Spatiotemporal dynamics of feature-based attention spread: evidence from combined electroencephalographic and magnetoencephalographic recordings

Attentional selection on the basis of nonspatial stimulus features induces a sensory gain enhancement by increasing the firing-rate of individual neurons tuned to the attended feature, while responses of neurons tuned to opposite feature-values are suppressed. Here we recorded event-related potentials (ERPs) and magnetic fields (ERMFs) in human observers to investigate the underlying neural correlates of feature-based attention at the population level. During the task subjects attended to a moving transparent surface presented in the left visual field, while task-irrelevant probe stimuli executing brief movements into varying directions were presented in the opposite visual field. ERP and ERMF amplitudes elicited by the unattended task-irrelevant probes were modulated as a function of the similarity between their movement direction and the task-relevant movement direction in the attended visual field. These activity modulations reflecting globally enhanced processing of the attended feature were observed to start not before 200 ms poststimulus and were localized to the motion-sensitive area hMT. The current results indicate that feature-based attention operates in a global manner but needs time to spread and provide strong support for the feature-similarity gain model.     

Time course of cross-hemispheric spatial updating in the human parietal cortex

In human parietal cortex, the retinal location of a just seen visual stimulus is updated from one hemisphere to the other, when a horizontal eye movement brings the representation of the stimulus into the opposite visual hemifield. The present study aimed to elucidate the time course of this process. Twelve subjects performed an updating task, in which a filled circle was shown before a horizontal saccade, requiring updating of stimulus location, and a control task without visual stimulation before the saccade. Electroencephalogram (EEG) and electrooculogram (EOG) were recorded while subjects performed the tasks and LORETA source analysis was performed on event-related potential (ERP) components. ERP amplitudes were more positive in the updating condition in comparison to the control condition in two latency windows. An early positive wave starting at about 50 ms after saccade offset and originating in the posterior parietal cortex contralateral to saccade direction probably reflects the integration of saccade-related and visual information and thus the updating process. A shift of the representation of the to-be-updated stimulus to the opposite hemisphere is reflected in a later component starting approximately 400 ms after saccade offset, which is related to memory and originates in the PPC ipsilateral to saccade direction and thus contralateral to the spatial location of the updated visual stimulus.     

Attention to motion enhances processing of both visual and auditory stimuli: an event-related potential study

The present event-related potential (ERP) study investigated whether attending to a particular direction of motion similarly enhances the processing of auditory and visual stimuli. ERPs were recorded while participants perceived horizontally moving visual and auditory stimuli. Attention was manipulated by asking participants to detect an infrequent target stimulus that was of a specified modality (either visual or auditory) and that moved in a specified direction (either leftward or rightward). Stimuli moving in the attended direction elicited ERPs that were more negative than ERPs to stimuli moving in the unattended direction. This difference started around 140 ms post stimulus onset for visual and around 120 ms for auditory stimuli. The auditory effect had a frontal scalp topography, whereas the visual effect was distributed parieto-occipitally. Later parts of the difference waves were maximal at centro-parietal electrodes for both modalities. Crossmodal effects of attention to motion from one modality to the other could not be detected. The results are discussed with regard to hierarchical models of attention.     

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.     

Spatiotemporal mapping of sex differences during attentional processing

Functional neuroimaging studies have increasingly aimed at approximating neural substrates of human cognitive sex differences elicited by visuospatial challenge. It has been suggested that females and males use different behaviorally relevant neurocognitive strategies. In females, greater right prefrontal cortex activation has been found in several studies. The spatiotemporal dynamics of neural events associated with these sex differences is still unclear. We studied 22 female and 22 male participants matched for age, education, and nicotine with 29‐channel‐electroencephalogram recorded under a visual selective attention paradigm, the Attention Network Test. Visual event‐related potentials (ERP) were topographically analyzed and neuroelectric sources were estimated. In absence of behavioral differences, ERP analysis revealed a novel frontal‐occipital second peak of visual N100 that was significantly increased in females relative to males. Further, in females exclusively, a corresponding central ERP component at around 220 ms was found; here, a strong correlation between stimulus salience and sex difference of the central ERP component amplitude was observed. Subsequent source analysis revealed increased cortical current densities in right rostral prefrontal (BA 10) and occipital cortex (BA 19) in female subjects. This is the first study to report on a tripartite association between sex differences in ERPs, visual stimulus salience, and right prefrontal cortex activation during attentional processing. Hum Brain Mapp 2009. © 2009 Wiley‐Liss, Inc.     

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.     

Brain mechanisms of involuntary visuospatial attention: an event-related potential study

The brain mechanisms mediating visuospatial attention were investigated by recording event-related potentials (ERPs) during a line-orientation discrimination task. Nonpredictive peripheral cues were used to direct participant's attention involuntarily to a spatial location. The earliest attentional modulation was observed in the P1 component (peak latency about 130 ms), with the valid trials eliciting larger P1 than invalid trials. Moreover, the attentional modulations on both the amplitude and latency of the P1 and N1 components had a different pattern as compared to previous studies with voluntary attention tasks. In contrast, the earliest visual ERP component, C1 (peak latency about 80 ms), was not modulated by attention. Low-resolution brain electromagnetic tomography (LORETA) showed that the earliest attentional modulation occurred in extrastriate cortex (middle occipital gyrus, BA 19) but not in the primary visual cortex. Later attention-related reactivations in the primary visual cortex were found at about 110 ms after stimulus onset. The results suggest that involuntary as well as voluntary attention modulates visual processing at the level of extrastriate cortex; however, at least some different processes are involved by involuntary attention compared to voluntary attention. In addition, the possible feedback from higher visual cortex to the primary visual cortex is faster and occurs earlier in involuntary relative to voluntary attention task.     

Attentional modulation of early-stage visual processing in schizophrenia

This study shows that paying attention to the color of a visual stimulus is manifested by an early endogenous scalp-positive event-related brain potential (ERP) component, referred to as "selection positivity", that emerges within the first 100 ms after stimulus onset in healthy observers. In contrast, recently ill and chronically ill schizophrenia patients as well as patients at high risk for schizophrenia all failed to show this early ERP component while attending to color. These results suggest that a relatively early stage of visual-selective processing in posterior extrastriate cortex is disrupted in schizophrenia.     

Attending to visual or auditory motion affects perception within and across modalities: an event-related potential study

The present event-related potential (ERP) study examined the role of dynamic features in multisensory binding. It was tested whether endogenous attention to the direction of motion affects processing of visual and auditory stimuli within and across modalities. Human participants perceived horizontally moving dot patterns and sounds that were presented either continuously (standards) or briefly interrupted (infrequent deviants). Their task was to detect deviants moving in a particular direction within a primary modality, but to detect all deviants irrespective of their motion direction within the secondary modality. Attending to the direction of visual motion resulted in a broad selection negativity (SN) starting at about 200 ms post-stimulus onset, and attending to the direction of auditory motion resulted in a positive difference wave at 150 ms that was followed by a broad negativity starting at about 200 ms (unimodal effects). Moreover, dot patterns moving in a direction that was attended within audition were detected faster and more accurately than oppositely moving stimuli and elicited a cross-modal SN wave. Corresponding cross-modal behavioural and ERP results were obtained for sounds moving in a direction that was attended within vision. Unimodal and cross-modal ERP attention effects partially differed in their scalp topography. The present study shows that dynamic features (direction of motion) may be used to link input across modalities and demonstrates for the first time that these multisensory interactions take place as early as about 200 ms after stimulus onset.     

Spatial attention triggered by eye gaze enhances and speeds up visual processing in upper and lower visual fields beyond early striate visual processing.

OBJECTIVE: The detection of a lateralized visual target is faster when preceded by a face gazing to the location of this stimulus. Here we aimed to clarify the time-course of the visual processing modulated by these reflexive shifts of attention. METHODS: ERPs were measured on 16 subjects performing a speeded location task on a circular checkerboard. The checkerboard target appeared either on the left or right of the upper or lower visual field, and was preceded by a central face orienting its gaze obliquely to one of the four possible corner locations for the target to appear. RESULTS: Congruently cued targets were located faster than incongruently cued targets and were associated with larger and earlier occipital P1 (approximately 110 ms) and occipito-parieto-temporal N1 (approximately 150 ms) components. However, no such attentional modulations were found on the earlier C1 visual component, best observed with a negative polarity for upper visual field stimulations, and thought to originate largely from primary visual cortex. CONCLUSIONS AND SIGNIFICANCE: These results show that reflexive shifts of attention following oblique eye gaze to upper and lower visual fields increase and speed up the processing of visual information beyond the feedforward flow of information in primary visual cortex.     

Attention to central and peripheral visual space in a movement detection task: an event-related potential and behavioral study. I. Normal hearing adults

The effects of focussed attention to peripherally and centrally located visual stimuli were compared via an analysis of event-related brain potentials (ERPs) while subjects detected the direction of motion of a white square in a specified location. While attention to both peripheral and foveal stimuli produced enhancements of the early ERP components, the distribution over the scalp of the attention-related changes varied according to stimulus location. The attention-related increase in the amplitude of the N1 wave (157 ms) to the peripheral stimuli was greater over the parietal region of the hemisphere contralateral to the attended visual field. By contrast, the largest effects of foveally directed attention occurred over the occipital regions where the increase was bilaterally symmetrical. Additionally, the effects of attention on the ERPs were significantly larger for moving than for stationary stimuli, and this effect was greater for peripheral than for central attention. A long-latency positive displacement component (300-600 ms) was larger over the right than the left hemisphere during attention to the lateral visual fields, but was symmetrical in amplitude when central stimuli were attended. These results suggest that different pathways are modulated when attention is deployed to different regions of the visual fields. Further, they suggest that the special role of the right hemisphere in spatial attention may be limited to analysis of information in the visual periphery.     

Neural correlates of within-level and across-level attention to multiple compound stimuli

Event-related potentials (ERPs) were recorded to investigate the neural mechanisms of attention to the same or different levels of two compound letters presented concurrently in the left and right visual fields, respectively. Relative to the condition when attention was allocated to the global level of one compound stimulus and the local level of another one (across-level attention), attention to the same level of the two compound stimuli (within-level attention) increased an early positivity between 100 and 140 ms (P1) over the occipito-parietal cortex. A long-latency positivity between 320 and 560 ms (P3) over the central-parietal area was also increased in the within-level relative to across-level attention conditions. The ERP results suggest that, relative to across-level attention, within-level attention to multiple compound stimuli facilitates both early sensory-perceptual processing and late process of stimulus evaluation and identification in hierarchical analysis.     

Evoked phase synchronization between adjacent high-density electrodes in human scalp EEG: duration and time course related to behavior.

OBJECTIVE: Data from a previous event-related potential (ERP) study in visual-perceptual grouping [Nikolaev AR, van Leeuwen C. Flexibility in spatial and non-spatial feature grouping: an event-related potentials study. Brain Res Cogn Brain Res 2004;22:13-25] were re-analyzed to identify event-related dynamics of phase-synchronization. METHODS: In 20 Hz activity, uniform spreading of phase synchronization in closely spaced (approximately 2 cm) scalp electrodes appears and disappears spontaneously. The lengths of synchronized activity intervals and how they vary as a function of stimulus presentation were compared between task and control conditions. RESULTS: Synchronization reached a maximum in the task condition about 180 ms post-stimulus onset, coinciding with the peak N180 ERP marking the deployment of task-specific attention. Synchronized intervals were longer in the task than in the control condition. Long (above 80 ms) intervals occurred at a stable rate before and just after stimulus onset, but steeply decreased 200-400 ms afterwards. CONCLUSIONS: Perceptual tasks lead to longer synchronized intervals in early visual areas. Attention deployment resets the ongoing synchronization. Event-related activity, besides low-frequency ERP, consists of high-frequency short and long synchronized intervals corresponding to evoked bursts and ongoing oscillations, respectively. SIGNIFICANCE: High-density scalp recorded EEG revealed synchronization dynamics in a local, early visual area of cortex that can be interpreted as modulation of spontaneous ongoing task-related processes by attention.     

Bimodal speech: early suppressive visual effects in human auditory cortex

While everyone has experienced that seeing lip movements may improve speech perception, little is known about the neural mechanisms by which audiovisual speech information is combined. Event-related potentials (ERPs) were recorded while subjects performed an auditory recognition task among four different natural syllables randomly presented in the auditory (A), visual (V) or congruent bimodal (AV) condition. We found that: (i) bimodal syllables were identified more rapidly than auditory alone stimuli; (ii) this behavioural facilitation was associated with cross-modal [AV-(A+V)] ERP effects around 120-190 ms latency, expressed mainly as a decrease of unimodal N1 generator activities in the auditory cortex. This finding provides evidence for suppressive, speech-specific audiovisual integration mechanisms, which are likely to be related to the dominance of the auditory modality for speech perception. Furthermore, the latency of the effect indicates that integration operates at pre-representational stages of stimulus analysis, probably via feedback projections from visual and/or polymodal areas.     

The time course of semantic category processing in the cerebral hemispheres: an electrophysiological study

Using visual half-field presentations of words to the right (RVF) and to the left visual field (LVF), this study investigated the time course of the hemispheric involvement in the processing of semantic category information. Multi-channel event related brain potentials (ERPs) were recorded from 15 healthy subjects during a categorisation task of sequentially presented word pairs. Subjects had to judge mentally after the appearance of the second word whether the words of a pair were semantically related (SR) or not (SU). ERPs were computed, from 100 ms before the onset of the second word to 600 ms, for SR and SU conditions in the LVF and in the RVF separately. The temporal segmentation of ERP map series into sequences of quasi-stable map configurations revealed a total of seven segments in each visual field of which only the first five (S1-S5, appearing between 70 and 400 ms) showed different map configurations as a function of visual field but presented a similar temporal sequence in both visual fields. By contrast, of the last two segments (S6 and S7) which appeared between approximately 400 and approximately 600 ms, only S7 differentiated SR and SU conditions in terms of its duration. Source localisation analysis of the segments showed that following the initial activation of posterior brain regions as a function of the visual field of presentation, a common neural network was activated in the left hemisphere (LH) although the dynamics of activation varied as a function of visual field. Concerning the role of the right hemisphere (RH) in lexico-semantic processing, the results presented here appear to be compatible with a 'callosal relay model' and suggest that, in healthy subjects, information is transferred rapidly ( approximately 150 ms) from the RH to the language dominant-LH.     

Asymmetry of the human visual field in magnetic response to apparent motion

Predominance of the lower visual field has been shown in various visual tasks, but whether the upper visual field is involved in a specific neural process is unknown. We used magnetoencephalography to study the effect of orientation and direction on the responses of five subjects to apparent motion from the human extrastriate cortex. The first magnetic response always was the largest, and the peak latency of about 200 ms did not change with the stimulus conditions. Amplitudes of the first responses were highest when motions were oriented at the horizontal meridian, decreasing with the degree of the angle between motion orientation and the horizontal meridian. There was no difference in amplitude between the two directions in the lower visual field, whereas the value of the response to downward motion in the upper visual field was significantly larger than that to upward motion. These amplitude changes are not due to differences in the anatomical distribution of neural activities because the estimated origins for the first responses always were in the same cortical area (around the occipito-parieto-temporal region) and the directions of the current vectors did not change with the stimulus conditions, and the estimated current strength changed with the stimulus conditions as did the response amplitude. These findings suggest that the human extrastriate cortex has a directional preference for downward versus upward motion in the upper visual field.     

Attention changes the peak latency of the visual gamma-band oscillation of the EEG.

To investigate the physiological role of visual gamma-band oscillation (GBO), we calculated the event-related dynamics of the EEG power-spectrum for paired visual stimuli (S1 and S2) with or without attention in 12 subjects. The visual stimuli elicited transient increases in the GBO power (around 40 Hz), which were maximal over the parietal area. The peak GBO increase appeared around 300 ms after stimulus onset, but its latency was shorter after S1 and longer after S2 under the 'with attention' than under the 'without attention' condition. This transient increase in the visual GBO is thought to reflect attention and to reset the activity of the visual system in preparation for a new stimulus.     

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.     

Spatio-temporal dynamics of attention to color: Evidence from human electrophysiology

This study characterized patterns of brain electrical activity associated with selective attention to the color of a stimulus. Multichannel recordings of event-related potentials (ERPs) were obtained while subjects viewed randomized sequences of checkerboards consisting of isoluminant red or blue checks superimposed on a grey background. Stimuli were presented foveally at a rapid rate, and subjects were required to attend to the red or blue checks in separate blocks of trials and to press a button each time they detected a dimmer target stimulus of the attended color. An early negative ERP component with an onset latency of 50 ms was sensitive to stimulus color but was unaffected by the attentional manipulation. ERPs elicited by attended and unattended stimuli began to diverge after approximately 100 ms following stimulus onset. Inverse dipole modelling of the attended-minus-unattended difference waveform indicated that an initial positive deflection with an onset latency of 100 ms had a source in lateral occipital cortex, while a subsequent negative deflection with an onset at 160 ms had a source in inferior occipito-temporal cortex. Longer-latency attention-sensitive components were localized to premotor frontal areas (onset at 190 ms) and to more anterior regions of the fusiform gyrus (onset at 240 ms). These source localizations correspond closely with cortical areas that were identified in previous neuroimaging studies as being involved in color-selective processing. The present ERP data thus provide information about the time course of stimulus selection processes in cortical areas that subserve attention to color. Hum. Brain Mapping 6:216–238, 1998. © 1998 Wiley-Liss, Inc.     

Covariations in ERP and PET measures of spatial selective attention in human extrastriate visual cortex

In a previous study using positron emission tomography (PET), we demonstrated that focused attention to a location in the visual field produced increased regional cerebral blood flow in the fusiform gyrus contralateral to the attended hemifield (Heinze et al. [1994]: Nature 372:543). We related these effects to modulations in the amplitude of the P1 component (80–130 msec latency) of the visual event-related brain potentials (ERPs) recorded from the same subjects, under the identical stimulus and task conditions. Here, we replicate and extend these findings by showing that attention effects in the fusiform gyrus and the P1 component were similarly modulated by the perceptual load of the task. When subjects performed a perceptually demanding symbol-matching task within the focus of spatial attention, the fusiform activity and P1 component of the ERP were of greater magnitude than when the subjects performed a less perceptually demanding task that required only luminance detection at the attended location. In the latter condition, both the PET and ERP attention effects were reduced. In addition, in the present data significant activations were also obtained in the middle occipital gyrus contralateral to the attended hemifield, thereby demonstrating that multiple regions of extrastriate visual cortex are modulated by spatial attention. The findings of covariations between the P1 attention effect and activity in the posterior fusiform gyrus reinforce our hypothesis that common neural sources exist for these complementary, but very different measures of human brain activity. Hum. Brain Mapping 5:273–279, 1997. © 1997 Wiley-Liss, Inc.