Spatial Attention

Event-related brain potentials (ERPs) provide high-resolution measures of the time course of neuronal activity patterns associated with perceptual and cognitive processes. New techniques for ERP source analysis and comparisons with data from blood-f low neuroimaging studies enable improved localization of cortical activity during visual selective attention. ERP modulations during spatial attention point toward a mechanism of gain control over information f low in extrastriate visual cortical pathways, starting about 80 ms after stimulus onset. Paying attention to nonspatial features such as color, motion, or shape is manifested by qualitatively different ERP patterns in multiple cortical areas that begin with latencies of 100-150 ms. The processing of nonspatial features seems to be contingent upon the prior selection of location, consistent with early selection theories of attention and with the hypothesis that spatial attention is ''special.''

We demonstrate a previously unknown gender difference in the distribution of spatial attention, a basic capacity that supports higher-level spatial cognition. More remarkably, we found that playing an action video game can virtually eliminate this gender difference in spatial attention and simultaneously decrease the gender disparity in mental rotation ability, a higher-level process in spatial cognition. After only 10 hr of training with an action video game, subjects realized substantial gains in both spatial attention and mental rotation, with women benefiting more than men. Control subjects who played a nonaction game showed no improvement. Given that superior spatial skills are important in the mathematical and engineering sciences, these findings have practical implications for attracting men and women to these fields.

The panoramic scenes of the world contain more information than we can take in with a single glance. To examine the fine details of a visual scene, we must rely on the brain's capability to focus attention in a spatially selective manner and thereby facilitate the perception of stimuli within a restricted zone of the visual field 1,2 . This covert focusing of attention has been likened metaphorically to a 'spotlight' 3 or 'zoom lens' 4 that can be shifted to relevant locations even when the eyes remain stationary. Psychophysical experiments have shown that stimuli falling within the spotlight of attention are detected and discriminated more rapidly and accurately than stimuli at unattended locations 1-4 .

The syndrome of spatial neglect is typically associated with focal injury to the temporoparietal or ventral frontal cortex. This syndrome shows spontaneous partial recovery, but the neural basis of both spatial neglect and its recovery is largely unknown. We show that spatial attention deficits in neglect (rightward bias and reorienting) after right frontal damage correlate with abnormal activation of structurally intact dorsal and ventral parietal regions that mediate related attentional operations in the normal brain. Furthermore, recovery of these attention deficits correlates with the restoration and rebalancing of activity within these regions. These results support a model of recovery based on the re-weighting of activity within a distributed neuronal architecture, and they show that behavioral deficits depend not only on structural changes at the locus of injury, but also on physiological changes in distant but functionally related brain areas.

The syndrome of contralesional neglect reflects a lateralized disruption of spatial attention. In the human, the left hemisphere shifts attention predominantly in the contralateral hemispace and in a contraversive direction whereas the right hemisphere distributes attention more evenly, in both hemispaces and both directions. As a consequence of this asymmetry, severe contralesional neglect occurs almost exclusively after right hemisphere lesions. Patients with left neglect experience a loss of salience in the mental representation and conscious perception of the left side and display a reluctance to direct orientating and exploratory behaviours to the left. Neglect is distributed according to egocentric, allocentric, world-centred, and object-centred frames of reference. Neglected events can continue to exert an implicit influence on behaviour, indicating that the attentional filtering occurs at the level of an internalized representation rather than at the level of peripheral sens...

Natural scenes contain more information than the human visual system can efficiently process at once. Visual attention is the perceptual mechanism by which observers select important aspects of a scene for further cognitive processing 1-5 . Evidence from neurophysiology and functional neuroimaging shows that the neural representation of a visual object is suppressed when it is presented along with other competing objects, relative to when it is presented alone, reflecting competitive interactions among sensory representations. This competitive suppression is thought to occur through inhibitory neural connections. When an object is attended through the action of a top-down biasing signal, this suppression is effectively lifted, and the item is 'selected' 6-8 .

Attention typically amplifies neuronal responses evoked by task-relevant stimuli while attenuating responses to task-irrelevant distracters. In this context, visual distracters constitute an external source of noise that is diminished to improve attended signal quality. Activity that is internal to the cortex itself, stimulus-independent ongoing correlated fluctuations in firing, might also act as task-irrelevant noise. To examine this, we recorded from area V4 of macaques performing an attention-demanding task. The firing of neurons to identically repeated stimuli was highly variable. Much of this variability originates from ongoing low-frequency (<5 Hz) fluctuations in rate correlated across the neuronal population. When attention is directed to a stimulus inside a neuron's receptive field, these correlated fluctuations in rate are reduced. This attentiondependent reduction of ongoing cortical activity improves the signal-to-noise ratio of pooled neural signals substantially more than attention-dependent increases in firing rate.

Functional MRI was used to test whether instructing subjects to attend to one or another location in a visual scene would affect neural activity in human primary visual cortex. Stimuli were moving gratings restricted to a pair of peripheral, circular apertures, positioned to the right and to the left of a central fixation point. Subjects were trained to perform a motion discrimination task, attending (without moving their eyes) at any moment to one of the two stimulus apertures. Functional MRI responses were recorded while subjects were cued to alternate their attention between the two apertures. Primary visual cortex responses in each hemisphere modulated with the alternation of the cue; responses were greater when the subject attended to the stimuli in the contralateral hemifield. The attentional modulation of the brain activity was about 25% of that evoked by alternating the stimulus with a uniform field.

We investigated the spatio-temporal dynamic of attentional bias towards fearful faces. Twelve participants performed a covert spatial orienting task while recording visual event-related brain potentials (VEPs). Each trial consisted of a pair of faces (one emotional and one neutral) briefly presented in the upper visual field, followed by a unilateral bar presented at the location of one of the faces. Participants had to judge the orientation of the bar. Comparing VEPs to bars shown at the location of an emotional (valid) versus neutral (invalid) face revealed an early effect of spatial validity: the lateral occipital P1 component (~130 ms post-stimulus) was selectively increased when a bar replaced a fearful face compared to when the same bar replaced a neutral face. This effect was not found with upright happy faces or inverted fearful faces. A similar amplification of P1 has previously been observed in electrophysiological studies of spatial attention using non-emotional cues. In a behavioural control experiment, participants were also better at discriminating the orientation of the bar when it replaced a fearful rather than a neutral face. In addition, VEPs time-locked to the face-pair onset revealed a C1 component (~90 ms) that was greater for fearful than happy faces. Source localization (LORETA) confirmed an extrastriate origin of the P1 response showing a spatial validity effect, and a striate origin of the C1 response showing an emotional valence effect. These data suggest that activity in primary visual cortex might be enhanced by fear cues as early as 90 ms post-stimulus, and that such effects might result in a subsequent facilitation of sensory processing for a stimulus appearing at the same location. These results provide evidence for neural mechanisms allowing rapid, exogenous spatial orienting of attention towards fear stimuli.

have demonstrated a non-spatial, feature-based attentional modulation of neuronal responses in direction-Ontario M3J 1P3 Canada selective cells of the macaque cortical middle temporal (MT) area . The magnitude of neuronal responses to 2 Cognitive Neuroscience Laboratory German Primate Center an unattended stimulus (the distractor) located inside the cells' receptive field (RF) and moving in the cells' Goettingen 37077 Germany preferred direction was dependent on the direction of an attended stimulus (the target) even though the target stimulus was located outside the RF. When attention was directed toward a target moving in the preferred Summary direction, responses were higher than when it was directed to the anti-preferred direction. This finding led to Background: Attending to the spatial location or to nonthe proposal of a feature-similarity gain model in which spatial features of visual stimuli can modulate neuronal attention changes responses in a multiplicative fashion, responses in primate visual cortex. The modulation by with the sign and strength of the modulation reflecting spatial attention changes the gain of sensory neurons the similarity between the attended stimulus feature and and strengthens the representation of attended locathe neuron's preferred feature [2]. (We use the term tions without changing neuronal selectivities such as "feature" to denote a particular value of a stimulus didirectionality, i.e., the ratio of responses to preferred mension. For example, two motion directions represent and anti-preferred directions of motion. Whether featwo different features).

Despite reports of improved auditory discrimination capabilities in blind humans and visually deprived animals, there is no general agreement as to the nature or pervasiveness of such compensatory sensory enhancements. Neuroimaging studies have pointed out differences in cerebral organization between blind and sighted humans, but the relationship between these altered cortical activation patterns and auditory sensory acuity remains unclear. Here we compare behavioural and electrophysiological indices of spatial tuning within central and peripheral auditory space in congenitally blind and normally sighted but blindfolded adults to test the hypothesis (raised by earlier studies of the effects of auditory deprivation on visual processing) that the effects of visual deprivation might be more pronounced for processing peripheral sounds. We find that blind participants displayed localization abilities that were superior to those of sighted controls, but only when attending to sounds in pe...

Recordings of event-related potentials (ERPs) were combined with structural and functional magnetic resonance imaging (fMRI) to study the spatio-temporal patterns of cortical activity that underlie visual-spatial attention. Small checkerboard stimuli were flashed in random order to the four quadrants of the visual field at a rapid rate while subjects attended to stimuli in one quadrant at a time. Attended stimuli elicited enhanced ERP components in the latency range 80-200 ms that were co-localized with fMRI activations in multiple extrastriate cortical regions. The earliest ERP component (C1 at 50-90 ms) was unaffected by attention and was localized by dipole modeling to calcarine cortex. A longer latency deflection in the 150-225 ms range that was accounted for by this same calcarine source, however, did show consistent modulation with attention. This late attention effect, like the C1, inverted in polarity for upper versus lower field stimuli, consistent with a neural generator in primary visual cortex (area V1). These results provide support to current hypotheses that spatial attention in humans is associated with delayed feedback to area V1 from higher extrastriate areas that may have the function of improving the salience of stimuli at attended locations.

Advance information about an impending stimulus facilitates its subsequent identification and ensuing behavioral responses. This facilitation is thought to be mediated by top-down control signals from frontal and parietal cortex that modulate sensory cortical activity. Here we show, using Granger causality measures on blood oxygen level-dependent time series, that frontal eye field (FEF) and intraparietal sulcus (IPS) activity predicts visual occipital activity before an expected visual stimulus. Top-down levels of Granger causality from FEF and IPS to visual occipital cortex were significantly greater than both bottom-up and mean cortex-wide levels in all individual subjects and the group. In the group and most individual subjects, Granger causality was significantly greater from FEF to IPS than from IPS to FEF, and significantly greater from both FEF and IPS to intermediate-tier than lower-tier ventral visual areas. Moreover, top-down Granger causality from right IPS to intermediate-tier areas was predictive of correct behavioral performance. These results suggest that FEF and IPS modulate visual occipital cortex, and FEF modulates IPS, in relation to visual attention. The current approach may prove advantageous for the investigation of interregional directed influences in other human brain functions.

We used functional magnetic resonance imaging (fMRI) to investigate the functional anatomical relationship between covert orienting of visual spatial attention and execution of saccadic eye movements. Brain areas engaged by shifting spatial attention covertly and by moving the eyes repetitively toward visual targets were compared and contrasted directly within the same subjects. The two tasks activated highly overlapping neural systems and showed that common parietal and frontal regions are more activated during the covert task than the overt oculomotor condition. The possible nature of the relationship between these two operations is discussed.

Temporal orienting of attention is the ability to focus resources at a particular moment in time in order to optimise behaviour, and is associated with activation of left parietal and premotor cortex [Coull, J. T., Nobre, A. C. Where and when to pay attention: the neural systems for directing attention to spatial locations and to time intervals as revealed by both PET and fMRI. Journal of Neuroscience, 1998, 18, 7426±7435]. In the present experiment, we explored the behavioural and anatomical correlates of temporal orienting to foveal visual stimuli, in order to eliminate any spatial attention confounds. We implemented a two-way factorial design in an event-related fMRI study to examine the factors of trial validity (predictability of target by cue), length of delay (cue-target interval), and their interaction. There were two distinct types of invalid trial: those where attention was automatically drawn to a premature target and those where attention was voluntarily shifted to a delayed time-point. Reaction times for valid trials were shorter than those for invalid trials, demonstrating appropriate allocation of attention to temporal cues. All trial-types activated a shared system, including frontoparietal areas bilaterally, showing that this network is consistently associated with attentional orienting and is not speci®c to spatial tasks. Distinct brain areas were sensitive to cuetarget delays and to trial validity. Long cue-target intervals activated areas involved in motor preparation: supplementary motor cortex, basal ganglia and thalamus. Invalid trials, where temporal expectancies were breached, showed enhanced activation of left parietal and frontal areas, and engagement of orbitofrontal cortex bilaterally. Finally, trial validity interacted with length of delay. Appearance of targets prematurely selectively activated visual extrastriate cortex; while postponement of target appearance selectively activated right prefrontal cortex. These ®ndings suggest that distinct brain areas are involved in redirecting attention based upon sensory events (bottom-up, exogenous shifts) and based upon cognitive expectations (top-down, endogenous shifts). #

The extent to which brief abrupt-onset visual stimuli involuntarily capture spatial attention was examined in five experiments. The paradigm used was intended to maximize the opportunity and incentive for subjects to ignore abrupt-onset distractor stimuli in nontarget locations. Subjects made a speeded two-choice response to a target letter appearing in one of four boxes. An abrupt-onset visual stimulus, easily discriminable from the target, was flashed briefly prior to the presentation of a target. In separate blocks, the flash stimulus marked the box in which the target would subsequently appear (SAME), a different box (DIFF), fixation (CENTER), or all four boxes (ALL). Prior to each block, subjects were informed of the flash-target relationship. In all five experiments, response time was elevated in the DIFF, CENTER, and ALL conditions. The interference effect was larger for the DIFF condition and persisted for longer flash-target SOAs. These results suggest that, under appropriate conditions, spatial attention can be involuntarily drawn to abrupt-onset events despite the intention of subjects' to ignore them.

Human event-related potentials (ERPs) were recorded from 10 subjects presented with visual target and nontarget stimuli at five screen locations and responding to targets presented at one of the locations. The late positive response complexes of 25-75 ERP average waveforms from the two task conditions were simultaneously analyzed with Independent Component Analysis, a new computational method for blindly separating linearly mixed signals. Three spatially fixed, temporally independent, behaviorally relevant, and physiologically plausible components were identified without reference to peaks in single-channel waveforms. A novel frontoparietal component (P3f) began at ϳ140 msec and peaked, in faster responders, at the onset of the motor command. The scalp distribution of P3f appeared consistent with brain regions activated during spatial orienting in functional imaging experiments. A longer-latency large component (P3b), positive over parietal cortex, was followed by a postmotor potential (Pmp) component that peaked 200 msec after the button press and reversed polarity near the central sulcus. A fourth component associated with a left frontocentral nontarget positivity (Pnt) was evoked primarily by target-like distractors presented in the attended location. When no distractors were presented, responses of five fasterresponding subjects contained largest P3f and smallest Pmp components; when distractors were included, a Pmp component appeared only in responses of the five slower-responding subjects. Direct relationships between component amplitudes, latencies, and behavioral responses, plus similarities between component scalp distributions and regional activations reported in functional brain imaging experiments suggest that P3f, Pmp, and Pnt measure the time course and strength of functionally distinct brain processes.

We investigated the effects of spatial-selective attention on oscillatory neuronal dynamics in a tactile delayed-match-to-sample task. Whole-head magnetoencephalography was recorded in healthy subjects while dot patterns were presented to their index fingers using Braille stimulators. The subjects' task was to report the reoccurrence of an initially presented sample pattern in a series of up to eight test stimuli that were presented unpredictably to their right or left index finger. Attention was cued to one side (finger) at the beginning of each trial, and subjects performed the task at the attended side, ignoring the unattended side.

Traditional studies of spatial attention consider only a single sensory modality at a time (e.g. just vision, or just audition). In daily life, however, our spatial attention often has to be coordinated across several modalities. This is a non-trivial problem, given that each modality initially codes space in entirely different ways. In the last five years, there has been a spate of studies on crossmodal attention. These have demonstrated numerous crossmodal links in spatial attention, such that attending to a particular location in one modality tends to produce corresponding shifts of attention in other modalities. The spatial coordinates of these crossmodal links illustrate that the internal representation of external space depends on extensive crossmodal integration. Recent neuroscience studies are discussed that suggest possible brain mechanisms for the crossmodal links in spatial attention.

It has recently been demonstrated that a cortical network of visuospatial and oculomotor control areas is active for covert shifts of spatial attention (shifts of attention without eye movements) as well as for overt shifts of spatial attention (shifts of attention with saccadic eye movements). Studies examining activity in this visuospatial network during attentional shifts at a single rate have given conflicting reports about how the activity differs for overt and covert shifts. To better understand how the network subserves attentional shifts, we performed a parametric study in which subjects made either overt attentional shifts or covert attentional shifts at three different rates (0.2, 1.0, and 2.0 Hz). At every shift rate, both overt and covert shifts of visuospatial attention induced activations in the precentral sulcus, intraparietal sulcus, and lateral occipital cortex that were of greater amplitude for overt than during covert shifting. As the rate of attentional shifts increased, responses in the visuospatial network increased in both overt and covert conditions but this parametric increase was greater during overt shifts. These results confirm that overt and covert attentional shifts are subserved by the same network of areas. Overt shifts of attention elicit more neural activity than do covert shifts, reflecting additional activity associated with saccade execution. An additional finding concerns the anatomical organization of the visuospatial network. Two distinct activation foci were observed within the precentral sulcus for both overt and covert attentional shifts, corresponding to specific anatomical landmarks. We therefore reappraise the correspondence of these two precentral areas with the frontal eye fields.

Research on sensory perception now often tal setup , and Figure 2 outlines the experimental conditions in which the view given to the participants was considers more than one sense at a time. This varied. Figure 3 shows typical data from two blocks of approach reflects real-world situations, such as trials showing how 2PDT estimates were arrived at. The when a visible object touches us. Indeed, vision results (Figure 4a,b) clearly show that tactile resolution and touch show great interdependence: the sight of was better when visibility of the arm was allowed coma body part can reduce tactile target detection pared to when the arm was in complete darkness. For times [1], visual and tactile attentional systems are those participants who performed binocularly, the mean spatially linked [2], and the texture of surfaces that tactile two-point discrimination threshold was signifiare actively touched with the fingertips is perceived cantly lower (mean ϭ 35 mm) when the arm could be seen using both vision and touch [3]. However, these than when the arm was in complete darkness (mean ϭ 42 previous findings might be mediated by spatial mm; t 8 ϭ 2.1, p ϭ 0.03). This central finding was replicated attention [1, 2] or by improved guidance of in a separate group of participants who performed all four movement [3] via visually enhanced body position conditions monocularly. Visibility of the arm once more sense [4-6]. Here, we investigate the direct effects provided lower thresholds (mean ϭ 27 mm) than when of viewing the body on passive touch. We measured the arm was in darkness (mean ϭ 31 mm; t 9 ϭ 2.9, p ϭ tactile two-point discrimination thresholds [7] on 0.009). This benefit could not be attributed to trivial visithe forearm while manipulating the visibility of the bility of the tactile stimulation, since the arm could not arm but holding gaze direction constant. The be seen during the moment of tactile stimulation. Indeed, spatial resolution of touch was better when the arm a control study revealed that the available visual informawas visible than when it was not. Tactile tion alone (replayed later from a video, without tactile performance was further improved when the view of stimulation) led to only chance performance on judgethe arm was magnified. In contrast, performance ments of the number of tactile stimuli. Moreover, gaze was not improved by viewing a neutral object at the direction was constant across all conditions. Therefore, arm's location, ruling out improved spatial the effects of gaze direction, which are known to affect orienting as a possible account. Controls confirmed tactile performance [8], cannot explain our results. that no information about the tactile stimulation was provided by visibility of the arm. This visual Importantly, visibility of the arm also gave rise to better enhancement of touch may point to online performance than did visibility of the neutral object (mean ϭ reorganization of tactile receptive fields. 30 mm; t 9 ϭ 1.9, p ϭ 0.04; Figure 4d). Also, visibility of

Theories of visual selective attention posit that both spatial location and nonspatial stimulus features (e.g., color) are elementary dimensions on which top-down attentional control mechanisms can selectively influence visual processing. Neuropsychological and neuroimaging studies have demonstrated that regions of superior frontal and parietal cortex are critically involved in the control of visual-spatial attention. This frontoparietal control network has also been found to be activated when attention is oriented to nonspatial stimulus features (e.g., motion). To test the generality of the frontoparietal network in attentional control, we directly compared spatial and nonspatial attention in a cuing paradigm. Event-related fMRI methods permitted the isolation of attentional control activity during orienting to a location or to a nonspatial stimulus feature (color). Portions of the frontoparietal network were commonly activated to the spatial and nonspatial cues. However, direct statistical comparisons of cue-related activity revealed subregions of the frontoparietal network that were significantly more active during spatial than nonspatial orienting when all other stimulus, task, and attentional factors were equated. No regions of the frontal-parietal network were more active for nonspatial cues in comparison to spatial cues. These findings support models suggesting that subregions of the frontal-parietal network are highly specific for controlling spatial selective attention.

Attention is the mechanism with which we select specific aspects of our environment for processing. Psychological experiments have shown that attention can be directed to a spatial location or to a particular object. Electrophysiological studies in trained macaque monkeys have found that attention can strengthen the responses of neurons in cortical area V4. Some of these studies have attributed these effects to spatial attention, whereas others have suggested that feature-directed attention may modulate the neuronal response. Here we report that neuronal correlates for both spatial and feature-directed attention exist in individual neurons in area V4 of behaving rhesus monkeys.

We examined neural activations during decision-making using fMRI paired with the wheel of fortune task, a newly developed two-choice decision-making task with probabilistic monetary gains. In particular, we assessed the impact of high-reward/risk events relative to low-reward/risk events on neural activations during choice selection and during reward anticipation. Seventeen healthy adults completed the study. We found, in line with predictions, that (i) the selection phase predominantly recruited regions involved in visuo-spatial attention (occipito-parietal pathway), conflict (anterior cingulate), manipulation of quantities (parietal cortex), and preparation for action (premotor area), whereas the anticipation phase prominently recruited regions engaged in reward processes (ventral striatum); and (ii) high-reward/risk conditions relative to low-reward/risk conditions were associated with a greater neural response in ventral striatum during selection, though not during anticipation. Following an a priori ROI analysis focused on orbitofrontal cortex, we observed orbitofrontal cortex activation (BA 11 and 47) during selection (particularly to high-risk/reward options), and to a more limited degree, during anticipation. These findings support the notion that (1) distinct, although overlapping, pathways subserve the processes of selection and anticipation in a two-choice task of probabilistic monetary reward; (2) taking a risk and awaiting the consequence of a risky decision seem to affect neural activity differently in selection and anticipation; and thus (3) common structures, including the ventral striatum, are modulated differently by risk/reward during selection and anticipation. Published by Elsevier Ltd.

We report two experiments designed to investigate the potential use of vibrotactile warning signals to present spatial information to car drivers. Participants performed an attention-demanding rapid serial visual presentation (RSVP) monitoring task. Meanwhile, whenever they felt a vibrotactile stimulus presented on either their front or back, they had to check the front and the rearview mirror for the rapid approach of a car, and brake or accelerate accordingly. We investigated whether speeded responses to potential emergency driving situations could be facilitated by the presentation of spatially-predictive (80% valid; Experiment 1) or spatially-nonpredictive (50% valid; Experiment 2) vibrotactile cues. Participants responded significantly more rapidly following both spatially-predictive and spatially-nonpredictive vibrotactile cues from the same rather than the opposite direction as the critical driving events. These results highlight the potential utility of vibrotactile warning signals in automobile interface design for directing a driver’s visual attention to time-critical events or information.

Throughout the history of mathematics, concepts of number and space have been tightly intertwined. We tested the hypothesis that cortical circuits for spatial attention contribute to mental arithmetic. We trained a multivariate classifier to infer the direction of an eye movement, left or right, from the brain activation measured in posterior parietal cortex. Without further training, the classifier then generalized to an arithmetic task. Its left versus right classification could be used to sort out subtraction versus addition trials, whether performed with symbols or with sets of dots. These findings are consistent with the suggestion that mental arithmetic co-opts parietal circuitry associated with spatial coding.

In a study of the neural processes that mediate visual attention in humans, 32-channel recordings of event-related potentials were obtained from 14 normal subjects while they performed a spatial attention task. The generator locations of the early C1, P1, and Nl components of the visual evoked response were estimated by means of topographic maps of voltage and current source density in conjunction with dipole modelling. The topography of the C1 component (ca. 85 ms post-stimulus) was consistent with a generator in striate cortex, and this component was unaffected by attention. In contrast, the P1 and Nl components (ca. 95 and 170ms) exhibited current density foci at scalp sites overlying lateral extrastriate cortex and were larger for attended stimuli than for unattended stimuli. The voltage topographies in the 75–175 ms latency range were modeled with a 5-dipole configuration consisting of a single striate dipole and left-right pairs of dipoles located in lateral extrastriate and inferior occipito-temporal areas. This model was found to account for the voltage topographies produced by both attended and unattended stimuli with low residual variance. These results support the proposal that visual-spatial attention modulates neural activity in extrastriate visual cortex but does not affect the initial evoked response in striate cortex.

The purpose of this study was to identify brain regions underlying internally generated anticipatory biases toward locations where significant events are expected to occur. Subjects fixated centrally and responded to peripheral targets preceded by a spatially valid (predictive), invalid (misleading), or neutral central cue while undergoing fMRI scanning. In some validly cued trials, reaction time was significantly shorter than in trials with neutral cues, indicating that the cue had successfully induced a spatial redistribution of motivational valence, manifested as expectancy. The largest cue benefits led to selectively greater activations within the posterior cingulate and medial prefrontal cortex. These two areas thus appear to establish a neural interface between attention and motivation. An inverse relationship to cue benefit was seen in the parietal cortex, suggesting that spatial expectancy may entail the inhibition of attention-related areas to reduce distractibility by events at irrelevant locations.

The authors report a series of 6 experiments investigating crossmodal links between vision and touch in covert endogenous spatial attention. When participants were informed that visual and tactile targets were more likely on 1 side than the other, speeded discrimination responses (continuous vs. pulsed, Experiments 1 and 2; or up vs. down, Experiment 3) for targets in both moralities were significantly faster on the expected side, even though target morality was entirely unpredictable. When participants expected a target on a particular side in just one morality, corresponding shifts of covert attention also took place in the other morality, as evidenced by faster elevation judgments on that side (Experiment 4). Larger attentional effects were found when directing visual and tactile attention to the same position rather than to different positions (Experiment 5). A final study with crossed hands revealed that these visuotactile links in spatial attention apply to common positions in external space.

This study investigated the cortical mechanisms of visual-spatial attention in a task where subjects discriminated patterned targets in one visual field at a time. Functional magnetic imaging (fMRI) was used to localize attention-related changes in neural activity within specific retinotopic visual areas, while recordings of event-related brain potentials (ERPs) traced the time course of these changes. The earliest ERP components enhanced by attention occurred in the time range 70 -130 ms post-stimulus onset, and their neural generators were estimated to lie in the dorsal and ventral extrastriate visual cortex. The anatomical areas activated by attention corresponded closely to those showing increased neural activity during passive visual stimulation. Enhanced neural activity was also observed in the primary visual cortex (area V1) with fMRI, but ERP recordings indicated that the initial sensory response at 50-90 ms that was localized to V1 was not modulated by attention. Modeling of ERP sources over an extended time range showed that attended stimuli elicited a long-latency (160-260 ms) negativity that was attributed to the dipolar source in area V1. This finding is in line with hypotheses that V1 activity may be modulated by delayed, reentrant feedback from higher visual areas.

Voluntary attention is the top-down selection process that focuses cortical processing resources on the most relevant sensory information. Spatial attention-that is, selection based on stimulus position-alters neuronal responsiveness throughout primate visual cortex. It has been hypothesized that it also changes receptive field profiles by shifting their centers toward attended locations and by shrinking them around attended stimuli. Here we examined, at high resolution, receptive fields in cortical area MT of rhesus macaque monkeys when their attention was directed to different locations within and outside these receptive fields. We found a shift of receptive fields, even far from the current location of attention, accompanied by a small amount of shrinkage. Thus, already in early extrastriate cortex, receptive fields are not static entities but are highly modifiable, enabling the dynamic allocation of processing resources to attended locations and supporting enhanced perception within the focus of attention by effectively increasing the local cortical magnification.

High-field (3 Tesla) functional magnetic resonance imaging (MRI) was used to investigate the cortical circuitry subserving pursuit tracking in humans and compare it to that for saccadic eye movements. Pursuit performance, relative to visual fixation, elicited activation in three areas known to contribute to eye movements in humans and in nonhuman primates: the frontal eye field, supplementary eye field, and intraparietal sulcus. It also activated three medial regions not previously identified in human neuroimaging studies of pursuit: the precuneus and the anterior and posterior cingulate cortices. All six areas were also activated during saccades. The spatial extent of activation was similar for saccades and pursuit in all but two regions: spatial extent was greater for saccades in the superior branch of the frontal eye field and greater for pursuit in posterior cingulate cortex. This set of activations for smooth pursuit parallels the network of oculomotor areas characterized in nonhuman primates and complements recent studies showing that common cortical networks subserve oculomotor functions and spatial attention in humans.

According to the activation-orienting hypothesis the distribution of attention in space is biased in the direction contralateral to the more activated hemisphere. The present investigation tested this proposal and evaluated the nature of hemispheric differences in orienting control. Activation imbalance was produced by a unilateral visual stimulus. The distribution of attention was measured using a modified line bisection task in which subjects judged the location of an intersect on a tachistoscopically presented horizontal line. The first three experiments suggest that (a) attention is biased in the direction contralateral to the stimulated
hemisphere, and (b) the biases do not depend on the task relevance or hemispatial position of the stimulus producing the activation imbalance. The final three experiments suggest that when orienting conflict is introduced the rightward bias becomes more robust than the leftward bias. The findings are consistent with the activation-orienting hypothesis. Each hemisphere generates a contralateralattentional bias and the rightward bias of the left hemisphere is stronger. The relevance of these findings to understanding unilateral neglect resulting from
parietal damage is discussed.

This study explored whether hand location affected spatial attention. The authors used a visual covertorienting paradigm to examine whether spatial attention mechanisms-location prioritization and shifting attention-were supported by bimodal, hand-centered representations of space. Placing 1 hand next to a target location, participants detected visual targets following highly predictive visual cues. There was no a priori reason for the hand to influence task performance unless hand presence influenced attention. Results showed that target detection near the hand was facilitated relative to detection away from the hand, regardless of cue validity. Similar facilitation was found with only proprioceptive or visual hand location information but not with arbitrary visual anchors or distant targets. Hand presence affected attentional prioritization of space, not the shifting of attention.

Recent imaging and clinical studies have challenged the concept that the functional role of the cerebellum is exclusively in the motor domain. We present evidence of slowed covert orienting of visuospatial attention in patients with developmental cerebellar abnormality (patients ...

The goal of this experiment was to determine whether the allocation of attention necessarily requires saccade preparation. To dissociate the focus of attention from the endpoint of a saccade, macaque monkeys were trained to perform visual search for a uniquely colored rectangle and shift gaze either toward or opposite this color singleton according to its orientation. A vertical singleton cued a prosaccade, a horizontal singleton, an antisaccade. Saccade preparation was probed by measuring the direction of saccades evoked by intracortical microstimulation of the frontal eye fields at variable times after presentation of the search array. Eye movements evoked on prosaccade trials deviated progressively toward the singleton that was also the endpoint of the correct eye movement. However, eye movements evoked on antisaccade trials never deviated toward the singleton but only progressively toward the location opposite the singleton. This occurred even though previous work showed that on antisaccade trials most neurons in frontal eye fields initially select the singleton while attention is allocated to distinguish its shape. Thus, sensorimotor structures can covertly orient attention without preparing a saccade.

These University of California, San Diego findings suggested that the visual pathways in V1 may La Jolla, California 92093 only subserve passive sensory transmission and are 4 Center for Cognitive Neuroscience and not subject to cognitive control under a wide range of Department of Psychological and Brain Sciences attentional tasks. Duke University During the past few years, however, several studies Durham, North Carolina 27708 in monkeys have affirmed Motter's (1993) earlier finding that neural activity in V1 may be modified by attention under certain circumstances, particularly in discrimina-Summary tion tasks when several competing stimuli are present in the visual field (Roelfsema et al., 1998; Vidyasagar,

Visual attention is a mechanism by which observers select relevant or important information from the current visual array. Previous investigations have focused primarily on the ability to select a region of space for further visual analysis. These studies have revealed a distributed frontoparietal circuit that is responsible for the control of spatial attention. However, vision must ultimately represent objects and in real scenes objects often overlap spatially; thus attention must be capable of selecting objects and their properties nonspatially.

Vol.3.No.1.,Haptics-e, 11-June-2003 http://www.haptics-e.org ... Hong Z. Tan,† Robert Gray,‡ J. Jay Young,† and Ryan Traylor† ... † Haptic Interface Research Laboratory Purdue University 1285 Electrical Engineering Building West Lafayette, IN 47907 {hongtan, ...

Laser evoked potentials (LEPs) are brain responses to activation of skin nociceptors by laser heat stimuli. LEPs consist of three components: N1, N2, and P2. Previous reports have suggested that in contrast to earlier activities (N1), LEPs responses after 230-250 ms (N2-P2) are modulated by attention to painful laser stimuli. However, the experimental paradigms used were not designed to specify the attentional processes involved in these LEP modulations. We investigated the effects of selective spatial attention and oddball tasks on LEPs. CO 2 laser stimuli of two different intensities were delivered on the dorsum of both hands of ten subjects. One intensity was frequently presented, and the other rarely. Subjects were asked to pay attention to stimuli delivered on one hand and to count rare stimuli, while ignoring stimuli on the other hand. Frequent and rare attended stimuli evoked enhanced N160 (N1) and N230 (N2) components in comparison to LEPs from unattended stimuli. Both components showed scalp distribution contralateral to the stimulus location. The vertex P400 (P2) was unaffected by spatial attention and stimulus location, but its amplitude increased after rare stimuli, whether attended or unattended. An additional parietal P600 component was induced by the attended rare stimuli. It is suggested that several attentional processes can modify nociceptive processing in the brain at different stages. LEP activities in the time-range of N1 and N2 (120-270 ms) showed evidence of processes modulated by the direction of spatial attention. Conversely, processes underlying P2 (400 ms) were not affected by spatial attention, but by the probability of the stimulus. This probability effect was not due to P3b-related processes that were observed at a later latency (600 ms). Indeed, P600 could be seen as a P3b evoked by conscious detection of rare targets.