toward the talker of interest. In dynamic auditory scenes, changes in talker settings require rapid shifts of attention, which is especially relevant when the position of a target talker switches from one location to another. Here, we explored electrophysiological correlates of shifts in spatial auditory attention, using a free-field speech perception task, in which sequences of short words (a company name, followed by a numeric value, e.g., "Bosch-6") were presented in the participants' left and right horizontal plane. Younger and older participants responded to the value of a pre-defined target company, while ignoring three simultaneously presented pairs of concurrent company names and values from different locations. All four stimulus pairs were spoken by different talkers, alternating from trial-to-trial. The location of the target company was within either the left or right hemisphere for a variable number of consecutive trials (between 3 and 42 trials) and then changed, switching from the left to the right hemispace or vice versa. Thus, when a switch occurred, the participants had to search for the new position of the target company among the concurrent streams of auditory information and re-focus their attention on the relevant location. As correlates of lateralized spatial auditory attention, the anterior contralateral N2 subcomponent (N2ac) and the posterior alpha power lateralization were analyzed in trials immediately before and after switches of the target location. Both measures were increased after switches, while only the increase in N2ac was related to better speech perception performance (i.e., a reduced post-switch decline in accuracy). While both age groups showed a similar pattern of switch-related attentional modulations, N2ac and alpha lateralization to the task-relevant stimulus (the target company's value) was overall greater in the younger, than older, group. The results suggest that N2ac and alpha lateralization reflect different attentional processes in multi-talker speech perception, the first being primarily associated with auditory search and the focusing of attention, and the second with the in-depth attentional processing of task-relevant information. Especially the second process appears to be prone to age-related cognitive decline.

A dual mechanism underlying retroactive shifts of auditory spatial attention: dissociating target- and distractor-related modulations of alpha lateralization.

Attention can be allocated to mental representations to select information from working memory. To date, it remains ambiguous whether such retroactive shifts of attention involve the inhibition of irrelevant information or the prioritization of relevant information. Investigating asymmetries in posterior alpha-band oscillations during an auditory retroactive cueing task, we aimed at differentiating those mechanisms. Participants were cued to attend two out of three sounds in an upcoming sound array. Importantly, the resulting working memory representation contained one laterally and one centrally presented item. A centrally presented retro-cue then indicated the lateral, the central, or both items as further relevant for the task (comparing the cued item(s) to a memory probe). Time-frequency analysis revealed opposing patterns of alpha lateralization depending on target eccentricity: A contralateral decrease in alpha power in target lateral trials indicated the involvement of target prioritization. A contralateral increase in alpha power when the central item remained relevant (distractor lateral trials) suggested the de-prioritization of irrelevant information. No lateralization was observed when both items remained relevant, supporting the notion that auditory alpha lateralization is restricted to situations in which spatial information is task-relevant. Altogether, the data demonstrate that retroactive attentional deployment involves excitatory and inhibitory control mechanisms.

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.

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.

Visually guided auditory attention in a dynamic "cocktail-party" speech perception task: ERP evidence for age-related differences.

Speech understanding in the presence of concurring sound is a major challenge especially for older persons. In particular, conversational turn-takings usually result in switch costs, as indicated by declined speech perception after changes in the relevant target talker. Here, we investigated whether visual cues indicating the future position of a target talker may reduce the costs of switching in younger and older adults. We employed a speech perception task, in which sequences of short words were simultaneously presented by three talkers, and analysed behavioural measures and event-related potentials (ERPs). Informative cues resulted in increased performance after a spatial change in target talker compared to uninformative cues, not indicating the future target position. Especially the older participants benefited from knowing the future target position in advance, indicated by reduced response times after informative cues. The ERP analysis revealed an overall reduced N2, and a reduced P3b to changes in the target talker location in older participants, suggesting reduced inhibitory control and context updating. On the other hand, a pronounced frontal late positive complex (f-LPC) to the informative cues indicated increased allocation of attentional resources to changes in target talker in the older group, in line with the decline-compensation hypothesis. Thus, knowing where to listen has the potential to compensate for age-related decline in attentional switching in a highly variable cocktail-party environment.

Driver state examination--Treading new paths.

A large proportion of crashes in road driving can be attributed to driver fatigue. Several types of fatigue are discussed, comprising sleep-related fatigue, active task-related fatigue (as a consequence of workload in demanding driving situations) as well as passive task-related fatigue (as related to monotonous driving situations). The present study investigated actual states of fatigue in a monotonous driving situation, using EEG measures and a long-lasting driving simulation experiment, in which drivers had to keep the vehicle on track by compensating crosswind of different strength. Performance data and electrophysiological correlates of mental fatigue (EEG Alpha and Theta power, Inter Trial Coherence (ITC), and auditory event-related potentials to short sound stimuli) were analyzed. Driving errors and driving lane variability increased with time on task and with increasing crosswind. The posterior Alpha and Theta power also increased with time on task, but decreased with stronger crosswind. The P3a to sound stimuli decreased with time on task when the crosswind was weak, but remained stable when the crosswind was strong. The analysis of ITC revealed less frontal Alpha and Theta band synchronization with time on task, but no effect of crosswind. The results suggest that Alpha power in monotonous driving situations reflects boredom or attentional withdrawal due to monotony rather than the decline of processing abilities as a consequence of high mental effort. A more valid indicator of declining mental resources with increasing time on task seems to be provided by brain oscillatory synchronization measures and event-related activity.

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.

Electrophysiological correlates of cocktail-party listening.

Detecting, localizing, and selectively attending to a particular sound source of interest in complex auditory scenes composed of multiple competing sources is a remarkable capacity of the human auditory system. The neural basis of this so-called "cocktail-party effect" has remained largely unknown. Here, we studied the cortical network engaged in solving the "cocktail-party" problem, using event-related potentials (ERPs) in combination with two tasks demanding horizontal localization of a naturalistic target sound presented either in silence or in the presence of multiple competing sound sources. Presentation of multiple sound sources, as compared to single sources, induced an increased P1 amplitude, a reduction in N1, and a strong N2 component, resulting in a pronounced negativity in the ERP difference waveform (N2d) around 260 ms after stimulus onset. About 100 ms later, the anterior contralateral N2 subcomponent (N2ac) occurred in the multiple-sources condition, as computed from the amplitude difference for targets in the left minus right hemispaces. Cortical source analyses of the ERP modulation, resulting from the contrast of multiple vs. single sources, generally revealed an initial enhancement of electrical activity in right temporo-parietal areas, including auditory cortex, by multiple sources (at P1) that is followed by a reduction, with the primary sources shifting from right inferior parietal lobule (at N1) to left dorso-frontal cortex (at N2d). Thus, cocktail-party listening, as compared to single-source localization, appears to be based on a complex chronology of successive electrical activities within a specific cortical network involved in spatial hearing in complex situations.

What does successful speech-in-noise perception in aging depend on? Electrophysiological correlates of high and low performance in older adults.

Aging usually decreases the ability to understand language under difficult listening conditions. However, aging is also associated with increased between-subject variability. Here, we studied potential sources of inter-individual differences and investigated spoken language understanding of younger and older adults (age ranges 21-35 and 57-74 years, respectively) in a simulated "cocktail-party" scenario. A naturalistic "stock-price monitoring" task was employed in which prices of listed companies were simultaneously recited by four speakers at different locations in space. The participants responded when prices of a target company exceeded specific values, while ignoring all other companies. According to their individual performance levels three subgroups of participants were composed, consisting of 12 high-performing and 12 low-performing older adults, and 12 young adults matching the high-performing older group. The analysis of the event-related brain potentials indicated that all older adults showed delayed attentional control (indicated by a later P2) and reduced speech processing (indicated by a reduced N400), relative to the younger adults. High-performing older adults differed in increased allocation of attention and inhibitory control (indicated by a stronger P2-N2 complex) from their low-performing counterparts. The results are consistent with the idea of an adjustment of mental resources that could help compensating potential deficiencies in peripheral and central auditory processing.

ERP correlates of auditory goal-directed behavior of younger and older adults in a dynamic speech perception task.

The ability to understand speech under adverse listening conditions deteriorates with age. In addition to genuine hearing deficits, age-related declines in attentional and inhibitory control are assumed to contribute to these difficulties. Here, the impact of task-irrelevant distractors on speech perception was studied in 28 younger and 24 older participants in a simulated "cocktail party" scenario. In a two-alternative forced-choice word discrimination task, the participants responded to a rapid succession of short speech stimuli ("on" and "off") that was presented at a frequent standard location or at a rare deviant location in silence or with a concurrent distractor speaker. Behavioral responses and event-related potentials (mismatch negativity MMN, P3a, and reorienting negativity RON) were analyzed to study the interplay of distraction, orientation, and refocusing in the presence of changes in target location. While shifts in target location decreased performance of both age groups, this effect was more pronounced in the older group. Especially in the distractor condition, the electrophysiological measures indicated a delayed attention capture and a delayed re-focussing of attention toward the task-relevant stimulus feature in the older group, relative to the young group. In sum, the results suggest that a delay in the attention switching mechanism contribute to the age-related difficulties in speech perception in dynamic listening situations with multiple speakers.

Allocentric or craniocentric representation of acoustic space: an electrotomography study using mismatch negativity.

The world around us appears stable in spite of our constantly moving head, eyes, and body. How this is achieved by our brain is hardly understood and even less so in the auditory domain. Using electroencephalography and the so-called mismatch negativity, we investigated whether auditory space is encoded in an allocentric (referenced to the environment) or craniocentric representation (referenced to the head). Fourteen subjects were presented with noise bursts from loudspeakers in an anechoic environment. Occasionally, subjects were cued to rotate their heads and a deviant sound burst occurred, that deviated from the preceding standard stimulus either in terms of an allocentric or craniocentric frame of reference. We observed a significant mismatch negativity, i.e., a more negative response to deviants with reference to standard stimuli from about 136 to 188 ms after stimulus onset in the craniocentric deviant condition only. Distributed source modeling with sLORETA revealed an involvement of lateral superior temporal gyrus and inferior parietal lobule in the underlying neural processes. These findings suggested a craniocentric, rather than allocentric, representation of auditory space at the level of the mismatch negativity.

Cortical processing of change in sound location: smooth motion versus discontinuous displacement.

The detection of a change in space is an essential prerequisite of adequate responding to dynamic aspects of our auditory environment, be it induced by smooth motion or abrupt and discontinuous displacements of ongoing sound sources. Here, we investigated the auditory processing of different types of spatial change using electroencephalography. While fixating straight ahead, participants listened to a free-field sound stimulus that, after an initial stationary phase in a central position, either (1) started to move horizontally (motion stimulus), or (2) changed repetitively its spatial position in random order within left or right hemispaces (scatter stimulus), or (3) shifted abruptly toward a lateral position (displacement stimulus). Irrespective of stimulus type, the onset of spatial change elicited a characteristic sequence of auditory evoked potentials that was similar to the so-called motion-onset response described in previous studies. Differences in response amplitudes and latencies to the different stimulus types were only gradual, with scatter and displacement producing generally stronger responses than motion. Also, inter-hemispheric asymmetry patterns in the responses to scatter differed somewhat from those obtained with the other types of changes in position. Rather than any substantial stimulus-specific differences, the findings suggest the existence of an auditory "spatial change response," that is, a common electrophysiological correlate of auditory processing of any spatial change in the environment that does not necessarily need to involve motion as such.

When and where of auditory spatial processing in cortex: a novel approach using electrotomography.

The modulation of brain activity as a function of auditory location was investigated using electro-encephalography in combination with standardized low-resolution brain electromagnetic tomography. Auditory stimuli were presented at various positions under anechoic conditions in free-field space, thus providing the complete set of natural spatial cues. Variation of electrical activity in cortical areas depending on sound location was analyzed by contrasts between sound locations at the time of the N1 and P2 responses of the auditory evoked potential. A clear-cut double dissociation with respect to the cortical locations and the points in time was found, indicating spatial processing (1) in the primary auditory cortex and posterodorsal auditory cortical pathway at the time of the N1, and (2) in the anteroventral pathway regions about 100 ms later at the time of the P2. Thus, it seems as if both auditory pathways are involved in spatial analysis but at different points in time. It is possible that the late processing in the anteroventral auditory network reflected the sharing of this region by analysis of object-feature information and spectral localization cues or even the integration of spatial and non-spatial sound features.

Understanding of spoken language under challenging listening conditions in younger and older listeners: a combined behavioral and electrophysiological study.

Numerous studies suggested an age-related decline in speech perception under difficult listening conditions. Here, spoken language understanding of two age groups of listeners was investigated in a naturalistic "stock price monitoring" task. Stock prices of listed companies were simultaneously recited by three speakers at different positions in space and presented via headphones to 14 younger and 14 older listeners (age ranges 19-25 and 54-64 years, respectively). The listeners had to respond when prices of target companies exceeded a specific value, but to ignore all other prices as well as beep sounds randomly interspersed within the stock prices. Older listeners did not produce more missing responses, or longer response times than younger listeners. However, differences in event-related potentials indicated a reduced parietal P3b of older, relative to younger, listeners. Separate analyses for those listeners who performed relatively high or low in the behavioral task revealed a right-frontal P3a that was pronounced especially in the group of high-performing older listeners. Correlational analyses indicated a direct relationship between P3a amplitude and spoken language comprehension in older, but not younger, listeners. Furthermore, younger (especially, low-performing) listeners showed a more pronounced P2 on irrelevant beep sounds than older listeners. These subtle differences in cortical processing between age groups suggest that high performance of older middle-aged listeners in demanding listening situations is associated with increased engagement of frontal brain areas, and thus the allocation of mental resources for compensation of potential declines in spoken language understanding.

Auditory motion perception: onset position and motion direction are encoded in discrete processing stages.

The neural processing of auditory motion information shows a pronounced interhemispheric asymmetry. In previous electrophysiological studies, the so-called motion-onset response (MOR), a prominent auditory-evoked potential to the onset of sound motion, was stronger over the hemisphere contralateral to the side of motion. Here, effects of lateral-onset position and direction of motion on MOR contralaterality were investigated. Eighteen listeners were presented with free-field sound stimuli that, after an initial stationary phase at a lateral spatial position within the left or right hemifield, started to move either left- or rightward. The early part of the MOR, the so-called change-N1, exhibited contralaterality that depended on the lateral motion-onset position with stronger activations over the hemisphere contralateral to the side of motion onset, whereas the contralaterality of the later part of the MOR, the so-called change-P2, merely depended on the direction of motion. Cortical source localization indicated that this pattern of contralaterality primarily resulted from asymmetric activation in primary auditory cortex and insula. These findings suggest that the early and late parts of the MOR reflect different phases in auditory motion perception, supporting the notion of a modular organization of discrete processing stages.