Retinotopic organization of early visual spatial attention effects as revealed by PET and ERPs

Cerebral blood flow PET scans and high-density event-related potentials (ERPs) were recorded (separate sessions) while subjects viewed rapidly-presented, lower-visual-field, bilateral stimuli. Active attention to a designated side of the stimuli (relative to passive-viewing conditions) resulted in an enhanced ERP positivity (P1 effect) from 80–150 msec over occipital scalp areas contralateral to the direction of attention. In PET scans, active attention vs. passive showed strong activation in the contralateral dorsal occipital cortex, thus following the retinotopic organization of the early extrastriate visual sensory areas, with some weaker activation in the contralateral fusiform. Dipole modeling seeded by the dorsal occipital PET foci yielded an excellent fit for the P1 attention effect. In contrast, dipoles constrained to the fusiform foci fit the P1 effect poorly, and, when the location constraints were released, moved upward to the dorsal occipital locations during iterative dipole fitting. These results argue that the early ERP P1 attention effects for lower-visual-field stimuli arise mainly from these dorsal occipital areas and thus also follow the retinotopic organization of the visual sensory input pathways. These combined PET/ERP data therefore provide strong evidence that sustained visual spatial attention results in a preset, top-down biasing of the early sensory input channels in a retinotopically organized way. Hum. Brain Mapping 5:280–286, 1997. © 1997 Wiley-Liss, Inc.     

Combining steady-state visual evoked potentials and f MRI to localize brain activity during selective attention

Brain activity was studied with fMRI and steady-state visual evoked potentials (SSVEPs) in separate sessions as subjects attended to letter sequences in either the right or left visual field. The two letter sequences were superimposed on small square backgrounds that flickered at 8.6 and 12.0 Hz, respectively, and elicited SSVEPs at the flicker frequencies. The amplitude of the frequency-coded SSVEP elicited by either of the flickering backgrounds was enlarged when attention was focused upon the letter sequence at the same location. Source analysis of the SSVEP waveforms localized the attentional modulation to ventral and lateral extrastriate visual cortex, which corresponded to zones of activation observed with fMRI. Hum. Brain Mapping 5:287–292, 1997. © 1997 Wiley-Liss, Inc.     

Hemifield effects of spatial attention in early human visual cortex

Early visual areas (V1, V2, V3/VP, V4v) contain representations of the contralateral hemifield within each hemisphere. Little is known about the role of the visual hemifields along the visuo-spatial attention processing hierarchy. It is hypothesized that attentional information processing is more efficient across the hemifields (known as bilateral field advantage) and that the integration of information is greater within one hemifield as compared with across the hemifields. Using functional magnetic resonance imaging we examined the effect of distance and hemifield on parallel attentional processing in the early visual areas (V1-V4v) at individually mapped retinotopic locations aligned adjacently or separately within or across the hemifields. We found that the bilateral field advantage in parallel attentional processing over separated attended locations can be assigned, at least partly, to differences in distractor position integration in early visual areas. These results provide evidence for a greater integration of locations between two attended locations within one hemifield than across both hemifields. This nicely correlates with behavioral findings of a bilateral field advantage in parallel attentional processing (when distractors in between cannot be excluded) and a unilateral field advantage if attention has to be shifted across separated locations (when locations in between were integrated).     

Electrophysiological measures and dual-task performance in Tourette syndrome indicate deficient divided attention mechanisms

Tourette syndrome has been associated with impairments of attentional functions such as distractability, even in subjects without co-morbid attention deficit hyperactivity disorder. Based on the results of earlier research we hypothesized that Tourette syndrome patients might employ altered control mechanisms of attentional processes and have concurrent difficulties in allocating their attentional resources among competing tasks. Event-related brain potentials (ERPs) were recorded in a group of Tourette syndrome patients and in a matched control group during a dual task experiment. This experiment required the simultaneous detection of visual and auditory target stimuli which were manipulated to yield two different difficulty levels each of which were varied orthogonally. The behavioural parameters confirmed the intended performance differences between difficult-to-detect targets and easy-to-detect targets. This was paralleled by lower amplitudes and longer latencies of the corresponding P3b-ERP subcomponents. Although Tourette syndrome patients were unimpaired in overall performance they showed an increased interference of visual task demands with auditory target perception. In parallel they also exhibited a reduced amplitude of the P3b component to auditory targets. The findings show that Tourette syndrome patients are not generally impaired in their dual task performance. The allocation of attentional resources to competing tasks however, is altered. We speculate that this may be related to deficient inhibitory functions.     

A shift of visual spatial attention is selectively associated with human EEG alpha activity

Event-related potentials and ongoing oscillatory electroencephalogram (EEG) activity were measured while subjects performed a cued visual spatial attention task. They were instructed to shift their attention to either the left or right visual hemifield according to a cue, which could be valid or invalid. Thereafter, a peripheral target had to be evaluated. At posterior parietal brain areas early components of the event-related potential (P1 and N1) were higher when the cue had been valid compared with invalid. An anticipatory attention effect was found in EEG alpha magnitude at parieto-occipital electrode sites. Starting 200 ms before target onset alpha amplitudes were significantly stronger suppressed at sites contralateral to the attended visual hemifield than ipsilateral to it. In addition, phase coupling between prefrontal and posterior parietal electrode sites was calculated. It was found that prefrontal cortex shows stronger phase coupling with posterior sites that are contralateral to the attended hemifield than ipsilateral sites. The results suggest that a shift of attention selectively modulates excitability of the contralateral posterior parietal cortex and that this posterior modulation of alpha activity is controlled by prefrontal regions.     

Attentional modulation of perceptual grouping in human visual cortex: ERP studies

We sought to identify the neural substrates underlying perceptual grouping, and examined whether the grouping-related neural activities are modulated by task relevance and attention by recording high-density event-related potentials (ERPs). Participants were presented with stimulus arrays, in which local elements were either evenly distributed or grouped into rows or columns by proximity or similarity, and had to discriminate orientations of the perceptual groups, or to identify the colors of dots around stimulus displays or of the fixation cross. We found that proximity grouping was indexed by a positive activity over the medial occipital cortex with a peak latency of about 100 ms after stimulus onset (Pd100), whereas grouping by similarity of shape was reflected in a negative activity with longer latency over the occipito-temporal areas. Dipole modeling based on a realistic boundary element head model localized the Pd100 to the right calcarine cortex. Moreover, we showed that the grouping-related activities were weakened when stimulus arrays were of low task relevance and fell outside an attended area of field. The results suggest that human calcarine cortex is engaged in early grouping operations defined by proximity, reinforcing the previous fMRI findings. Moreover, our ERP results indicate that the neural bases underlying perceptual grouping in human visual cortex can be modulated by task relevance and attention as early as 100 ms after sensory stimulation. Hum Brain Mapp, 2005. (c) 2005 Wiley-Liss, Inc.     

Timing the fearful brain: unspecific hypervigilance and spatial attention in early visual perception

Numerous studies suggest that anxious individuals are more hypervigilant to threat in their environment than nonanxious individuals. In the present event-related potential (ERP) study, we sought to investigate the extent to which afferent cortical processes, as indexed by the earliest visual component C1, are biased in observers high in fear of specific objects. In a visual search paradigm, ERPs were measured while spider-fearful participants and controls searched for discrepant objects (e.g. spiders, butterflies, flowers) in visual arrays. Results showed enhanced C1 amplitudes in response to spatially directed target stimuli in spider-fearful participants only. Furthermore, enhanced C1 amplitudes were observed in response to all discrepant targets and distractors in spider-fearful compared with non-anxious participants, irrespective of fearful and non-fearful target contents. This pattern of results is in line with theoretical notions of heightened sensory sensitivity (hypervigilance) to external stimuli in high-fearful individuals. Specifically, the findings suggest that fear facilitates afferent cortical processing in the human visual cortex in a non-specific and temporally sustained fashion, when observers search for potential threat cues.     

Viewing the body modulates neural mechanisms underlying sustained spatial attention in touch

Cross-modal links between vision and touch have been extensively shown with a variety of paradigms. The present event-related potential (ERP) study aimed to clarify whether neural mechanisms underlying sustained tactile-spatial attention may be modulated by visual input, and the sight of the stimulated body part (i.e. hands) in particular. Participants covertly attended to one of their hands throughout a block to detect infrequent tactile target stimuli at that hand while ignoring tactile targets at the unattended hand, and all tactile non-targets. In different blocks, participants performed this task under three viewing conditions: full vision; hands covered from view; and blindfolded. When the participants' hands were visible attention was found to modulate somatosensory ERPs at early latencies (i.e. in the time range of the somatosensory P100 and the N140 components), as well as at later time intervals, from 200 ms after stimulus onset. By contrast, when participants were blindfolded and, crucially, even when only their hands were not visible, attentional modulations were found to arise only at later intervals (i.e. from 200 ms post-stimulus), while earlier somatosensory components were not affected by spatial attention. The behavioural results tallied with these electrophysiological findings, showing faster response times to tactile targets under the full vision condition compared with conditions when participants' hands were covered, and when participants were blindfolded. The results from this study provide the first evidence of the profound impact of vision on mechanisms underlying sustained tactile-spatial attention, which is enhanced by the sight of the body parts (i.e. hands).     

Developmental dyslexics show altered allocation of attention in visual classification tasks

OBJECTIVES: Event-related brain potentials (ERPs) were recorded to investigate allocation of attention in adult developmental dyslexics. SUBJECTS AND METHODS: Twelve adult developmental dyslexics and 12 matched normal controls performed three visual choice reaction tasks. In the passive oddball condition, subjects watched two different simple visual stimuli presented with 87.5 and 12.5% probability. In the active oddball condition, participants responded to the low-probability target stimulus. In the active 50/50-condition, both stimuli were presented with 50% probability and a response was required to the target stimulus only. RESULTS: No group differences emerged for performance, P300 latency or laterality and for N200 amplitude, latency or laterality. An enhancement of P300 amplitude with a frontal distribution was found for NoGo (standard)-stimuli in both active conditions for the dyslexic sample. CONCLUSION: Results are discussed in the context of deviances in allocation of attentional resources in dyslexic readers.     

Brain dynamics of visual attention during anticipation and encoding of threat- and safe-cues in spider-phobic individuals

This study systematically investigated the sensitivity of the phobic attention system by measuring event-related potentials (ERPs) in spider-phobic and non-phobic volunteers in a context where spider and neutral pictures were presented (phobic threat condition) and in contexts where no phobic but unpleasant and neutral or only neutral pictures were displayed (phobia-irrelevant conditions). In a between-group study, participants were assigned to phobia-irrelevant conditions either before or after the exposure to spider pictures (pre-exposure vs post-exposure participants). Additionally, each picture was preceded by a fixation cross presented in one of three different colors that were informative about the category of an upcoming picture. In the phobic threat condition, spider-phobic participants showed a larger P1 than controls for all pictures and signal cues. Moreover, individuals with spider phobia who were sensitized by the exposure to phobic stimuli (i.e. post-exposure participants) responded with an increased P1 also in phobia-irrelevant conditions. In contrast, no group differences between spider-phobic and non-phobic individuals were observed in the P1-amplitudes during viewing of phobia-irrelevant stimuli in the pre-exposure group. In addition, cues signaling neutral pictures elicited decreased stimulus-preceding negativity (SPN) compared with cues signaling emotional pictures. Moreover, emotional pictures and cues signaling emotional pictures evoked larger early posterior negativity (EPN) and late positive potential (LPP) than neutral stimuli. Spider phobics showed greater selective attention effects than controls for phobia-relevant pictures (increased EPN and LPP) and cues (increased LPP and SPN). Increased sensitization of the attention system observed in spider-phobic individuals might facilitate fear conditioning and promote generalization of fear playing an important role in the maintenance of anxiety disorders.     

The wandering mind of men: ERP evidence for gender differences in attention bias towards attractive opposite sex faces

To examine the time course and automaticity of our attention bias towards attractive opposite sex faces, event-related potentials (ERPs) were recorded from 20 males and 20 females while they carried out a covert orienting task. Faces that were high, low or average in attractiveness, were presented in focus of attention, but were unrelated to task goals. Across the entire sample larger P2 amplitudes were found in response to both attractive and unattractive opposite sex faces, presumably reflecting early implicit selective attention to distinctive faces. In male but not female participants this was followed by an increased late slow wave for the attractive faces, signifying heightened processing linked to motivated attention. This latter finding is consistent with sexual strategy theory, which suggests that men and women have evolved to pursue different mating strategies with men being more attentive to cues such as facial beauty. In general, our ERP results suggest that, in addition to threat-related stimuli, other evolutionary-relevant information is also prioritized by our attention systems.     

Visual mismatch negativity highlights abnormal pre-attentive visual processing in mild cognitive impairment and Alzheimer's disease

Alzheimer's disease (AD) and mild cognitive impairment (MCI) are traditionally characterised in relation to abnormalities in higher-level brain function, particularly memory. However, in the study presented here we report significant abnormality in pre-attentive visual processing in both MCI and AD compared to healthy ageing. The functional integrity of the automatic change detection system was determined by measuring the visual mismatch negativity (vMMN) component of the N2 visual event-related potential (ERP) in 8 individuals with amnestic MCI, 10 individuals with AD and 10 healthy older adult controls. Compared to healthy ageing, both the MCI and AD groups displayed a significant abnormally increased vMMN over the 140–250 ms measurement epoch although such effects were not apparent over the later 250–400 ms epoch. These novel findings support the view that deficits in brain function in both AD and MCI are not exclusively high-level and indeed that abnormal output from ‘low level’ or pre-attentive processes (as evinced by the abnormal vMMN) is likely to contribute to the high-level processing deficits so characteristic of these clinical disorders.     

Selective auditory attention effects in tonotopically organized cortical areas: A topographic ERP study

This study reinvestigates some aspects of the neurophysiological mechanisms of auditory selective attention, by testing the hypothesis that auditory attention can exert a selective control over the sensory processing of acoustic stimuli in tonotopic auditory cortex. Event-related potentials (ERPs) were recorded from human subjects while they listened selectively to a tone sequence in one ear and ignored a concurrent sequence of a different frequency in the opposite ear. The tone frequencies were 500, 1,000, 2,000, or 4,000 Hz in separate sequences, and were delivered at a constant rate of one every 800 ms. The effects of attention were analyzed in the difference waves obtained by subtracting ERPs to ignore tones from those of the same tones when attended. The earliest effect of attention (70–80 ms post-stimulus) was found to present the same spatio-temporal organization as the obligatory, sensory-evoked N1 wave, with similar tonotopic changes in scalp distribution with the tone frequencies. At longer latencies, two other attentional effects were observed, of probable endogenous origin: one around 175 ms post-stimulus, possibly originating from non-tonotropic auditory cortex, and the latest one (after 300 ms) from non-specific areas. The results support the hypothesis of a genuine sensory gating mechanism for auditory attention, taking place in tonotopic auditory cortex at an early stage of sensory processing, even in low attentional load conditions. © 1995 Wiley-Liss, Inc.     

Topography of attention in the primary visual cortex

Previous research suggests that feedback circuits mediate the effect of attention to the primary visual cortex (V1). This inference is mainly based on temporal information of the responses, where late modulation is associated with feedback signals. However, temporal data alone are inconclusive because the anatomical hierarchy between cortical areas differs significantly from the temporal sequence of activation. In the current work, we relied on recent physiological and computational models of V1 network architecture, which have shown that the thalamic feedforward, local horizontal and feedback contribution are reflected in the spatial spread of responses. We used multifocal functional localizer and quantitative analysis in functional magnetic resonance imaging to determine the spatial scales of attention and sensory responses. Representations of 60 visual field regions in V1 were functionally localized and four of these regions were targets in a subsequent attention experiment, where human volunteers fixated centrally and performed a visual discrimination task at the attended location. Attention enhanced the peak amplitudes significantly more in the lower than in the upper visual field. This enhancement by attention spread with a 2.4 times larger radius (approximately 10 mm, assuming an average magnification factor) compared with the unattended response. The corresponding target region of interest was on average 20% stronger than that caused by the afferent sensory stimulation alone. This modulation could not be attributed to eye movements. Given the contemporary view of primate V1 connections, the activation spread along the cortex provides further evidence that the signal enhancement by spatial attention is dependent on feedback circuits.     

Age-related changes in the attentional control of visual cortex: a selective problem in the left visual hemifield

To what extent does our visual-spatial attention change with age? In this regard, it has been previously reported that relative to young controls, seniors show delays in attention-related sensory facilitation. Given this finding, our study was designed to examine two key questions regarding age-related changes in the effect of spatial attention on sensory-evoked responses in visual cortex--are there visual field differences in the age-related impairments in sensory processing, and do these impairments co-occur with changes in the executive control signals associated with visual spatial orienting? Therefore, our study examined both attentional control and attentional facilitation in seniors (aged 66-74 years) and young adults (aged 18-25 years) using a canonical spatial orienting task. Participants responded to attended and unattended peripheral targets while we recorded event-related potentials (ERPs) to both targets and attention-directing spatial cues. We found that not only were sensory-evoked responses delayed in seniors specifically for unattended events in the left visual field as measured via latency shifts in the lateral occipital P1 elicited by visual targets, but seniors also showed amplitude reductions in the anterior directing attentional negativity (ADAN) component elicited by cues directing attention to the left visual field. At the same time, seniors also had significantly higher error rates for targets presented in the left vs. right visual field. Taken together, our data thus converge on the conclusion that age-related changes in visual spatial attention involve both sensory-level and executive attentional control processes, and that these effects appear to be strongly associated with the left visual field.     

Motion direction tuning in human visual cortex

A number of electrophysiological studies have been conducted in recent years in order to clarify the dynamics of visual motion processing in the human brain. Using a variety of event-related potential (ERP) measures, several parameters such as onset, offset, contrast and velocity have been investigated, while a critical aspect of visual motion, that of direction, has received less attention. Here we used multichannel electroencephalography and distributed source localization to study brain activity for different directions of visual motion using random dot stimuli. Our data reveal differential extrastriate activation at 164–226 ms after motion onset that coded for motion direction with different ERP maps and underlying electrical generators for each tested direction. This activation was paralleled initially (164–186 ms) by a distinct extrastriate activation encoding whether the motion stimulus consisted of directed motion stimuli (as above) or contained undirected incoherent motion (control stimulus). Application of a linear inverse solution localized the brain activity for each tested motion direction to distinct brain regions within the same larger network of extrastriate brain regions. These regions included bilateral temporo-occipital and bilateral parieto-occipital cortex. The present data in healthy subjects are compatible with extrastriate activity that is tuned to different directions of visual motion. This extends previous clinical data and suggests the presence of distributed macroscopic motion direction tuning in primate extrastriate cortex that may complement the classical microscopic motion tuning at the columnar level.     

Cross-modal selective attention to visual and auditory stimuli modulates endogenous ERP components

In two experiments event-related potentials (ERPs) to visual and auditory stimuli were measured in 12 healthy subjects. A cross-modal and delayed response paradigm was used that allows ERPs to be obtained separately to attended and unattended stimuli under conditions in which unattended stimuli are less likely to be covertly or randomly attended. The results showed: (1) N1 enhancement with attention for standard stimuli in auditory and visual modalities and for deviant stimuli in the visual modality; (2) The onset time and scalp distribution of both the N1 for attend condition and Nd1 were similar regardless of standard or deviant stimuli in the auditory and visual modality; the onset time of Nd1 elicited by auditory and visual deviant stimuli was earlier than that of the unattended N1, and their scalp distributions were different; and (3) The Nd1 components elicited by auditory and visual deviant stimuli were distributed over the respective primary sensory areas, but Nd1 components evoked by auditory and visual standard stimuli were distributed over the frontal scalp. These results suggest that the attended N1 enhancement is primarily caused by a component with endogenous origins and that the early attention effect occurs before the exogenous components. The results support the view that the cross-modal attention to deviant stimuli modulates modality-specific processing in the brain, whereas attention to standard stimuli affects modality-nonspecific or supramodal brain systems.     

Attentional modulation of perceptual grouping in human visual cortex: functional MRI studies

When presented with a complex visual scene, our visual system has to organize the discrete entities present into useful perceptual units. The current work investigated the neural substrates of perceptual grouping defined by Gestalt laws of proximity and similarity of shape, and whether the neural substrates underlying perceptual grouping are modulated by task relevance and spatial attention. In visual discrimination tasks, subjects identified the orientations of perceptual groups formed by proximity or similarity of local elements or alternatively identified colors of either dots around the grouped stimuli or the fixation cross. Using functional magnetic resonance imaging (fMRI), we identified that the calcarine cortex was involved in proximity grouping but not in the grouping process defined by similarity of shape. Moreover, we showed evidence that the neural correlates of proximity grouping in the calcarine cortex were weakened when the elements were of low task relevance and fell outside an attended area of field. The findings reveal the neural basis for basic grouping operations, as well as illustrating how attention and proximity grouping interact in human visual cortex.     

Changes of effective connectivity between the lateral and medial parts of the prefrontal cortex during a visual task

Structural equation modelling was used to study the change of connectivity during a visual task with continuous variation of the attention load. The model was based on areas defined by the haemodynamic responses described elsewhere [Mazoyer, P., Wicker, B. & Fonlupt, P. (2002) A neural network elicited by parametric manipulation of the attention load. Neuroreport, 13, 2331-2334], including occipitotemporal, parietal, temporal and prefrontal (lateral and medial areas) cortices. We have studied stationary- (which does not depend on the attentional load) and attention-related coupling between areas. This allowed the segregation of two subsystems. The first could reflect a system performing the integration step of the visual signal and the second a system participating in response selection. The major finding is the mutual negative influence between the lateral and medial parts of the prefrontal cortex. This negative influence between these two brain regions increased with the attention load. This is interpreted as a modification of the balance between integration and decision processes that are needed for the task to be efficiently completed.     

Executive control of spatial attention shifts in the auditory compared to the visual modality

Voluntary orienting of visual spatial attention has been shown to be associated with activation in a distributed network of frontal and parietal brain areas. Neuropsychological data suggest that at least some of these areas should be sensitive to the direction in which attention is shifted. The aim of this study was to use rapid event-related functional magnetic resonance imaging to investigate whether spatial attention in the auditory modality is subserved by the same or different brain areas as in the visual modality, and whether the auditory and visual attention networks show any degree of hemispheric lateralisation or sensitivity to the direction of attention shifts. The results suggest that auditory and visual spatial attention shifts are controlled by a supramodal network of frontal, parietal and temporal areas. Areas activated included the precuneus and superior parietal cortex, the inferior parietal cortex and temporo-parietal junction, as well as the premotor and supplementary motor areas and dorsolateral prefrontal cortex (DLPFC). In the auditory task, some of these areas, in particular the precuneus as well as the inferior parietal cortex and temporo-parietal junction, showed 'relative' asymmetry, in that they responded more strongly to attention shifts towards the contralateral than the ipsilateral hemispace. Some areas, such as the right superior parietal cortex and left DLPFC, showed 'absolute' asymmetry, in that they responded more strongly in one than in the other hemisphere.