Causal topography of visual cortex in perceptual learning.

Individuals are able to improve their visual skill with practice, a phenomenon called Visual Perceptual Learning (VPL). We previously observed that after training on a difficult shape identification task, the dorsal visual regions (i.e. right V2d/V3 and right lateral occipital, LO) corresponding to the trained visual quadrant, and their homologous in the opposite hemisphere, exhibited a selective activation at the end of the learning. By contrast, such modulation was not observed in the ventral visual regions, corresponding to the untrained quadrants. The causal role of the trained visual cortex was previously showed in a TMS study as its inactivation impaired behavioral performance to learned stimuli. Here, using the same experimental design, we employed TMS over the homologous of the trained area (i.e. left V2d/V3) as well as over the untrained region (i.e. right V4) to causally map the visual network during the perceptual learning. We report a decrease of accuracy after TMS over left V2d/V3 as compared to both right V4 and Sham (inactive stimulation) conditions. Importantly, TMS effect was correlated with the degree of learning, such that subjects with lower accuracy at the end of the training exhibited stronger TMS impairment. These results provide evidence that segregated regions within the visual network are causally involved in visual perceptual learning.

Magnetic stimulation selectively affects pre-stimulus EEG microstates.

Different electrophysiological (EEG) correlates may provide specific important assessment of the period that anticipates an imperative stimulus. Previous study of our group showed that a local (i.e. parietal) anticipatory EEG marker (i.e. the event related de-synchronization of the alpha rhythms; ERD) is selectively affected when transcranial magnetic stimulation (TMS) is delivered over crucial nodes belonging to well-known human networks involved in different cognitive domains. Here, we investigated whether such distinction is also present in the whole brain activity as seen through the pre-stimulus microstate's topography, representing a global and reference-free measure of the neural activity. First, when subjects received a pseudo-stimulation (sham), we found two distinct pre-stimulus topographies during perceptual or memory task, respectively. Second, we reported that, during the visuo-spatial attention task, stimulation of left intraparietal sulcus (IPS), but not left angular gyrus (AG), significantly modifies the topography observed in the Sham condition. Conversely, stimulation of AG, but not IPS, changes the topography observed in the Sham condition during a semantic memory task. These findings provide the first causal evidence for the task and region specificity of the pre-stimulus EEG microstates, thus proposing this EEG index as of particular interest for the assessment of the period that precedes a predictable event.

Task and Regions Specific Top-Down Modulation of Alpha Rhythms in Parietal Cortex.

Alpha (8-12 Hz) power desynchronization is strongly associated to visual perception but has been observed in a large variety of tasks, indicating a general role in task anticipation. We previously reported in human observers that interference by repetitive transcranial magnetic stimulation (rTMS) of core regions of the dorsal attention network (DAN) disrupts both anticipatory alpha desynchronization and performance during a visuospatial attention (VSA) task. Here, we test the hypothesis that alpha desynchronization is task specific, and can be selectively modulated by interfering with activity in different higher-order parietal regions. We contrast the effects of rTMS on alpha rhythms and behavior on 2 different tasks: a VSA and a semantic decision task, by targeting the posterior intraparietal sulcus (pIPS), a core region of the DAN, or the angular gyrus (AG), a core region of the default mode network (DMN). We found that both performance and anticipatory alpha desynchronization were affected by stimulation of IPS only during VSA, and of AG only during semantic decisions. These findings indicate the existence of multiple dedicated parietal channels for the modulation of anticipatory alpha rhythms, which in turn reflect task-specific modulation of excitability in human parieto-occipital cortex.

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.

Magnetic stimulation of visual cortex impairs perceptual learning.

The ability to learn and process visual stimuli more efficiently is important for survival. Previous neuroimaging studies have shown that perceptual learning on a shape identification task differently modulates activity in both frontal-parietal cortical regions and visual cortex (Sigman et al., 2005;Lewis et al., 2009). Specifically, fronto-parietal regions (i.e. intra parietal sulcus, pIPS) became less activated for trained as compared to untrained stimuli, while visual regions (i.e. V2d/V3 and LO) exhibited higher activation for familiar shape. Here, after the intensive training, we employed transcranial magnetic stimulation over both visual occipital and parietal regions, previously shown to be modulated, to investigate their causal role in learning the shape identification task. We report that interference with V2d/V3 and LO increased reaction times to learned stimuli as compared to pIPS and Sham control condition. Moreover, the impairment observed after stimulation over the two visual regions was positive correlated. These results strongly support the causal role of the visual network in the control of the perceptual learning.

The cerebral correlates of subliminal emotions: an eleoencephalographic study with emotional hybrid faces.

In a high-resolution electroencephalographic study, participants evaluated the friendliness level of upright and inverted 'hybrid faces', i.e. facial photos containing a subliminal emotional core in the low spatial frequencies (< 6 cycles/image), superimposed on a neutral expression in the rest of the spatial frequencies. Upright happy and angry faces were judged as more friendly or less friendly than neutral faces, respectively. We observed the time course of cerebral correlates of these stimuli with event-related potentials (ERPs), confirming that hybrid faces elicited the posterior emotion-related and face-related components (P1, N170 and P2), previously shown to be engaged by non-subliminal emotional stimuli. In addition, these components were stronger in the right hemisphere and were both enhanced and delayed by face inversion. A frontal positivity (210-300 ms) was stronger for emotional than for neutral faces, and for upright than for inverted faces. Hence, hybrid faces represent an original approach in the study of subliminal emotions, which appears promising for investigating their electrophysiological correlates.

Anatomical segregation of visual selection mechanisms in human parietal cortex.

Visual selection requires mechanisms for representing object salience and for shifting the focus of processing to novel objects. It is not clear from computational or neural models whether these operations are performed within the same or different brain regions. Here, we use repetitive transcranial magnetic stimulation to briefly interfere with neural activity in individually localized regions of human posterior parietal cortex (PPC) that are putatively involved in attending to contralateral locations or shifting attention between locations. Stimulation over right ventral intraparietal sulcus impaired target discrimination at contralateral locations, whereas stimulation over right medial superior parietal lobule impaired target discrimination after a shift of attention regardless of its location. This double dissociation is consistent with neuroimaging studies and indicates that mechanisms of visual selection are partly anatomically segregated in human PPC.

Resting-state modulation of α rhythms by interference with angular gyrus activity.

The default mode network is active during restful wakefulness and suppressed during goal-driven behavior. We hypothesize that inhibitory interference with spontaneous ongoing, that is, not task-driven, activity in the angular gyrus (AG), one of the core regions of the default mode network, will enhance the dominant idling EEG alpha rhythms observed in the resting state. Fifteen right-handed healthy adult volunteers underwent to this study. Compared with sham stimulation, magnetic stimulation (1 Hz for 1 min) over both left and right AG, but not over FEF or intraparietal sulcus, core regions of the dorsal attention network, enhanced the dominant alpha power density (8-10 Hz) in occipitoparietal cortex. Furthermore, right AG-rTMS enhanced intrahemispheric alpha coherence (8-10 Hz). These results suggest that AG plays a causal role in the modulation of dominant low-frequency alpha rhythms in the resting-state condition.

Differential contribution of right and left parietal cortex to the control of spatial attention: a simultaneous EEG-rTMS study.

We have recently shown that interference with repetitive transcranial magnetic stimulation (rTMS) of right posterior intraparietal sulcus (IPS) cortex during the allocation of spatial attention leads to abnormal desynchronization of anticipatory (pretarget) electroencephalographic alpha rhythms (8-12 Hz) in occipital-parietal cortex and the detection of subsequently presented visual targets (Capotosto et al. 2009). Since lesion data suggest that lesions of the right frontoparietal cortices produce more severe and long-lasting deficits of visual spatial attention than lesions of the left hemisphere, here, we used the mentioned rTMS-electroencephalographic procedure to test if the control of anticipatory alpha rhythms by IPS is asymmetrically organized in the 2 hemispheres. Results showed that interference with either left or right IPS during covert spatial attention equally disrupted the normally lateralized anticipatory modulation of occipital visual cortex, with stronger alpha desynchronization contralaterally to the attended visual field. In contrast, only interference with right IPS induced a paradoxical pretarget synchronization of alpha rhythms and bilateral deficits of target identification. These results suggest that the control of spatial topography of anticipatory alpha rhythms in occipital-parietal cortex is shared between left and right IPS cortex, but that right IPS uniquely contributes to a bilateral prestimulus activation of occipital visual cortex.

Visuo-spatial attention and semantic memory competition in the parietal cortex.

Neuroimaging studies associate specific functional roles to distinct brain regions investigating separate cognitive processes using dedicated tasks. For example, using both correlative (i.e., fMRI) and causal (i.e., TMS) approaches it has been shown the involvement of intra-parietal sulcus (IPS), as part of the dorsal attention network, in spatial attentional tasks as well as the importance of the angular gyrus (AG), as part of the default mode network, during the selection of relevant information in semantic memory. Nonetheless, in our daily life attention and semantic memory are rarely needed in isolation. In the present TMS study we investigate how the brain combines attentional and semantic memory demands in a single task. Results showed that, compared to a pseudo-TMS, stimulation of IPS, but not AG, affects behavioral performance, thus suggesting its preponderant role in such a combined task. Moreover, the lack of difference between the effect of IPS and AG stimulations seems to suggest that the two regions may be coactivated or that a third-party source might indirectly mediate the interaction between the two networks.

Neuroimaging evidence supporting a dual-network architecture for the control of visuospatial attention in the human brain: a mini review.

Neuroimaging studies conducted in the last three decades have distinguished two frontoparietal networks responsible for the control of visuospatial attention. The present review summarizes recent findings on the neurophysiological mechanisms implemented in both networks and describes the evolution from a model centered on the distinction between top-down and bottom-up attention to a model that emphasizes the dynamic interplay between the two networks based on attentional demands. The role of the dorsal attention network (DAN) in attentional orienting, by boosting behavioral performance, has been investigated with multiple experimental approaches. This research effort allowed us to trace a distinction between DAN regions involved in shifting vs. maintenance of attention, gather evidence for the modulatory influence exerted by the DAN over sensory cortices, and identify the electrophysiological correlates of the orienting function. Simultaneously, other studies have contributed to reframing our understanding of the functions of the ventral attention network (VAN) and its relevance for behavior. The VAN is not simply involved in bottom-up attentional capture but interacts with the DAN during reorienting to behaviorally relevant targets, exhibiting a general resetting function. Further studies have confirmed the selective rightward asymmetry of the VAN, proposed a functional dissociation along the anteroposterior axis, and suggested hypotheses about its emergence during the evolution of the primate brain. Finally, novel models of network interactions explain the expression of complex attentional functions and the emergence and restorations of symptoms characterizing unilateral spatial neglect. These latter studies emphasize the importance of considering patterns of network interactions for understanding the consequences of brain lesions.

Brain Topological Reorganization Associated with Visual Neglect After Stroke.

Background/Purpose: To identify brain hubs that are behaviorally relevant for neglect after stroke as well as to characterize their functional architecture of communication. Methods: Twenty acute right hemisphere damaged patients underwent neuropsychological and resting-state functional magnetic resonance imaging sessions. Spatial neglect was assessed by means of the Center of Cancellation on the Bells Cancellation Test. For each patient, resting-state functional connectivity matrices were derived by adopting a brain parcellation scheme consisting of 153 nodes. For every node, we extracted its betweenness centrality (BC) defined as the portion of all shortest paths in the connectome involving such node. Then, neglect hubs were identified as those regions showing a high correlation between their BC and neglect scores. Results: A first set of neglect hubs was identified in multiple systems including dorsal attention and ventral attention, default mode, and frontoparietal executive-control networks within the damaged hemisphere as well as in the posterior and anterior cingulate cortex. Such cortical regions exhibited a loss of BC and increased (i.e., less efficient) weighted shortest path length (WSPL) related to severe neglect. Conversely, a second group of neglect hubs found in visual and motor networks, in the undamaged hemisphere, exhibited a pathological increase of BC and reduction of WSPL associated with severe neglect. Conclusion: The topological reorganization of the brain in neglect patients might reflect a maladaptive shift in processing spatial information from higher level associative-control systems to lower level visual and sensory-motor processing areas after a right hemisphere lesion.

Reduced Segregation of Brain Networks in Spatial Neglect After Stroke.

Background/Purpose: To investigate the association between the degree of spatial neglect and the changes of brain system segregation (SyS; i.e., the ratio of the extent to which brain networks interact internally and with each other) after stroke. Methods: A cohort of 20 patients with right hemisphere lesion was submitted to neuropsychological assessment as well as to resting-state functional magnetic resonance imaging session at acute stage after stroke. The severity of spatial neglect was quantified using the Center of Cancellation (CoC) scores of the Bells cancellation test. For each patient, resting-state functional connectivity (FC) matrices were assessed by implementing a brain parcellation of nine networks that included the visual network, dorsal attention network (DAN), ventral attention network (VAN), sensorimotor network (SMN), auditory network, cingulo-opercular network, language network, frontoparietal network, and default mode network (DMN). For each patient and each network, we then computed the SyS derived by subtracting the between-network FC from the within-network FC (normalized by the within-network FC). Finally, for each network, the CoC scores were correlated with the SyS. Results: The correlational analyses indicated a negative association between CoC and SyS in the DAN, VAN, SMN, and DMN (q < 0.05 false discovery rate [FDR]-corrected). Patients with more severe spatial neglect exhibited lower SyS and vice versa. Conclusion: The loss of segregation in multiple and specific networks provides a functional framework for the deficits in spatial and nonspatial attention and motor/exploratory ability observed in neglect patients.

Electrophysiological correlates of stimulus-driven reorienting deficits after interference with right parietal cortex during a spatial attention task: a TMS-EEG study.

TMS interference over right intraparietal sulcus (IPS) causally disrupts behaviorally and EEG rhythmic correlates of endogenous spatial orienting before visual target presentation [Capotosto, P., Babiloni, C., Romani, G. L., & Corbetta, M. Differential contribution of right and left parietal cortex to the control of spatial attention: A simultaneous EEG-rTMS study. Cerebral Cortex, 22, 446-454, 2012; Capotosto, P., Babiloni, C., Romani, G. L., & Corbetta, M. Fronto-parietal cortex controls spatial attention through modulation of anticipatory alpha rhythms. Journal of Neuroscience, 29, 5863-5872, 2009]. Here we combine data from our previous studies to examine whether right parietal TMS during spatial orienting also impairs stimulus-driven reorienting or the ability to efficiently process unattended stimuli, that is, stimuli outside the current focus of attention. Healthy volunteers (n = 24) performed a Posner spatial cueing task while their EEG activity was being monitored. Repetitive TMS (rTMS) was applied for 150 msec simultaneously to the presentation of a central arrow directing spatial attention to the location of an upcoming visual target. Right IPS-rTMS impaired target detection, especially for stimuli presented at unattended locations; it also caused a modulation of the amplitude of parieto-occipital positive ERPs peaking at about 480 msec (P3) post-target. The P3 significantly decreased for unattended targets and significantly increased for attended targets after right IPS-rTMS as compared with sham stimulation. Similar effects were obtained for left IPS stimulation albeit in a smaller group of volunteers. We conclude that disruption of anticipatory processes in right IPS has prolonged effects that persist during target processing. The P3 decrement may reflect interference with postdecision processes that are part of stimulus-driven reorienting. Right IPS is a node of functional interaction between endogenous spatial orienting and stimulus-driven reorienting processes in human vision.

Exploring brain activity for positive and negative emotions by means of EEG microstates.

Microstate analysis applied to electroencephalographic signals (EEG) allows both temporal and spatial imaging exploration and represents the activity across the scalp. Despite its potential usefulness in understanding brain activity during a specific task, it has been mostly exploited at rest. We extracted EEG microstates during the presentation of emotional expressions, presented either unilaterally (a face in one visual hemifield) or bilaterally (two faces, one in each hemifield). Results revealed four specific microstate's topographies: (i) M1 involves the temporal areas, mainly in the right hemisphere, with a higher occurrence for stimuli presented in the left than in the right visual field; (ii) M2 is localized in the left temporal cortex, with higher occurrence and coverage for unilateral than bilateral presentations; (iii) M3, with a bilateral temporo-parietal localization, shows higher coverage for bilateral than unilateral presentation; (iv) M4, mainly localized in the right fronto-parietal areas and possibly representing the hemispheric specialization for the peculiar stimulus category, shows higher occurrence and coverage for unilateral stimuli presented in the left than in the right visual field. These results suggest that microstate analysis is a valid tool to explore the cerebral response to emotions and can add new insights on the cerebral functioning, with respect to other EEG markers.

rTMS affects EEG microstates dynamic during evoked activity.

Electrophysiological (EEG) correlates both at time (i.e., event-related potentials, ERP) and frequency (i.e., event-related desynchronization, ERD) domains have been shown to be modulated by external magnetic interference. Parallel studies reported a similar interference also for the EEG microstate at rest and in the period that anticipates a task. Here we investigated whether such interference was prolonged during the evoked activity in the framework of the semantic decision task. To this aim, rTMS was delivered over a core region of both the Default mode network and the language network (i.e., left angular gyrus, AG), previously associated to the current task, and as active control we stimulated the left IPS. When subjects received a non-active stimulation (i.e., Sham), in the period that follows the target onset (i.e., 2 sec after the rTMS) we found an interesting alternation of two dominant microstates (MS1, MS3), previously associated to the phonological network and the Cingulo-Opercular Network (CON), respectively. This dynamic was not altered when TMS was delivered over the left IPS. On the contrary, rTMS over left AG selectively suppressed the phonological-related microstate. These findings provide the first causal evidence of region specificity of the EEG microstates topography during the evoked activity corroborating the idea of a crucial role of AG in the semantic memory. Moreover, the present results might provide insight for understanding the neurophysiological correlates of language disorders e.g., aphasia as well as for planning non-invasive brain stimulation protocols for the rehabilitation.

Directed Flow of Beta Band Communication During Reorienting of Attention Within the Dorsal Attention Network.

Background: The endogenous allocation of spatial attention to selected environmental stimuli is controlled by prefrontal (frontal eye fields [FEFs]) and parietal (superior parietal lobe [SPL] and intraparietal sulcus [IPS]) regions belonging to the dorsal attention network (DAN) with a subdivision in subsystems devoted to reorienting (or shifting) of attention between locations (SPL) or maintaining attention at contralateral versus ipsilateral locations (ventral IPS [vIPS]). Although previous studies suggested a leading role of prefrontal regions over parietal sites in orienting attention, the spectral signature of communication flow within the DAN for different attention processes is still debated. Methods: We used the directed transfer function (DTF) on magnetoencephalography (MEG) data to examine the causal interaction between prefrontal and parietal regions of the DAN when subjects shifted versus maintained attention to a stream of cued visual stimuli. Results: In the beta band, we found that shift versus stay cues induced stronger connectivity (DTF values) from right FEF to right SPL, in the early phase of reorienting. Conversely, when considering stay versus shift cues, an increase of DTF values and stronger directionality was observed between bilateral vIPS and from right vIPS to FEF. Similar analyses carried out in theta, alpha, and gamma showed no significant frontoparietal increases of DTF for shift versus stay cues, whereas the stay-related increase of DTF observed in beta between ventral parietal areas was preserved in the alpha band. Conclusions: These findings suggest that control processes in DAN regions (in particular between FEF and SPL) can be associated to a beta frequency channel during shift of attention. Impact statement In the present study, we compared the reorienting response to novel stimuli with respect to maintaining response. Results provided new insights into understanding the neural mechanisms of control attention processes by identifying the frequency-specific causal interactions between frontal and parietal regions belonging to the dorsal attention network supporting spatial reorienting response.

Automatic coding of environmental distance for walking-related locomotion in the foot-related sensory-motor system: A TMS study on macro-affordances.

We have recently described a facilitation effect for the execution of a walking-related action in response to distant objects/locations in the extrapersonal space. Based on the parallelism with the well-known effect of "micro-affordance", observed during the execution of functionally appropriate hand-related actions towards manipulable objects, we have referred to this effect in terms of "macro-affordance". Here we used transcranical magnetic stimulation (TMS) to investigate whether a foot-related region located in the human dorsal precuneate cortex plays a causal role in the generation and maintenance of such behavioral effect. This question was addressed by comparing the magnitude of the facilitation effect during an incidental go/no-go task, i.e. advantage for walking-related actions to pictures framing an environment from a far vs. near distance, during three different TMS conditions. The three TMS conditions were collected in all subjects in a randomized order and included stimulation of: i. a foot-related region in the anterior precuneus, ii. a control region in the middle intraparietal sulcus (mIPS), and iii. a sham condition. Enrollment in the TMS protocol was based on analysis of individual performance during a preliminary session conducted using a sham stimulation. TMS was administered at a low frequency range before the beginning of each condition. The results showed that stimulation of the foot-related region in the anterior precuneus produced a significant reduction of the walking-related facilitation effect as compared to both stimulation of the active-control region and the non-active sham stimulation. These findings suggest that the foot-related sensory-motor system directly participates in the process of extraction of the spatial features (i.e. distance) from an environmental scene that are useful for locomotion. More in general, these findings support an automatic coding of environmental affordance or "macro-affordances" in the walking-related sensory-motor system.

Cortical Hyper-Connectivity in a Stroke Patient with Rotated Drawing.

In the present case report, we investigated the cortical networks of a patient (DDA) affected by right parietal stroke who showed a constructional phenomenon, in which when coping and recalling from memory a complex figure, the model was reproduced rotated of 90° along the vertical axis. Previous studies suggested that rotation on copy is associated with visuospatial impairments and abnormalities in parietal cortex, whereas rotation on recall might be related to executive deficits and dysfunction of frontal regions. Here, we computed the DDA's resting-state functional connectivity (FC) derived from cortical regions of the dorsal attention (DAN) and the frontal portion of the executive-control network (fECN), which are involved in the control of visuospatial attention and multiple executive functions, respectively. We observed that, as compared to a control group of right stroke patients without drawing rotation, DDA exhibited selective increased FC of the DAN and fECN, but not of task-irrelevant language network, within the undamaged hemisphere. These patterns might reflect a pathological communication in such networks leading to impaired attentional and executive operations required to reproduce the model in the correct orientation. Notably, such enhancement of FC was not detected in a patient with a comparable neuropsychological profile as DDA, yet without rotated drawing, suggesting that network-specific modulations in DDA might be ascribed to the constructional phenomenon of rotated drawing.

Frontoparietal cortex controls spatial attention through modulation of anticipatory alpha rhythms.

A dorsal frontoparietal network, including regions in intraparietal sulcus (IPS) and frontal eye field (FEF), has been hypothesized to control the allocation of spatial attention to environmental stimuli. One putative mechanism of control is the desynchronization of electroencephalography (EEG) alpha rhythms (approximately 8-12 Hz) in visual cortex in anticipation of a visual target. We show that brief interference by repetitive transcranial magnetic stimulation (rTMS) with preparatory activity in right IPS or right FEF while subjects attend to a spatial location impairs identification of target visual stimuli approximately 2 s later. This behavioral effect is associated with the disruption of anticipatory (prestimulus) alpha desynchronization and its spatially selective topography in parieto-occipital cortex. Finally, the disruption of anticipatory alpha rhythms in occipital cortex after right IPS- or right FEF-rTMS correlates with deficits of visual identification. These results support the causal role of the dorsal frontoparietal network in the control of visuospatial attention, and suggest that this is partly exerted through the synchronization of occipital visual neurons.