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Neurologically normal observers misperceive the midpoint of horizontal lines as systematically leftward of veridical center, a phenomenon known as pseudoneglect. Pseudoneglect is attributed to a tonic asymmetry of visuospatial attention favoring left hemispace. Whereas visuospatial attention is biased toward left hemispace, some evidence suggests that audiospatial attention may possess a right hemispatial bias. If spatial attention is supramodal, then the leftward bias observed in visual line bisection should also be expressed in auditory bisection tasks. If spatial attention is modality specific then bisection errors in visual and auditory spatial judgments are potentially dissociable. Subjects performed a bisection task for spatial intervals defined by auditory stimuli, as well as a tachistoscopic visual line bisection task. Subjects showed a significant leftward bias in the visual line bisection task and a significant rightward bias in the auditory interval bisection task. Performance across both tasks was, however, significantly positively correlated. These results imply the existence of both modality specific and supramodal attentional mechanisms where visuospatial attention has a prepotent leftward vector and audiospatial attention has a prepotent rightward vector of attention. In addition, the biases of both visuospatial and audiospatial attention are correlated., (2010 Elsevier Inc. All rights reserved.)

Parietal neuronal populations have been found which respond bimodally to visual and somatosensory input regarding one's own limbs or even perceived haptic input of a false limb (Graziano et al., Science 290:1782-1785, 2000). Further, neuronal populations have been observed which respond preferentially to visual stimuli presented in spatial congruence with our hands (Graziano, Proc Natl Acad Sci USA 96:10418-10421, 1999). In this study, we examined event-related potentials (ERPs) elicited by laser dots projected onto or above participants' index and middle fingers during a sustained-attention task. We hypothesized that visual stimuli projected onto the hand would elicit differences in ERP deflections related to sensory gating and categorization in comparison to when projected close to the hand. Participants responded via a footswitch to rare target flashes of light occurring on or directly above the middle finger of the attended hand. We found enhanced amplitudes of the N1 and P3 deflections when the stimuli fell onto the finger tips as opposed to above them. Furthermore, the N1 for unattended stimuli was less suppressed when the lasers were projected onto the fingers. Behaviorally, participants were less accurate to targets when the lasers fell onto the fingers. We conclude that when the lasers appear to "touch" the participants, they act to automatically draw participants' attention. Thus visual stimuli projected onto the fingers of the 'unattended' hand are harder to filter out, leading to decreases in accuracy during task performance.

Further processing of auditory stimuli in the free field is attenuated when participants are in contact with speakers versus not touching them. Studies in the visual domain have found that men and women use different strategies for processing spatial information. In this study, we examined sex-related differences in event-related potentials while men and women performed an auditory discrimination task in peripersonal space when either holding speakers or resting their hands in their laps. We found that men responded more accurately than women to targets in attended locations, and that the sexes exhibited different event-related potential patterns during task performance. These differences are consistent with existing predictions of female top-down and male bottom-up strategies in spatial processing.

Multisensory integration and interaction occur when bimodal stimuli are presented as either spatially congruent or incongruent, but temporally coincident. We investigated whether proprioceptive cues interact with auditory attention to one of two sound sources in free-field. The participant's task was to attend to either the left or right speaker and to respond to occasional increased-bandwidth targets via a footswitch. We recorded high-density EEG in three experimental conditions: the participants either held the speakers in their hands (Hold), reached out close to them (Reach), or had their hands in their lap (Lap). In the last two conditions, the auditory event-related potentials (ERPs) revealed a prominent negativity around 200 ms post-stimulus (N2 wave) over fronto-central areas, which is a reliable index of further processing of spatial stimulus features in free-field. The N2 wave was markedly attenuated in the 'Hold' condition, which suggests that proprioceptive cues apparently solidify spatial information computed by the auditory system, in so doing alleviating the need for further processing of spatial coordinates solely based on auditory information.

Imaging, transcranial magnetic stimulation, and psychophysiological recordings of the congenitally blind have confirmed functional activation of the visual cortex but have not extensively explained the functional significance of these activation patterns in detail. This review systematically examines research on the role of the visual cortex in processing spatial and non-visual information, highlighting research on individuals with early and late onset blindness. Here, we concentrate on the methods utilized in studying visual cortical activation in early blind participants, including positron emissions tomography (PET), functional magnetic resonance imaging (fMRI), transcranial magnetic stimulation (TMS), and electrophysiological data, specifically event-related potentials (ERPs). This paper summarizes and discusses findings of these studies. We hypothesize how mechanisms of cortical plasticity are expressed in congenitally in comparison to adventitiously blind and short-term visually deprived sighted participants and discuss potential approaches for further investigation of these mechanisms in future research.

Integration of information across time is an essential part of auditory processing. Evidence from a variety of experiments support the notion of an approximately 200-ms long time window following the onset of a sound, during which a unitary sound representation is formed (the temporal window of integration, TWI). The temporal resolution in the auditory system is assumed to decrease with aging suggesting that the duration of the TWI may be longer in elderly than young adults. The TWI duration was assessed in young and elderly adults using the oddball paradigm in which a regular auditory event (standard) is occasionally exchanged for a different event (deviant). Previous studies showed that when the stimulus onset asynchrony (SOA) exceeds the duration of the TWI, two successive deviations occurring infrequently in a repetitive sound sequence elicit two separate mismatch negativity (MMN) event-related brain potentials. However, only one MMN is elicited when the SOA is shorter than the TWI. Experiment 1 tested MMN elicitation for the second of two successive deviant sounds as a function of the SOA. Experiment 2 used the sound omission paradigm, in which MMN is only elicited by omissions when the SOA is shorter than the TWI. Again, MMN elicitation was tested by infrequent tone omissions as a function of the SOA. Results showed no significant differences between elderly and younger participants as a function of SOA. This suggests that the duration of the TWI is approximately between 200 and 250 ms in both groups of subjects. On the other hand, the lower MMN amplitudes elicited by frequency deviation in the elderly compared with the younger participants suggest that the specificity of frequency representation deteriorates with aging.

The neural systems that mediate voluntary shifts of attention to visual and auditory stimuli were investigated by examining the patterns of human brain electricity elicited by attention-directing cues in auditory and visual tasks. Several lateralized event-related potential (ERP) components were observed when participants shifted attention in expectation of visual targets (experiment 1). One component was focused over frontal cortex and a second was focused primarily over the occipital-temporal cortex but also spread to parietal regions of the scalp. Previous work has indicated that the frontal component reflects supramodal processes involved in the executive control of attention and that the posterior component reflects either spatial attentional control processes in the posterior parietal lobe or modulation of processes in visual cortex. Here, the posterior component was observed when participants shifted attention in expectation of auditory targets (experiments 2-4), but the frontal component was found only in the visual task. The posterior component seemed to be generated in parietal and occipital areas even when there was no visual information about the to-be-attended locations. These results are consistent with the view that voluntary shifts of attention are mediated by supramodal processes in the parietal lobe and that the spatial coordinates of the to-be-attended location are based on visual representations of space.

Attended objects are perceived to occur before unattended objects even when the two objects are presented simultaneously. This finding has led to the widespread view that attention modulates the speed of neural transmission in the various perceptual pathways. We recorded event-related potentials during a time-order judgment task to determine whether a reflexive shift of attention to a sudden sound modulates the speed of sensory processing in the human visual system. Attentional cueing influenced the perceived order of lateralized visual events but not the timing of event-related potentials in visual cortex. Attentional cueing did, however, enhance the amplitude of neural activity in visual cortex, which shows that attention-induced shifts in visual time-order perception can arise from modulations of signal strength rather than processing speed in the early visual-cortical pathways.

Spatial constraints on multisensory integration of auditory (A) and visual (V) stimuli were investigated in humans using behavioral and electrophysiological measures. The aim was to find out whether cross-modal interactions between A and V stimuli depend on their spatial congruity, as has been found for multisensory neurons in animal studies (Stein & Meredith, 1993). Randomized sequences of unimodal (A or V) and simultaneous bimodal (AV) stimuli were presented to right- or left-field locations while subjects made speeded responses to infrequent targets of greater intensity that occurred in either or both modalities. Behavioral responses to the bimodal stimuli were faster and more accurate than to the unimodal stimuli for both same-location and different-location AV pairings. The neural basis of this cross-modal facilitation was studied by comparing event-related potentials (ERPs) to the bimodal AV stimuli with the summed ERPs to the unimodal A and V stimuli. These comparisons revealed neural interactions localized to the ventral occipito-temporal cortex (at 190 msec) and to the superior temporal cortical areas (at 260 msec) for both same- and different-location AV pairings. In contrast, ERP interactions that differed according to spatial congruity included a phase and amplitude modulation of visual-evoked activity localized to the ventral occipito-temporal cortex at 100-400 msec and an amplitude modulation of activity localized to the superior temporal region at 260-280 msec. These results demonstrate overlapping but distinctive patterns of multisensory integration for spatially congruent and incongruent AV stimuli.

The objective of this study was to compare autistic adults and matched control subjects in their ability to focus attention selectively on a sound source in a noisy environment. Event-related brain potentials (ERPs) were recorded while subjects attended to a fast paced sequence of brief noise bursts presented in free-field at a central or peripheral location. Competing sequences of noise bursts at adjacent locations were to be ignored. Both behavioral measures of target detection and auditory ERP amplitudes indicated that control subjects were able to focus their attention more sharply on the relevant sound source than autistic subjects. These findings point to a fundamental deficit in the spatial focusing of auditory attention in autism, which may be a factor that impedes social interactions and sensory-guided behavior, particularly in noisy environments.

The brain organizes sound into coherent sequences, termed auditory streams. We asked whether task-irrelevant sounds would be detected as separate auditory streams in a natural listening environment that included three simultaneously active sound sources. Participants watched a movie with sound while street-noise and sequences of naturally varying footstep sounds were presented in the background. Occasional deviations in the footstep sequences elicited the mismatch negativity (MMN) event-related potential. The elicitation of MMN showed that the regular features of the footstep sequences had been registered and their violations detected, which could only occur if the footstep sequence had been detected as a separate auditory stream. Our results demonstrate that sounds are organized into auditory streams irrespective of their relevance to ongoing behavior.

Orienting attention involuntarily to the location of a sudden sound improves perception of subsequent visual stimuli that appear nearby. The neural substrates of this cross-modal attention effect were investigated by recording event-related potentials to the visual stimuli using a dense electrode array and localizing their brain sources through inverse dipole modeling. A spatially nonpredictive auditory precue modulated visual-evoked neural activity first in the superior temporal cortex at 120-140 msec and then in the ventral occipital cortex of the fusiform gyrus 15-25 msec later. This spatio-temporal sequence of brain activity suggests that enhanced visual perception produced by the cross-modal orienting of spatial attention results from neural feedback from the multimodal superior temporal cortex to the visual cortex of the ventral processing stream.

AC amplifiers can introduce significant distortions into the low frequency and DC components of recorded electrophysiological data such as event-related potentials (ERPs). Methods for correcting such distortions (i.e., estimating the waveform of the original data) after the data have been amplified and recorded rely on an accurate estimate of the amplifier's time constant (TC). We show that the filter characteristics of AC amplifiers in at least some commercially available ERP recording instruments may vary considerably across individual channels, even when each houses an identical AC amplifier circuit. Clearly, distortion correction methods must take this variability into account. We propose an empirical means of estimating the correct TC value. This approach yields more accurate correction than those based on TCs calculated analytically.

Crossmodal integration was studied in humans by presenting random sequences of auditory (brief noise bursts), visual (flashes), and audiovisual (simultaneous noise bursts and flashes) stimuli from a central location at irregular intervals between 600 and 800 ms. The subjects' task was to press a button to infrequent and unpredictable (P=0.15) target stimuli that could be either a more intense noise burst, a brighter flash, or a combination of the two. In accordance with previous studies, behavioral data showed that bimodal target stimuli were responded to much faster and were identified more accurately than the unimodal target stimuli. The neural basis of this crossmodal interaction was investigated by subtracting the ERPs to the auditory (A) and the visual (V) stimuli alone from the ERP to the combined audiovisual (AV) stimuli (i.e. interaction=AV-(A+V)). Using this approach, we replicated previous reports of both early (at around 40 ms) and late (after 100 ms) ERP interaction effects. However, it appears that the very early interaction effects can be largely accounted for by an anticipatory ERP that precedes both the unimodal and bimodal stimuli. In calculating the ERP interaction this slow shift is subtracted twice, resulting in an apparent shift of the opposite polarity that may be confounded with actual crossmodal interactions.

Orienting attention involuntarily to the location of a sensory event influences responses to subsequent stimuli that appear in different modalities with one possible exception: orienting attention involuntarily to a sudden light sometimes fails to affect responses to subsequent sounds (e.g., Spence & Driver, 1997). Here we investigated the effects of involuntary attention to a brief flash on the processing of subsequent sounds in a design that eliminates stimulus-response compatibility effects and criterion shifts as confounding factors. In addition, the neural processes mediating crossmodal attention were studied by recording event-related brain potentials. Our data show that orienting attention to the location of a spatially nonpredictive visual cue modulates behavioural and neural responses to subsequent auditory targets when the stimulus onset asynchrony is short (between 100 and 300 ms). These findings are consistent with the hypothesis that involuntary shifts of attention are controlled by supramodal brain mechanisms rather than by modality-specific ones.

Event-related brain potentials were recorded from healthy human subjects while they attended to one of two auditory stimulus channels (defined by location, left and right of a fixation point, and pitch) in order to detect rare target events. The distracting properties of periodic noise (vs. continuous noise, experiment 1) and backward speech (vs. forward speech, experiment 2) presented from a third speaker located behind the subjects were investigated. A typical attention effect with a larger negativity for attended tones was observed in both experiments. Backward speech led to a significantly reduced target detection rate for the first four stimuli after onset of the distractor accompanied by a reduced event-related brain potential (ERP)-attention effect and a reduced fronto-central N2b component for the target stimuli. This indicates that irrelevant information leads to an attention decrement of about 1 s duration.

To perceive real-world objects and events, we need to integrate several stimulus features belonging to different sensory modalities. Although the neural mechanisms and behavioural consequences of intersensory integration have been extensively studied, the processes that enable us to pay attention to multimodal objects are still poorly understood. An important question is whether a stimulus in one sensory modality automatically attracts attention to spatially coincident stimuli that appear subsequently in other modalities, thereby enhancing their perceptual salience. The occurrence of an irrelevant sound does facilitate motor responses to a subsequent light appearing nearby. However, because participants in previous studies made speeded responses rather than psychophysical judgements, it remains unclear whether involuntary auditory attention actually affects the perceptibility of visual stimuli as opposed to postperceptual decision and response processes. Here we provide psychophysical evidence that a sudden sound improves the detectability of a subsequent flash appearing at the same location. These data show that the involuntary orienting of attention to sound enhances early perceptual processing of visual stimuli.

Young adult subjects attended selectively to brief noise bursts delivered in free-field via central and peripheral arrays of four loudspeakers each that were arranged along a semi-circle extending from the midline to 90 degrees right of center. Frequent "standard" stimuli (90%) and infrequent "target/deviant" stimuli (10%) of increased bandwidth were delivered at a fast rate in random order and equiprobably from all eight speakers. In separate runs, the subject's task was to selectively attend to the center or rightmost speaker, and to press a button to the infrequent "target" stimuli occurring at the designated (spatial) location. Behavioral detection rates and concurrently recorded event-related potentials (ERPs) indicated that auditory attention was deployed as a finely tuned gradient around the attended source. The attentional gradients were steeper for the central than the peripheral array, indicating that attention can be more sharply focused upon sound sources directly in front of the listener. The ERP data suggested that selection for location is accomplished in two distinct stages, with an initial broadly tuned filtering, followed by a more narrowly focused selection of attended-location deviants.

This study investigated cross-modal interactions in spatial attention by means of recording event-related brain potentials (ERPs). Noise bursts and light flashes were presented in random order to both left and right field locations separated by 60 degrees in free-field. One group of subjects was instructed to attend selectively to the noise bursts (attend-auditory group), and a second group attended only to the flashes (attend-visual group). On different runs attention was directed to either the right or left field stimuli of the designated modality. In the attend-auditory group, noise bursts at the attended location elicited a broad, biphasic negativity (Nd) beginning at 70 ms. The cross-modal spatial attention effect on the auditory ERPs in the attend-visual group was very similar in morphology, but the Nd was reduced in amplitude relative to the intra-modal effect. In the attend-visual group, flashes at the attended location elicited enhanced early (100-200 ms) and late (200-350 ms) ERP components relative to unattended-location flashes. The cross-modal effect in the attend-auditory group included small but significant enhancements of early components of the visual ERPs. It was concluded that spatial attention has a multi-modal organization such that the processing of stimuli at attended locations is facilitated at an early, sensory level, even for stimuli of an unattended modality.

Young adult subjects attended selectively to brief noise bursts delivered in free field via a horizontal array of seven loudspeakers spaced apart by 9 degrees of angle. Frequent "standard" stimuli (90%) and infrequent "target/deviant" stimuli (10%) of increased bandwidth were delivered at a fast rate in a random sequence equiprobably from each speaker. In separate runs, the subjects' task was to selectively attend to the leftmost, center, or rightmost speaker and to press a button to the infrequent "target" stimuli occurring at the designated spatial location. Behavioral detection rates and concurrently recorded event-related potentials (ERPs) indicated that auditory attention was deployed as a finely tuned gradient around the attended sound source, thus providing support for gradient models of auditory spatial attention. Furthermore, the ERP data suggested that the spatial focusing of attention was achieved in two distinct stages, with an early more broadly tuned filtering of inputs occurring over the first 80-200 msec after stimulus onset, followed by a more narrowly focused selection of attended-location deviants that began at around 250 msec and closely resembled the behavioral gradient of target detections.

Steady-state visual evoked potentials (SSVEPs) were recorded from the scalp of subjects who attended to a flickering LED display in one visual field while ignoring a similar display (flickering at a different frequency) in the opposite visual field. The flicker frequencies were 20.8 Hz in the left-field display and 27.8 Hz in the right-field display. The SSVEP to the flicker in either field was enhanced in amplitude when attention was directed to its location. The scalp distribution of this SSVEP enhancement was narrowly focused over the posterior scalp contralateral to the visual field of stimulation. A source analysis using Variable Resolution Electromagnetic Tomography (VARETA) indicated that the source current densities for the SSVEP attention effect had a focal origin in the contralateral parieto-occipital cortex., (Copyright 1998 Elsevier Science B.V.)

Recordings of electrical and magnetic brain responses to sensory stimulation provide high-resolution measures of the time course of early perceptual processing. Spatio-temporal analyses of brain activity patterns during the first 200 ms after stimulus presentation have characterized the timing of attentional selection processes and different stages of feature encoding and pattern analyses. Recent studies that incorporate blood flow neuroimaging techniques provide support for mechanisms of early selection of attended visual inputs in extrastriate cortical pathways. The spatial tuning properties of early auditory selection have also been delineated. Electrical and magnetic responses that index the encoding of higher-order pattern information have been identified in both visual and auditory modalities and localized to specific cortical areas.