Attended Location Research Articles

Looking away to see: The acquisition of a search habit away from the saccade direction

Growing evidence has shown that attention can be habit-like, unconsciously and persistently directed toward locations that have frequently contained search targets in the past. The attentional preference typically arises when the eye gaze aligns with the attended location. Here we tested whether this spatial alignment is necessary for the acquisition of a search habit. To divert eye movements away from an attended location, we used gaze-contingent eye tracking, restricting the visible portion of the screen to an area opposite to the current gaze. Participants searched for a T target amidst a circular array of L distractors. Unbeknownst to them, the target appeared more frequently in one screen quadrant. Despite fixating on a location diametrically opposite to the visible, attended region, participants acquired probability cuing, producing quicker responses when the target appeared in the high-probability quadrant. They also showed a speed advantage in the diagonal quadrant. The attentional preference for the high-probability quadrant persisted during a testing phase in which the target’s location was unbiased, but only when participants continued to search with the restricted view. These results indicate that a search habit can be acquired even when participants are required to look away from the high-probability locations. The finding suggests that the learned search habit is not solely a result of oculomotor learning.

Distinct roles of forward and backward alpha-band waves in spatial visual attention.

Previous research has associated alpha-band [8-12 Hz] oscillations with inhibitory functions: for instance, several studies showed that visual attention increases alpha-band power in the hemisphere ipsilateral to the attended location. However, other studies demonstrated that alpha oscillations positively correlate with visual perception, hinting at different processes underlying their dynamics. Here, using an approach based on traveling waves, we demonstrate that there are two functionally distinct alpha-band oscillations propagating in different directions. We analyzed EEG recordings from three datasets of human participants performing a covert visual attention task (one new dataset with N = 16, two previously published datasets with N = 16 and N = 31). Participants were instructed to detect a brief target by covertly attending to the screen's left or right side. Our analysis reveals two distinct processes: allocating attention to one hemifield increases top-down alpha-band waves propagating from frontal to occipital regions ipsilateral to the attended location, both with and without visual stimulation. These top-down oscillatory waves correlate positively with alpha-band power in frontal and occipital regions. Yet, different alpha-band waves propagate from occipital to frontal regions and contralateral to the attended location. Crucially, these forward waves were present only during visual stimulation, suggesting a separate mechanism related to visual processing. Together, these results reveal two distinct processes reflected by different propagation directions, demonstrating the importance of considering oscillations as traveling waves when characterizing their functional role.

Spatial Attention Tunes Temporal Processing in Early Visual Cortex by Speeding and Slowing Alpha Oscillations.

The perception of dynamic visual stimuli relies on two apparently conflicting perceptual mechanisms: rapid visual input must sometimes be integrated into unitary percepts but at other times must be segregated or parsed into separate objects or events. Though they have opposite effects on our perceptual experience, the deployment of spatial attention benefits both operations. Little is known about the neural mechanisms underlying this impact of spatial attention on temporal perception. Here, we record magnetoencephalography (MEG) in male and female humans to demonstrate that the deployment of spatial attention for the purpose of segregating or integrating visual stimuli impacts prestimulus oscillatory activity in retinotopic visual brain areas where the attended location is represented. Alpha band oscillations contralateral to an attended location are therefore faster than ipsilateral oscillations when stimuli appearing at this location will need to be segregated, but slower in expectation of the need for integration, consistent with the idea that α frequency is linked to perceptual sampling rate. These results demonstrate a novel interaction between temporal visual processing and the allocation of attention in space.SIGNIFICANCE STATEMENT Our environment is dynamic and visual input therefore varies over time. To make sense of continuously changing information, our visual system balances two complementary processes: temporal segregation in order to identify changes, and temporal integration to identify consistencies in time. When we know that a circumstance requires use of one or the other of these operations, we are able to prepare for this, and this preparation can be tracked in oscillatory brain activity. Here, we show how this preparation for temporal processing can be focused spatially. When we expect to integrate or segregate visual stimuli that will appear at a specific location, oscillatory brain activity changes in visual areas responsible for the representation of that location. In this way, spatial and temporal mechanisms interact to support adaptive, efficient perception.

Age-related Changes in Cortical Excitability Linked to Decreased Attentional and Inhibitory Control

Understanding age-related changes in cortical excitability and their relation to cognitive functions will help to improve interventions based on non-invasive brain stimulation that aim to support cognitive function in older adults. Here, we investigate the relationship between cortical excitability, executive function, and underlying neural activity in samples of healthy young and older adults. These participants performed a Simon task during electroencephalogram (EEG) recording. During the task, participants had to respond to the colour of a lateralized stimulus while ignoring its spatial location. We studied event-related brain potential correlates of attentional and inhibitory control [i.e., the posterior contralateral negativity (N2pc) and central contralateral negativity (N2cc), respectively] related to the Simon task performance. We also used transcranial magnetic stimulation (TMS) EEG coregistration. In detail, we applied single-pulse TMS during EEG recording in order to analyse global mean field power (GMFP) and TMS-evoked potentials (TEPs) as correlates of cortical excitability. We found lower GMFP amplitude within 101–200 ms in older compared to young adults. Moreover, older adults showed smaller N45 amplitude and slower P180 latency. These findings suggest cortical excitability alterations related to ageing. Older adults also exhibited longer reaction times and N2pc and N2cc latencies, indicating that it took them longer to allocate attention to the target stimulus and inhibit the tendency to respond to the attended location. Finally, in older adults, cortical excitability alterations correlated with longer reaction times and N2pc latencies. These results suggest that age-related alterations in cortical excitability represent a dysfunctional change associated with physiological ageing.

Sustained Attention and Spatial Attention Distinctly Influence Long-term Memory Encoding.

Our attention is critically important for what we remember. Prior measures of the relationship between attention and memory, however, have largely treated "attention" as a monolith. Here, across three experiments, we provide evidence for two dissociable aspects of attention that influence encoding into long-term memory. Using spatial cues together with a sensitive continuous report procedure, we find that long-term memory response error is affected by both trial-by-trial fluctuations of sustained attention and prioritization via covert spatial attention. Furthermore, using multivariate analyses of EEG, we track both sustained attention and spatial attention before stimulus onset. Intriguingly, even during moments of low sustained attention, there is no decline in the representation of the spatially attended location, showing that these two aspects of attention have robust but independent effects on long-term memory encoding. Finally, sustained and spatial attention predicted distinct variance in long-term memory performance across individuals. That is, the relationship between attention and long-term memory suggests a composite model, wherein distinct attentional subcomponents influence encoding into long-term memory. These results point toward a taxonomy of the distinct attentional processes that constrain our memories.

Selective auditory attention modulates cortical responses to sound location change in younger and older adults.

The present study measured scalp potentials in response to low-frequency, narrowband noise bursts changing location in the front, azimuthal plane. At question was whether selective auditory attention has a modulatory effect on the cortical encoding of spatial change and whether older listeners with normal-hearing thresholds would show depressed cortical representation for spatial changes relative to younger listeners. Young and older normal-hearing listeners were instructed to either passively listen to the stimulus presentation or actively attend to a single location (either 30° left or right of midline) and detect when a noise stream moved to the attended location. Prominent peaks of the electroencephalographic scalp waveforms were compared across groups, locations, and attention conditions. In addition, an opponent-channel model of spatial coding was performed to capture the effect of attention on spatial-change tuning. Younger listeners showed not only larger responses overall but a greater dynamic range in their response to location changes. Results suggest that younger listeners were acquiring and encoding key spatial cues at early cortical processing areas. On the other hand, each group exhibited modulatory effects of attention to spatial-change tuning, indicating that both younger and older listeners selectively attend to space in a manner that amplifies the available signal.NEW & NOTEWORTHY In complex acoustic scenes, listeners take advantage of spatial cues to selectively attend to sounds that are deemed immediately relevant. At the neural level, selective attention amplifies electrical responses to spatial changes. We tested whether older and younger listeners have comparable modulatory effects of attention to stimuli moving in the free field. Results indicate that although older listeners do have depressed overall responses, selective attention enhances spatial-change tuning in younger and older listeners alike.

Cross-modal orienting of exogenous attention results in visual-cortical facilitation, not suppression

Attention may be oriented exogenously (i.e., involuntarily) to the location of salient stimuli, resulting in improved perception. However, it is unknown whether exogenous attention improves perception by facilitating processing of attended information, suppressing processing of unattended information, or both. To test this question, we measured behavioral performance and cue-elicited neural changes in the electroencephalogram as participants (N = 19) performed a task in which a spatially non-predictive auditory cue preceded a visual target. Critically, this cue was either presented at a peripheral target location or from the center of the screen, allowing us to isolate spatially specific attentional activity. We find that both behavior and attention-mediated changes in visual-cortical activity are enhanced at the location of a cue prior to the onset of a target, but that behavior and neural activity at an unattended target location is equivalent to that following a central cue that does not direct attention (i.e., baseline). These results suggest that exogenous attention operates via facilitation of information at an attended location.

What factors influence the switch from unitary to divided attention?

The focus of attention can be either unitary or divided and can transition from unitary to divided while performing a task. In Experiment 1, we investigated whether alerting hastens the transition from unitary to divided attention. To this end, we employed a dual-RSVP-stream Attentional Blink task (AB; impaired perception of the second of two rapidly sequential targets) with two pairs of letter targets (T1-pair and T2-pair). One component of the AB known as Lag-1 sparing (unimpaired perception of the T2-pair when it is presented directly after the T1-pair) occurs only when the T2-pair falls in an attended location. When the T2-pair falls in an unattended location, the converse pattern occurs (Lag-1 deficit). Accordingly, we used the incidence of Lag-1 sparing/deficit to index whether a location was attended or unattended. We found that presenting a brief brightening flash of the screen (alerting) just before the T1-pair hastened the transition from the initial unitary focus to a divided focus. In Experiment 2, we pitted the hastening account against an alternative hypothesis that the flash triggers phasic activation of the Locus Coeruleus-norepinephrine neuromodulatory system, thus resetting the underlying neural networks that mediate the distribution of attention, triggering a switch from unitary to divided attention. The results of Experiment 2 were incompatible with the hastening account, but consistent with the network-reset account.

Effects of eccentricity on the attention-related N2pc component of the event-related potential waveform.

The N2pc ERP component has been widely used as a measure of lateralized visual attention. It is characterized by a negativity contralateral to the attended location or target, and it is thought to reflect contralaterally enhanced processing of attended information in intermediate to high levels of the ventral visual pathway. Given that the receptive fields in these areas often extend a few degrees into the ipsilateral hemifield, we might expect that near-midline stimuli would be processed by both the contralateral and ipsilateral hemispheres, resulting in a diminished N2pc. However, little is known about the effect of eccentricity on the N2pc component. To address this gap in knowledge, we recorded the EEG while participants performed a discrimination task with stimuli presented at one of five eccentricities (0°, 0.05°, 1°, 2°, 4° and 8° between the inner edge of the stimulus and the midline). We found that the N2pc amplitude remained relatively constant across eccentricities, including when the inner edge was at the midline, except that N2pc amplitude was reduced by more than 50% at the greatest eccentricity (8°). We also examined the contralateral positivity that often follows the N2pc. This positivity became progressively larger, and the transition from negative to positive occurred progressively later, as the eccentricity increased. These findings suggest that future experiments looking at the N2pc can use near-midline stimuli without compromising the N2pc amplitude, but should avoid large eccentricities. Implications about the neural generators of the N2pc are also discussed.

Attentional set and the gradient of visual spatial attention

There are a number of factors that may influence the shape and scope of the gradient of visual attention. This study examined the role of attentional set in determining the scope and presence of suppression in the gradient of visual selective attention observed in the P1 event related potential waveform. Twenty-two participants were asked to attend to a specific location and respond to a target stimulus. However, thirty percent of the time, the stimulus could also appear at one of 7 other locations (three ipsilateral to the attended location and four contralateral). Attentional set was varied by changes in the instructions to avoid confounds created by changing the stimuli. In half of the study participants were asked to respond only to stimuli that appeared at the designated location. In the other half of the study participants were asked to attend to the designated location but respond to all stimuli. Results demonstrate a significant main effect of attentional set, with greater processing when participants were asked to respond to all stimuli as compared to when they were only asked to respond to stimuli at the designated location. Additionally a significant interaction between visual field and location shows greater differences in processing (i.e. an attentional effect) at locations closer to the designated location. Findings are discussed in relation to the scope and gradient of attention.

Neural Activation During Visual Attention Differs in Individuals with High versus Low Macular Pigment Density.

The neural efficiency hypothesis for lutein (L) and zeaxanthin (Z) suggests that higher levels of L+Z in the central nervous system (CNS) are predictive of stronger stimulus-specific brain responses. Past research suggests that supplementing L+Z can improve neural processing speed and cognitive function across multiple domains, which supports this hypothesis. The purpose of this study is to determine the extent to which CNS L+Z levels predict brain responses using an attentionally taxing task. Macular pigment optical density (MPOD) is measured at baseline in 85 participants ranging in age from 18-92 years. Brain activation is measured using dense array electroencephalography. Stimuli evoking the signal include a grating array of vertical bars, oscillating at four driving frequencies. Significant stimulus-specific interactions are detected between attend condition, location, and age (p < .002) for unattended image locations, and between age and location (p < .008) for attended locations. Although no differences are found across age by MPOD, this measure is found to be predictive of neural power at parafoveal bar locations (R2 .080). CNS L+Z status is related to differences in brain activation in conditions designed to stress visual attention. These differences are strongest for older subjects.

Attending to What and Where: Background Connectivity Integrates Categorical and Spatial Attention

Top-down attention prioritizes the processing of goal-relevant information throughout visual cortex based on where that information is found in space and what it looks like. Whereas attentional goals often have both spatial and featural components, most research on the neural basis of attention has examined these components separately. Here we investigated how these attentional components are integrated by examining the attentional modulation of functional connectivity between visual areas with different selectivity. Specifically, we used fMRI to measure temporal correlations between spatially selective regions of early visual cortex and category-selective regions in ventral temporal cortex while participants performed a task that benefitted from both spatial and categorical attention. We found that categorical attention modulated the connectivity of category-selective areas, but only with retinotopic areas that coded for the spatially attended location. Similarly, spatial attention modulated the connectivity of retinotopic areas only with the areas coding for the attended category. This pattern of results suggests that attentional modulation of connectivity is driven both by spatial selection and featural biases. Combined with exploratory analyses of frontoparietal areas that track these changes in connectivity among visual areas, this study begins to shed light on how different components of attention are integrated in support of more complex behavioral goals.

Influence of spatial and non-spatial feature continuity on cortical alpha oscillations

In everyday environments, listeners face the challenge of parsing the sound mixture reaching their ears into individual sources. Perceptual continuity of acoustic features, such as pitch or location, has an obligatory effect on auditory attention even when a feature is not task relevant. Cortical alpha oscillations (8-12 Hz) are thought to functionally inhibit the processing of task-irrelevant information. Here, we hypothesize that discontinuities in a task-irrelevant feature disrupt the attentional modulation of alpha rhythms. Using electroencephalography in humans, we compare physiological measures during a selective auditory attention task where listeners were asked to attend to either talker, based on gender (male or female) or location (left or right). On half of the trials, a discontinuity was introduced in the task-irrelevant acoustic feature. When listeners attended to the talker, there was no evidence of alpha power lateralization, and no effect of a discontinuity in location. In contrast, when listeners attended to location, parieto-occipital alpha power increased ipsilateral to the attended location; moreover, a discontinuity in talker reduced alpha power and disrupted alpha lateralization. Our findings support the importance of parieto-occipital alpha in suppressing sources when listeners focus spatial, but not non-spatial, attention, and show that task-irrelevant discontinuities affect these alpha rhythms.

Mechanisms for attentional modulation by threatening emotions of fear, anger, and disgust

Appropriately attending to threatening environmental stimuli is evolutionarily adaptive and crucial for survival. This study revealed that nonconscious attentional modulation of disgust has different behavioral and event-related potential (ERP) patterns, as compared to fear and anger. To facilitate its evolutionary purpose of avoidance, disgust first diverts rather than attracts attention. Accordingly, the N1 was smaller in a validly than in an invalidly disgust-cued condition. Furthermore, the frontal P3a for disgust, anger, and fear was found to be larger in the valid than in the invalid condition, which was interpreted as an involuntary switching of attention toward threat-related events to mobilize cognitive resources for action or defense. On the contrary, the parietal P3b only occurred at the conscious level; the enhanced P3b indicated that more cognitive resources were being allocated toward the task-relevant but previously less attended location, to ensure the effective achievement of task goals. In addition, group comparisons between individuals with low and high disgust sensitivity showed that the ERP differences between the disgust and the anger/fear conditions at the unconscious level may be attributed only to individuals with high disgust sensitivity. These findings, together with previous knowledge of the effects of fear and anger on attention, strengthen our confidence in the two-stage scheme of attentional modulation by threats, which consists of an early stage of bottom-up response scaling of sensory processing (reflected by the P1 and N1) and a later stage of top-down integration and regulation of emotion and behavior (reflected by the P3).

Decoding and Reconstructing the Focus of Spatial Attention from the Topography of Alpha-band Oscillations.

Many aspects of perception and cognition are supported by activity in neural populations that are tuned to different stimulus features (e.g., orientation, spatial location, color). Goal-directed behavior, such as sustained attention, requires a mechanism for the selective prioritization of contextually appropriate representations. A candidate mechanism of sustained spatial attention is neural activity in the alpha band (8-13 Hz), whose power in the human EEG covaries with the focus of covert attention. Here, we applied an inverted encoding model to assess whether spatially selective neural responses could be recovered from the topography of alpha-band oscillations during spatial attention. Participants were cued to covertly attend to one of six spatial locations arranged concentrically around fixation while EEG was recorded. A linear classifier applied to EEG data during sustained attention demonstrated successful classification of the attended location from the topography of alpha power, although not from other frequency bands. We next sought to reconstruct the focus of spatial attention over time by applying inverted encoding models to the topography of alpha power and phase. Alpha power, but not phase, allowed for robust reconstructions of the specific attended location beginning around 450 msec postcue, an onset earlier than previous reports. These results demonstrate that posterior alpha-band oscillations can be used to track activity in feature-selective neural populations with high temporal precision during the deployment of covert spatial attention.

Retrospective triggering of conscious perception by an interstimulus interaction.

Attention facilitates conscious perception of a visual stimulus at an attended location. Interestingly, a recent study (using the Posner spatial-cueing task) reported that attention facilitated conscious perception even when it was cued after a stimulus was gone (postcued-attention or retroperception effect). Here, we show that this effect can be induced without any contribution of attention. Contrary to previous situations, we fixed a position of a target (Gabor patch) and cue (luminance change of a circle encompassing the target) across trials so that subjects always could allocate their full attention to the target position. The cue (luminance change) improved objective and subjective visibility of the nearby target even when it was given ∼200 ms after the target's offset. This retrospective improvement was diminished when a shape of the cue was changed from a circle to a dot pattern, suggesting that the improvement emerged from a visual interaction (combinations of shapes) between the circular cue and target. Those results indicated that a local visual interaction between the target and cue is sufficient to trigger consciousness of the target, revealing a new type of retroperception effect mediated by sensory (nonattentional) mechanisms.

The Role of Right Inferior Parietal Cortex in Auditory Spatial Attention: A Repetitive Transcranial Magnetic Stimulation Study.

Behavioral studies support the concept of an auditory spatial attention gradient by demonstrating that attentional benefits progressively diminish as distance increases from an attended location. Damage to the right inferior parietal cortex can induce a rightward attention bias, which implicates this region in the construction of attention gradients. This study used event-related potentials (ERPs) to define attention-related gradients before and after repetitive transcranial magnetic stimulation (rTMS) to the right inferior parietal cortex. Subjects (n = 16) listened to noise bursts at five azimuth locations (left to right: -90°, -45°, 0° midline, +45°, +90°) and responded to stimuli at one target location (-90°, +90°, separate blocks). ERPs as a function of non-target location were examined before (baseline) and after 0.9 Hz rTMS. Results showed that ERP attention gradients were observed in three time windows (frontal 230–340, parietal 400–460, frontal 550–750 ms). Significant transient rTMS effects were seen in the first and third windows. The first window had a voltage decrease at the farthest location when attending to either the left or right side. The third window had on overall increase in positivity, but only when attending to the left side. These findings suggest that rTMS induced a small contraction in spatial attention gradients within the first time window. The asymmetric effect of attended location on gradients in the third time window may relate to neglect of the left hemispace after right parietal injury. Together, these results highlight the role of the right inferior parietal cortex in modulating frontal lobe attention network activity.

Motor preparation and attentional benefits: dependencies on the number of possible saccade targets

Growing evidence suggests movements are prepared through widespread and complex modulation in multiple brain centers, where inhibitory components assure execution is not triggered prematurely (Cohen, Sherman, Zinger, Perlmutter & Prut, 2010). It has been suggested that preparing an eye movement towards few possible targets, compared to many, results in longer latencies because of the activation of these inhibitory components (Lawrence, John, Abrams & Snyder, 2008). In contrast, Belopolsky & Theeuwes (2009) suggest that attending to a location that has a high probability of being a saccade target, because of only few available options, results in the activation of the oculomotor system and facilitation. Our goal was to clarify this discrepancy and to measure attention at object locations when saccade target alternatives were few or many. While the total number of presented stimuli was kept constant (twelve circles), the saccade target could be at only two possible circle locations in the few alternatives condition, while the many alternatives condition featured six options. We employed a dual-task paradigm, primarily requiring the rapid execution of saccades to the only circle that became the target colour. The secondary task was to discriminate an asymmetric cross flashed at one of the circles. Our results show that slower saccade initiation is found when the possible saccade targets are two, indeed suggesting inhibitory motor components may be at play. However, the discrimination scores show that perception at the saccade target and the (never fixated) alternative saccade target were both facilitated compared to an attended control location. This suggests motor suppression may not necessarily affect perception. Recently Dhawan, Deubel & Jonikaitis (2013) showed that inhibition of saccades led to perceptual suppression, while in our study perception appeared unaffected. We put forward attention-based interpretations focusing on attentional window size and fuzzy loci leading to more stimulus-driven eye movements. Meeting abstract presented at VSS 2015. Language: en

Rapid cortical dynamics associated with auditory spatial attention gradients

Behavioral and EEG studies suggest spatial attention is allocated as a gradient in which processing benefits decrease away from an attended location. Yet the spatiotemporal dynamics of cortical processes that contribute to attentional gradients are unclear. We measured EEG while participants (n = 35) performed an auditory spatial attention task that required a button press to sounds at one target location on either the left or right. Distractor sounds were randomly presented at four non-target locations evenly spaced up to 180° from the target location. Attentional gradients were quantified by regressing ERP amplitudes elicited by distractors against their spatial location relative to the target. Independent component analysis was applied to each subject's scalp channel data, allowing isolation of distinct cortical sources. Results from scalp ERPs showed a tri-phasic response with gradient slope peaks at ~300 ms (frontal, positive), ~430 ms (posterior, negative), and a plateau starting at ~550 ms (frontal, positive). Corresponding to the first slope peak, a positive gradient was found within a central component when attending to both target locations and for two lateral frontal components when contralateral to the target location. Similarly, a central posterior component had a negative gradient that corresponded to the second slope peak regardless of target location. A right posterior component had both an ipsilateral followed by a contralateral gradient. Lateral posterior clusters also had decreases in α and β oscillatory power with a negative slope and contralateral tuning. Only the left posterior component (120–200 ms) corresponded to absolute sound location. The findings indicate a rapid, temporally-organized sequence of gradients thought to reflect interplay between frontal and parietal regions. We conclude these gradients support a target-based saliency map exhibiting aspects of both right-hemisphere dominance and opponent process models.

Dynamics of EEG rhythms support distinct visual selection mechanisms in parietal cortex: a simultaneous transcranial magnetic stimulation and EEG study.

Using repetitive transcranial magnetic stimulation (rTMS), we have recently shown a functional anatomical distinction in human parietal cortex between regions involved in maintaining attention to a location [ventral intraparietal sulcus (vIPS)] and a region involved in shifting attention between locations [medial superior parietal lobule (mSPL)]. In particular, while rTMS interference over vIPS impaired target discrimination at contralateral attended locations, interference over mSPL affected performance following shifts of attention regardless of the visual field (Capotosto et al., 2013). Here, using rTMS interference in conjunction with EEG recordings of brain rhythms during the presentation of cues that indicate to either shift or maintain spatial attention, we tested whether this functional anatomical segregation involves different mechanisms of rhythm synchronization. The transient inactivation of vIPS reduced the amplitude of the expected parieto-occipital low-α (8-10 Hz) desynchronization contralateral to the cued location. Conversely, the transient inactivation of mSPL, compared with vIPS, reduced the high-α (10-12 Hz) desynchronization induced by shifting attention into both visual fields. Furthermore, rTMS induced a frequency-specific delay of task-related modulation of brain rhythms. Specifically, rTMS over vIPS or mSPL during maintenance (stay cues) or shifting (shift cues) of spatial attention, respectively, caused a delay of α parieto-occipital desynchronization. Moreover, rTMS over vIPS during stay cues caused a delay of δ (2-4 Hz) frontocentral synchronization. These findings further support the anatomo-functional subdivision of the dorsal attention network in subsystems devoted to shifting or maintaining covert visuospatial attention and indicate that these mechanisms operate in different frequency channels linking frontal to parieto-occipital visual regions.