Modulation of visual perception by eye gaze direction in patients with spatial neglect and extinction.

Parietal neurons have retinotopic receptive fields whose response is modulated by eye gaze signals. We examined the role of eye position and eye movement direction on visual extinction in three patients with parietal damage and left neglect, characterized by impaired perception of contralesional field stimuli despite intact visual cortex. Patients tracked a horizontally moving fixation point, while discriminating fine changes at the fixated point and detecting more salient targets flashed on either side of fixation. Eye position modulated extinction, with worse detection of left field targets during eccentric fixation towards the left. Direction of eye movement also modulated extinction, with worse detection of left targets during rightward scanning. These results demonstrate that extinction in contralesional field is influenced by extra-retinal factors related to eye position and eye movements, consistent with a convergence of these signals in parietal cortex.     

Infants' attention is biased by emotional expressions and eye gaze direction

This study investigates infants' processing of emotional expressions in combination with referential eye gaze cues. In experiment 1, 7-month-old infants' neural responses to fearful and neutral faces, which were looking at a novel object, were assessed. Infants' attention, as indexed by the negative central component of the event-related potential, was enhanced when the adult gazed at the object with a fearful expression compared with a neutral expression. In experiment 2, no effect of emotion on amplitude of the negative central was found when the face directed eye gaze at the infant and away from the object. We conclude that by 7 months, infants use emotional expressions in triadic person-object-person contexts to detect threat in the environment.     

Watching where you look: modulation of visual processing of foveal stimuli by spatial attention

Two experiments investigated the effect of sustained selective spatial attention upon the perceptual analysis of stimuli within the center of gaze. Spatial attention has typically been studied in relation to peripheral stimuli, and its relevance to the processing of central stimuli has remained relatively ignored. Here, behavioral measures in normal human volunteer participants showed that focused spatial attention can also influence responses to central stimuli, over and beyond the advantage conferred by their foveation. Event-related potentials (ERPs) showed that the action of attention upon foveal stimuli included the modulation of perceptual processing in extrastriate visual areas. Surprisingly, the visual modulation revealed an intriguing and consistent pattern of hemispheric asymmetry, in both experiments. These findings show that in addition to the established right hemisphere dominance of the brain areas that direct attention, the consequences of directed attention may also be asymmetrical.     

A left visual field advantage in perception of gaze direction

Previous work has found a left visual field (LVF) advantage for various judgements on faces, including identity and emotional expression. This has been related to possible right-hemisphere specialisation for face processing, and it has been proposed that this might reflect configural processing. We sought to determine whether a similar LVF advantage may also exist for gaze perception. In two experiments, normal adult subjects made judgements for seen gaze direction (left, right or straight). To assess how visual field may influence perception of gaze direction, eye stimuli were briefly presented unilaterally or bilaterally. In the latter case, the gaze direction of the two seen eyes could be congruent or incongruent (i.e. the two eyes could gaze in the same or different directions). For unilateral displays, performance was more accurate for LVF stimuli than RVF. With bilateral incongruent gaze, the LVF eye influenced judgements more strongly than the RVF eye. No such LVF advantage was found in a control experiment, in which subjects judged pupil size for similar eye stimuli. Taken together, these results reveal a LVF advantage for perception of gaze direction. Since only the eye region was visible, our results cannot be due to a LVF bias in processing the entire face context. Instead they suggest lateralisation specifically in processing the direction of seen gaze.     

Extinguishing extinction: hemispheric differences in the modulation of TMS-induced visual extinction by directing covert spatial attention

The topic of spatial attention is of great relevance for researchers in various fields, including neuropsychology, cognitive neuroscience, and cognitive psychology, as well as for clinical practice. Deficits of spatial attentional arising from parietal brain damage remain largely confined to the left visual field. The mechanisms underlying this hemispheric asymmetry are still elusive. We mimicked the neuropsychological syndrome of contralesional extinction by temporarily inducing a spatial attentional bias in healthy volunteers with TMS. We investigated whether directing covert spatial attention could enhance or, more importantly, counteract the resulting behavioral deficits. Although both the left and right parietal TMS induced contralateral extinction, only left hemifield extinction following right parietal TMS was severely aggravated by a competing stimulus in the ipsilesional (right) hemifield. We put forward the hypothesis that an asymmetry with respect to the ability of detaching attention from a distractor is contributing to the right hemispheric lateralization with regard to extinction. On a broader level, we suggest that "virtual patients" might be used for evaluating neuropsychological treatment in an early stage of development, reducing the burden on actual patients.     

Temporal modulation of spatial tactile extinction in right-brain-damaged patients

Unilateral and bilateral electrotactile stimuli were delivered to both hands of 11 right-brain-damaged (RBD) patients with left tactile extinction and 20 healthy subjects. Bimanual stimuli could be presented simultaneously or with varying stimulus onset asynchronies (SOAs). Subjects indicated their detection of unilateral or bilateral stimuli, their judgements of whether stimuli were simultaneous or successive, and, in the latter case, which side came first. In RBD patients, extinction was maximal with simultaneous presentations and decreased as SOA increased. With short SOAs, omissions of left-sided stimuli occurred with both right-side and left-side stimulus precedence, suggesting a forward and backward interference of the right stimulus on the processing of the left stimulus within a time window of at least 100 msec. In contrast, there was no interference of the left stimulus on the detection of the right stimulus. Unlike controls, extinction patients rarely expressed simultaneity judgements, but those that were produced tended to be veridical or nearly so, like in normal controls. Whereas controls expressed generally accurate judgements of right or left precedence, patients showed a bias toward a right precedence and a maximal uncertainty between left-first and right-first choices when the left stimulus had a lead between 100 and 200 msec. The results are consistent with the hypothesis that, as in other sensory modalities, tactile extinction is associated with an abnormal persistent bias of attention toward the ipsilesional side that delays the processing of contralesional stimuli. However, the finding that both extinction and explicit judgements of simultaneity tended to occur with simultaneous bilateral stimuli suggest the presence of some residual neural capacity to detect precise temporal coincidence.     

Looking you in the mouth: abnormal gaze in autism resulting from impaired top-down modulation of visual attention

People with autism are impaired in their social behavior, includingtheir eye contact with others, but the processes that underliethis impairment remain elusive. We combined high-resolutioneye tracking with computational modeling in a group of 10 high-functioningindividuals with autism to address this issue. The group fixatedthe location of the mouth in facial expressions more than didmatched controls, even when the mouth was not shown, even infaces that were inverted and most noticeably at latencies of200–400 ms. Comparisons with a computational model ofvisual saliency argue that the abnormal bias for fixating themouth in autism is not driven by an exaggerated sensitivityto the bottom-up saliency of the features, but rather by anabnormal top-down strategy for allocating visual attention.     

Supramodal effects of covert spatial orienting triggered by visual or tactile events

Event-related functional magnetic resonance imaging was used to identify brain areas involved in spatial attention and determine whether these operate unimodally or supramodally for vision and touch. On a trial-by-trial basis, a symbolic auditory cue indicated the most likely side for the subsequent target, thus directing covert attention to one side. A subsequent target appeared in vision or touch on the cued or uncued side. Invalidly cued trials (as compared with valid trials) activated the temporo-parietal junction and regions of inferior frontal cortex, regardless of target modality. These brain areas have been associated with multimodal spatial coding in physiological studies of the monkey brain and were linked to a change in the location that must be attended to in the present study. The intraparietal sulcus and superior frontal cortex were also activated in our task, again, regardless of target modality, but did not show any specificity for invalidly cued trials. These results identify a supramodal network for spatial attention and reveal differential activity for inferior circuits involving the temporo-parietal junction and inferior frontal cortex (specific to invalid trials) versus more superior intraparietal-frontal circuits (common to valid and invalid trials).     

Nonspatial cueing of tactile STM causes shift of spatial attention

The focus of attention can be flexibly altered in mnemonic representations of past sensory events. We investigated the neural mechanisms of selection in tactile STM by applying vibrotactile sample stimuli of different intensities to both hands, followed by a symmetrically shaped visual retro-cue. The retro-cue indicated whether the weak or strong sample was relevant for subsequent comparison with a single tactile test stimulus. Locations of tactile stimuli were randomized, and the required response did not depend upon the spatial relation between cued sample and test stimulus. Selection between spatially segregated items in tactile STM was mirrored in lateralized activity following visual retro-cues (N2pc) and influenced encoding of task-irrelevant tactile probe stimuli (N140). Our findings support four major conclusions. First, retrospective selection results in transient shifts of spatial attention. Second, retrospective selection is functionally dissociable from attention-based rehearsal of locations. Third, selection mechanisms are linked across processing stages, as attention shifts in STM influence encoding of sensory signals. Fourth, selection in tactile STM recruits attentional control mechanisms that are, at least partially, supramodal.     

Split of spatial attention as predicted by a systems-level model of visual attention

Can we attend to multiple distinct spatial locations at the same time? According to a recent psychophysical study [J. Dubois et?al. (2009)Journal of Vision, 9, 3.1-11] such a split of spatial attention might be limited to short periods of time. Following N. P. Bichot et?al. [(1999)Perception & Psychophysics, 61, 403-423] subjects had to report the identity of multiple letters that were briefly presented at different locations, while two of these locations (targets) were relevant for a concurrent shape comparison task. In addition to the design used by Bichot et?al. stimulus onset asynchrony between shape onset and letters was systematically varied. In general, the performance of subjects was superior at target locations. Furthermore, for short stimulus onset asynchronies, performance was simultaneously increasing at both target locations. For longer stimulus onset asynchronies, however, performance deteriorated at one of the target locations while increasing at the other target location. It was hypothesized that this dynamic deployment of attention might be caused by competitive processes in saccade-related structures such as the frontal eye field. Here we simulated the task of Dubois et?al. using a systems-level model of attention. Our results are consistent with recent findings in the frontal eye field obtained during covert visual search, and they support the view of a transient deployment of spatial attention to multiple stimuli in the early epoch of target selection.     

Spatial attention triggered by eye gaze increases and speeds up early visual activity

What are the neuronal correlates of reflexive shifts of attention triggered by eye gaze direction? Event related potentials (ERPs) were measured on 14 subjects performing a spatial attention task where eye gaze direction of a face cued the location of a forthcoming target. Subjects were faster in detecting a validly cued target, i.e. one appearing at the location the eye was gazing at, compared to invalidly cued targets, despite the non-predictive value of the eye cues. ERP results showed an enhanced and earlier occipito-parietal P1 and N1 for valid trials, demonstrating the early modulation of visual input by attentional allocation. These findings provide the first evidence that social attention can rapidly modify the processing of visual information in extrastriate cortex.     

Perceived gaze direction in faces and spatial attention: a study in patients with parietal damage and unilateral neglect

Perceived gaze in faces is an important social signal that may influence orienting of attention in normal observers. Would such effects of gaze still occur in patients with right parietal damage and left neglect who usually fail to attend to contralesional space? Two experiments tested for effects of perceived gaze on visual extinction. Face or shape stimuli were presented in the right, left, or both hemifields, with faces looking either straight ahead or toward the opposite field. On bilateral trials, patients extinguished a left shape much less often when a concurrent right face looked leftward rather than straight ahead. This occurred, even though gaze was not relevant to the task and processing of facial signals implied attention to a competing ipsilesional stimulus. By contrast, rightward gaze in faces presented on the left side had no effect on extinction, suggesting that gaze cues are not extracted without attention. Two other experiments examined effects of perceived gaze on the detection of peripheral targets. Targets appeared at one of four possible locations to the right or left of a central face looking either toward the target location, another location on the same side, the opposite side, or straight ahead. Face and gaze were not relevant to the task and not predictive of target location. Patients responded faster when the face looked toward the target on both the contralesional and ipsilesional sides. In contralesional space, gaze allowed shifting of attention in a specific quadrant direction, but only to the first target along the scan path when there were different possible locations on the same side. By contrast, in intact ipsilesional space, attention was selectively directed to one among different eccentric locations. Control experiments showed that symbolic arrow cues did not produce similar effects. These results indicate that even though parietal damage causes spatial neglect and impairs the representation of location on the contralesional side, perceived gaze in faces can still trigger automatic shifts of attention in the contralesional direction, suggesting the existence of specific and anatomically distinct attentional mechanisms.     

Rapid targeting followed by sustained deployment of visual spatial attention

We investigated preparatory attention processes when a spatial discrimination was required at a cued location, by measuring electroencephalography following a central symbolic cue to deploy spatial attention. Electroencephalography activity in response to the cue revealed three cue-related activations: an early-onset positivity following the P1 at posterior scalp sites contralateral to the cued location, followed by cue-related frontal scalp activity and later-onset sustained activity at posterior scalp sites contralateral to the cued location. The early contralateral positivity may reflect rapid targeting of the cued location. Our results also extend the findings of cue-related frontal activity followed by posterior activity contralateral to the cued location, found with nonspatial feature discriminations, to a task requiring a spatial discrimination.     

Neuronal modulation of auditory attention by informative and uninformative spatial cues

Sounds provide important information about the spatial environment, including the location of approaching objects. Attention to sounds can be directed through automatic or more controlled processes, which have been well studied in the visual modality. However, little is known about the neural underpinnings of attentional control mechanisms for auditory signals. We studied healthy adults who underwent event-related FMRI while performing a task that manipulated automatic and more controlled auditory orienting by varying the probability that cues correctly predicted target location. Specifically, we examined the effects of uninformative (50% validity ratio) and informative (75% validity ratio) auditory cues on reaction time (RT) and neuronal functioning. The stimulus-onset asynchrony (SOA) between the cue and the target was either 100 or 800 ms. At the 100 ms SOA, RT was faster for valid than invalid trials for both cue types, and frontoparietal activation was greater for invalid than valid trials. At the 800 ms SOA, RT and functional activation depended on whether cues were informative or uninformative, and whether cues correctly or incorrectly predicted the target location. Contrary to our prediction, activation in a frontoparietal network was greater for uninformative than informative cues across several different comparisons and at both SOAs. This finding contrasts with similar research of visual orienting, and suggests that the auditory modality may be more biased toward automatic shifts of attention following uninformative cues.     

Perception of static eye gaze direction facilitates subsequent early visual processing.

OBJECTIVE: Using event-related potentials (ERPs), it has been recently shown that a reflexive shift of attention following the observation of a dynamic eye gaze cue enhances and speeds up early visual processing of a target presented at the gazed-at location. Here we investigate whether similar early sensory modulations are also elicited by static gaze cues, or if previously described attentional effects were caused mainly by visual motion cues and not by eye gaze direction per se. Furthermore, we explore if these possible attentional orienting effects reflect facilitation of the processing of cued stimuli, inhibition of the unattended stimuli, or both. METHODS: Subjects were presented with a face looking to the right or left visual field (VF), or straight away, before the occurrence of a lateralized target to detect. There were 3 conditions in this nonpredictive cueing task: (1) target presented in the VF indicated by the eye gaze direction (congruent); (2) opposite to the eye gaze direction (incongruent); or (3) preceded by a straight gazing face (neutral). RESULTS: Subjects were faster at detecting congruently than incongruently and neutrally cued targets. Facilitation effects were observed on early ERP components: the occipital P1 and occipito-temporal N1 components were speeded up as early as approximately 100 ms following stimulus onset (P1), and enhanced (P1 and N1) in response to congruent trials, particularly in the right hemisphere. CONCLUSIONS: Spatial attention triggered by static eye gaze direction produces response facilitations - predominantly lateralized to the right hemisphere - from the early sensory stages of visual processing. Significance: This study provides the first evidence of a speeding up and amplification of early visual processing following attention triggered by static eye gaze perception.     

Luminance and spatial attention effects on early visual processing

Event-related potentials (ERPs) were recorded from healthy subjects in response to unilaterally flashed high and low luminance bar stimuli presented randomly to left and right field locations. Their task was to covertly and selectively attend to either the left or right stimulus locations (separate blocks) in order to detect infrequent shorter target bars of either luminance. Independent of attention, higher stimulus luminance resulted in higher ERP amplitudes for the posterior N95 (80–110 ms), occipital P1 (110–140 ms), and parietal N1 (130–180 ms). Brighter stimuli also resulted in shorter peak latency for the occipital N1 component (135–220 ms); this effect was not observed for the N1 components over parietal, central or frontal regions. Significant attention-related amplitude modulations were obtained for the occipital P1, occipital, parietal and central N1, the occipital and parietal P2, and the parietal N2 components; these components were larger to stimuli at the attended location. In contrast to the relatively short latencies of both spatial attention and luminance effects, the first interaction between luminance and spatial attention effects was observed for the P3 component to the target stimuli (350–750 ms). This suggests that interactions of spatial attention and stimulus luminance previously reported for reaction time measures may not reflect the earliest stages of sensory/perceptual processing. Differences in the way in which luminance and attention affected the occipital P1, occipital N1 and parietal N1 components suggest dissociations among these ERPs in the mechanisms of visual and attentional processing they reflect. Nonetheless, scalp current density mappings of the attention effects throughout the latency ranges of the P1 and N1 components show the most prominent attention-related activity to be in lateral occipital scalp areas. Such a pattern is consistent with the spatially selective filtering of information into the ventral stream of visual processing which is reponsible for complex feature analysis and object identification.     

Influence of final search direction on tactile line bisection in normal subjects

BACKGROUND: Many variables influence the tactile bisection performance of normal subjects. Of these, studies that investigated the starting point effect have reported inconsistent results. OBJECTIVE: To determine if the final search direction rather than the initial search direction (starting point) may have an effect on tactile bisection performance, especially when subjects are making multiple searches. METHODS: The authors divided the experiment into single- and multiple-search tasks. Thirty-two blindfolded normal subjects were asked to indicate the midpoint of a rod after a one-way search (single-search task) or after two or more one-way searches (multiple-search task). In each task, the subject started on either end of the horizontally placed rods. The same procedure was also conducted with rods oriented vertically or radially. Therefore, there were six conditions (two starting points x three orientations). RESULTS: In the single-search task, there was a significant effect of starting point or movement direction in all six conditions, with biases occurring toward the starting point from the true midpoint (this is termed the "overshoot phenomenon"). In the multiple-search task, however, there was no significant effect of the starting point in all six conditions. Rather, biases depended on the final movement direction in five of the six conditions, occurring toward the starting point of the final search. CONCLUSION: Future research is needed to understand the mechanism of the overshoot phenomenon in tactile bisection by normal subjects and how the overshoot phenomenon influences patients with neglect.     

Differential modulation of word recognition by semantic and spatial orienting of attention

In the present study, we investigated the ability to orient attention to abstract associative features of complex stimuli, more specifically, to the semantic categories of visual word stimuli. We compared the behavioral and electrophysiological effects of semantic orienting with those elicited by spatial orienting to word stimuli. Two parallel, cued lexical-decision tasks, with semantic- or spatial-orienting cues, were used. Results showed that both semantic and spatial orienting facilitated behavioral performance. The event-related potential analysis revealed different and non-overlapping patterns of modulation of word processing by semantic and spatial orienting. Modulation by semantic orienting started later, affecting only the potentials linked to conceptual or semantic processing (N300 and N400). The pattern of N300/N400 modulation in the semantic-orienting condition was similar to that observed in semantic-priming tasks, and was compatible with the operation of controlled semantic processes. Spatial orienting significantly enhanced the amplitude of the early visual potential P1 as well as the language-related N400 potential. These findings showed that the similar end-result of behavioral facilitation by semantic and spatial orienting is achieved through largely distinct mechanisms acting upon separate levels of stimulus analysis.     

Effects of feature and spatial attention on visual change detection

In event-related brain potential studies using a visual S1-S2 matching task, stimulus changes elicit change-related positivity, which reflects the detection of visual changes. To investigate the effects of attention on change detection, we tested the elicitation of change-related positivity in response to changes in color and spatial frequency under three attention conditions: (i) changes in an unattended feature at an attended location, (ii) in an attended feature at an unattended location, and (iii) in an unattended feature at an unattended location. The results suggest that stimulus changes can be detected even when both feature and spatial attention are withdrawn, but change detection can also be inhibited, which might be because of biased-competition determined by the combination of feature and spatial attention conditions.