Biases of spatial attention in vision and audition.

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.

"Touched" by light: event-related potentials (ERPs) to visuo-haptic stimuli in peri-personal space.

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.

Sex differences in auditory processing in peripersonal space: an event-related potential study.

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.

Psychophysiology and imaging of visual cortical functions in the blind: a review.

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.

Spatial attention facilitates selection of illusory objects: evidence from event-related brain potentials.

The relationship between spatial attention and object-based attention has long been debated. On the basis of behavioral evidence it has been hypothesized that these two forms of attention share a common mechanism, such that directing spatial attention to one part of an object facilitates the selection of the entire object. In a previous study (Martinez, A., Teder-Salejarvi, W., Vazquez, M., Molholm, S., Foxe, J.J., Javitt, D.C., Di Russo, F., Worden, M.S., Hillyard, S.A., 2006. "Objects are highlighted by spatial attention." J. Cogn. Neurosci. 18(2): 298-310) we used recordings of event-related potentials (ERPs) during a paradigm modeled after that of Egly et al. (Egly, R., Driver, J., Rafal, D.R., 1994. Shifting visual attention between objects and locations: evidence from normal and parietal lesion subjects. J. Exp. Psychol. Gen. 123(2) 161-77) to investigate this relationship. As reported in numerous studies of spatial attention, we found the typical pattern of enhanced neural activity in visual cortex elicited by attended stimuli. Unattended stimuli belonging to the same object as the attended stimuli elicited a very similar spatiotemporal pattern of enhanced neural activity that was localized to lateral occipital cortex (LOC). This similarity was taken as evidence that spatial- and object-selective attention share, at least in part, a common neural mechanism. In the present study we further investigate this relationship by examining whether this spread of spatial attention within attended objects can be guided by objects defined by illusory contours. Subjects viewed a display consisting of two illusory rectangular objects and directed attention to continuous sequences of stimuli (brief onsets) at one end of one of the objects. Stimuli occurring at irrelevant locations but belonging to the same attended object elicited larger posterior N1 amplitudes than that elicited by unattended objects forming part of a different object. This object-selective N1 enhancement was localized to lateral occipital cortex. The present data support the hypothesis that the allocation of spatial attention can be guided by illusory object boundaries and that this allocation strengthens the perceptual representations of attended objects at the level of visual area LOC.

The temporal window of integration in elderly and young adults.

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.

Auditory spatial tuning in late-onset blindness in humans.

Blind individuals who lost their sight as older children or adults were compared with normally sighted controls in their ability to focus auditory spatial attention and to localize sounds in a noisy acoustic environment. Event-related potentials (ERPs) were recorded while participants attended to sounds presented in free field from either central or peripheral arrays of speakers with the task of detecting infrequent targets at the attended location. When attending to the central array of speakers, the two groups detected targets equally well, and their spatial tuning curves for both ERPs and target detections were highly similar. By contrast, late blind participants were significantly more accurate than sighted participants at localizing sounds in the periphery. For both groups, the early N1 amplitude to peripheral standard stimuli displayed no significant spatial tuning. In contrast, the amplitude of the later P3 elicited by targets/deviants displayed a more sharply tuned spatial gradient during peripheral attention in the late blind than in the sighted group. These findings were compared with those of a previous study of congenitally blind individuals in the same task [Röder, B., Teder-Sälejärvi, W., Sterr, A., Rösler, F., Hillyard, S. A., & Neville, H. J. Improved auditory spatial tuning in blind humans. Nature, 400, 162-166, 1999]. It was concluded that both late blind and congenitally blind individuals demonstrate an enhanced capability for focusing auditory attention in the periphery, but they do so via different mechanisms: whereas congenitally blind persons demonstrate a more sharply tuned early attentional filtering, manifested in the N1, late blind individuals show superiority in a later stage of target discrimination and recognition, indexed by the P3.

Control mechanisms mediating shifts of attention in auditory and visual space: a spatio-temporal ERP analysis.

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.

Neural basis of auditory-induced shifts in visual time-order perception.

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.

Auditory spatial localization and attention deficits in autistic adults.

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.

Long-term exposure to noise impairs cortical sound processing and attention control.

Long-term exposure to noise impairs human health, causing pathological changes in the inner ear as well as other anatomical and physiological deficits. Numerous individuals are daily exposed to excessive noise. However, there is a lack of systematic research on the effects of noise on cortical function. Here we report data showing that long-term exposure to noise has a persistent effect on central auditory processing and leads to concurrent behavioral deficits. We found that speech-sound discrimination was impaired in noise-exposed individuals, as indicated by behavioral responses and the mismatch negativity brain response. Furthermore, irrelevant sounds increased the distractibility of the noise-exposed subjects, which was shown by increased interference in task performance and aberrant brain responses. These results demonstrate that long-term exposure to noise has long-lasting detrimental effects on central auditory processing and attention control.

Human auditory cortex tracks task-irrelevant sound sources.

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.

Preattentive evaluation of multiple perceptual streams in human audition.

To analyze an auditory scene, the segregation of the input into separate streams of information is necessary. Here, the mismatch negativity (MMN) of the event-related potential was used to trace the number of simultaneously monitored streams in auditory scene analysis. Subjects passively listened to sounds belonging to either one, two, or three auditory streams defined by spatial position and pitch. An MMN was obtained for deviants within either one, two or three auditory streams albeit with a smaller amplitude in the latter condition. Control experiments ruled out that this effect was due to variations in stimulation rate. Thus, a preattentive deviant detection system based on simple auditory features provides the basis for subsequent attentive processing directed to a selected stream.

Neural substrates of perceptual enhancement by cross-modal spatial attention.

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.