Gray Matter Volume in Different Cortical Structures Dissociably Relates to Individual Differences in Capacity and Precision of Visual Working Memory.

Visual working memory (VWM) refers to our ability to selectively maintain visual information in a mental representation. While cognitive limits of VWM greatly influence a variety of mental operations, it remains controversial whether the quantity or quality of representations in mind constrains VWM. Here, we examined behavior-to-brain anatomical relations as well as brain activity to brain anatomy associations with a "neural" marker specific to the retention interval of VWM. Our results consistently indicated that individuals who maintained a larger number of items in VWM tended to have a larger gray matter (GM) volume in their left lateral occipital region. In contrast, individuals with a superior ability to retain with high precision tended to have a larger GM volume in their right parietal lobe. These results indicate that individual differences in quantity and quality of VWM may be associated with regional GM volumes in a dissociable manner, indicating willful integration of information in VWM may recruit separable cortical subsystems.

Functional connectivity between prefrontal and parietal cortex drives visuo-spatial attention shifts.

It is well established that the frontal eye-fields (FEF) in the dorsal attention network (DAN) guide top-down selective attention. In addition, converging evidence implies a causal role for the FEF in attention shifting, which is also known to recruit the ventral attention network (VAN) and fronto-striatal regions. To investigate the causal influence of the FEF as (part of) a central hub between these networks, we applied thetaburst transcranial magnetic stimulation (TBS) off-line, combined with functional magnetic resonance (fMRI) during a cued visuo-spatial attention shifting paradigm. We found that TBS over the right FEF impaired performance on a visual discrimination task in both hemifields following attention shifts, while only left hemifield performance was affected when participants were cued to maintain the focus of attention. These effects recovered ca. 20min post stimulation. Furthermore, particularly following attention shifts, TBS suppressed the neural signal in bilateral FEF, right inferior and superior parietal lobule (IPL/SPL) and bilateral supramarginal gyri (SMG). Immediately post stimulation, functional connectivity was impaired between right FEF and right SMG as well as right putamen. Importantly, the extent of decreased connectivity between right FEF and right SMG correlated with behavioural impairment following attention shifts. The main finding of this study demonstrates that influences from right FEF on SMG in the ventral attention network causally underly attention shifts, presumably by enabling disengagement from the current focus of attention.

Effective Connectivity from Early Visual Cortex to Posterior Occipitotemporal Face Areas Supports Face Selectivity and Predicts Developmental Prosopagnosia.

Face processing is mediated by interactions between functional areas in the occipital and temporal lobe, and the fusiform face area (FFA) and anterior temporal lobe play key roles in the recognition of facial identity. Individuals with developmental prosopagnosia (DP), a lifelong face recognition impairment, have been shown to have structural and functional neuronal alterations in these areas. The present study investigated how face selectivity is generated in participants with normal face processing, and how functional abnormalities associated with DP, arise as a function of network connectivity. Using functional magnetic resonance imaging and dynamic causal modeling, we examined effective connectivity in normal participants by assessing network models that include early visual cortex (EVC) and face-selective areas and then investigated the integrity of this connectivity in participants with DP. Results showed that a feedforward architecture from EVC to the occipital face area, EVC to FFA, and EVC to posterior superior temporal sulcus (pSTS) best explained how face selectivity arises in both controls and participants with DP. In this architecture, the DP group showed reduced connection strengths on feedforward connections carrying face information from EVC to FFA and EVC to pSTS. These altered network dynamics in DP contribute to the diminished face selectivity in the posterior occipitotemporal areas affected in DP. These findings suggest a novel view on the relevance of feedforward projection from EVC to posterior occipitotemporal face areas in generating cortical face selectivity and differences in face recognition ability. SIGNIFICANCE STATEMENT: Areas of the human brain showing enhanced activation to faces compared to other objects or places have been extensively studied. However, the factors leading to this face selectively have remained mostly unknown. We show that effective connectivity from early visual cortex to posterior occipitotemporal face areas gives rise to face selectivity. Furthermore, people with developmental prosopagnosia, a lifelong face recognition impairment, have reduced face selectivity in the posterior occipitotemporal face areas and left anterior temporal lobe. We show that this reduced face selectivity can be predicted by effective connectivity from early visual cortex to posterior occipitotemporal face areas. This study presents the first network-based account of how face selectivity arises in the human brain.

Causal evidence that intrinsic beta-frequency is relevant for enhanced signal propagation in the motor system as shown through rhythmic TMS.

Correlative evidence provides support for the idea that brain oscillations underpin neural computations. Recent work using rhythmic stimulation techniques in humans provide causal evidence but the interactions of these external signals with intrinsic rhythmicity remain unclear. Here, we show that sensorimotor cortex follows externally applied rhythmic TMS (rTMS) stimulation in the beta-band but that the elicited responses are strongest at the intrinsic individual beta peak frequency. While these entrainment effects are of short duration, even subthreshold rTMS pulses propagate through the network and elicit significant cortico-spinal coupling, particularly when stimulated at the individual beta-frequency. Our results show that externally enforced rhythmicity interacts with intrinsic brain rhythms such that the individual peak frequency determines the effect of rTMS. The observed downstream spinal effect at the resonance frequency provides evidence for the causal role of brain rhythms for signal propagation.

Local but not long-range microstructural differences of the ventral temporal cortex in developmental prosopagnosia.

Individuals with developmental prosopagnosia (DP) experience face recognition impairments despite normal intellect and low-level vision and no history of brain damage. Prior studies using diffusion tensor imaging in small samples of subjects with DP (n=6 or n=8) offer conflicting views on the neurobiological bases for DP, with one suggesting white matter differences in two major long-range tracts running through the temporal cortex, and another suggesting white matter differences confined to fibers local to ventral temporal face-specific functional regions of interest (fROIs) in the fusiform gyrus. Here, we address these inconsistent findings using a comprehensive set of analyzes in a sample of DP subjects larger than both prior studies combined (n=16). While we found no microstructural differences in long-range tracts between DP and age-matched control participants, we found differences local to face-specific fROIs, and relationships between these microstructural measures with face recognition ability. We conclude that subtle differences in local rather than long-range tracts in the ventral temporal lobe are more likely associated with developmental prosopagnosia.

The perception of touch and the ventral somatosensory pathway.

In humans, touching the skin is known to activate, among others, the contralateral primary somatosensory cortex on the postcentral gyrus together with the bilateral parietal operculum (i.e. the anatomical site of the secondary somatosensory cortex). But which brain regions beyond the postcentral gyrus specifically contribute to the perception of touch remains speculative. In this study we collected structural magnetic resonance imaging scans and neurological examination reports of patients with brain injuries or stroke in the left or right hemisphere, but not in the postcentral gyrus as the entry site of cortical somatosensory processing. Using voxel-based lesion-symptom mapping, we compared patients with impaired touch perception (i.e. hypoaesthesia) to patients without such touch impairments. Patients with hypoaesthesia as compared to control patients differed in one single brain cluster comprising the contralateral parietal operculum together with the anterior and posterior insular cortex, the putamen, as well as subcortical white matter connections reaching ventrally towards prefrontal structures. This finding confirms previous speculations on the 'ventral pathway of somatosensory perception' and causally links these brain structures to the perception of touch.

fMRI correlates of object-based attentional facilitation vs. suppression of irrelevant stimuli, dependent on global grouping and endogenous cueing.

Theories of object-based attention often make two assumptions: that attentional resources are facilitatory, and that they spread automatically within grouped objects. Consistent with this, ignored visual stimuli can be easier to process, or more distracting, when perceptually grouped with an attended target stimulus. But in past studies, the ignored stimuli often shared potentially relevant features or locations with the target. In this fMRI study, we measured the effects of attention and grouping on Blood Oxygenation Level Dependent (BOLD) responses in the human brain to entirely task-irrelevant events. Two checkerboards were displayed each in opposite hemifields, while participants responded to check-size changes in one pre-cued hemifield, which varied between blocks. Grouping (or segmentation) between hemifields was manipulated between blocks, using common (vs. distinct) motion cues. Task-irrelevant transient events were introduced by randomly changing the color of either checkerboard, attended or ignored, at unpredictable intervals. The above assumptions predict heightened BOLD signals for irrelevant events in attended vs. ignored hemifields for ungrouped contexts, but less such attentional modulation under grouping, due to automatic spreading of facilitation across hemifields. We found the opposite pattern, in primary visual cortex. For ungrouped stimuli, BOLD signals associated with task-irrelevant changes were lower, not higher, in the attended vs. ignored hemifield; furthermore, attentional modulation was not reduced but actually inverted under grouping, with higher signals for events in the attended vs. ignored hemifield. These results challenge two popular assumptions underlying object-based attention. We consider a broader biased-competition framework: task-irrelevant stimuli are suppressed according to how strongly they compete with task-relevant stimuli, with intensified competition when the irrelevant features or locations comprise the same object.

Within-hemifield posture changes affect tactile-visual exogenous spatial cueing without spatial precision, especially in the dark.

We investigated the effects of seen and unseen within-hemifield posture changes on crossmodal visual-tactile links in covert spatial attention. In all experiments, a spatially nonpredictive tactile cue was presented to the left or the right hand, with the two hands placed symmetrically across the midline. Shortly after a tactile cue, a visual target appeared at one of two eccentricities within either of the hemifields. For half of the trial blocks, the hands were aligned with the inner visual target locations, and for the remainder, the hands were aligned with the outer target locations. In Experiments 1 and 2, the inner and outer eccentricities were 17.5º and 52.5º, respectively. In Experiment 1, the arms were completely covered, and visual up-down judgments were better when on the same side as the preceding tactile cue. Cueing effects were not significantly affected by hand or target alignment. In Experiment 2, the arms were in view, and now some target responses were affected by cue alignment: Cueing for outer targets was only significant when the hands were aligned with them. In Experiment 3, we tested whether any unseen posture changes could alter the cueing effects, by widely separating the inner and outer target eccentricities (now 10º and 86º). In this case, hand alignment did affect some of the cueing effects: Cueing for outer targets was now only significant when the hands were in the outer position. Although these results confirm that proprioception can, in some cases, influence tactile-visual links in exogenous spatial attention, they also show that spatial precision is severely limited, especially when posture is unseen.

Direct evidence for attention-dependent influences of the frontal eye-fields on feature-responsive visual cortex.

Voluntary selective attention can prioritize different features in a visual scene. The frontal eye-fields (FEF) are one potential source of such feature-specific top-down signals, but causal evidence for influences on visual cortex (as was shown for "spatial" attention) has remained elusive. Here, we show that transcranial magnetic stimulation (TMS) applied to right FEF increased the blood oxygen level-dependent (BOLD) signals in visual areas processing "target feature" but not in "distracter feature"-processing regions. TMS-induced BOLD signals increase in motion-responsive visual cortex (MT+) when motion was attended in a display with moving dots superimposed on face stimuli, but in face-responsive fusiform area (FFA) when faces were attended to. These TMS effects on BOLD signal in both regions were negatively related to performance (on the motion task), supporting the behavioral relevance of this pathway. Our findings provide new causal evidence for the human FEF in the control of nonspatial "feature"-based attention, mediated by dynamic influences on feature-specific visual cortex that vary with the currently attended property.

Spatial attention, precision, and Bayesian inference: a study of saccadic response speed.

Inferring the environment's statistical structure and adapting behavior accordingly is a fundamental modus operandi of the brain. A simple form of this faculty based on spatial attentional orienting can be studied with Posner's location-cueing paradigm in which a cue indicates the target location with a known probability. The present study focuses on a more complex version of this task, where probabilistic context (percentage of cue validity) changes unpredictably over time, thereby creating a volatile environment. Saccadic response speed (RS) was recorded in 15 subjects and used to estimate subject-specific parameters of a Bayesian learning scheme modeling the subjects' trial-by-trial updates of beliefs. Different response models-specifying how computational states translate into observable behavior-were compared using Bayesian model selection. Saccadic RS was most plausibly explained as a function of the precision of the belief about the causes of sensory input. This finding is in accordance with current Bayesian theories of brain function, and specifically with the proposal that spatial attention is mediated by a precision-dependent gain modulation of sensory input. Our results provide empirical support for precision-dependent changes in beliefs about saccade target locations and motivate future neuroimaging and neuropharmacological studies of how Bayesian inference may determine spatial attention.

Sight and sound out of synch: fragmentation and renormalisation of audiovisual integration and subjective timing.

The sight and sound of a person speaking or a ball bouncing may seem simultaneous, but their corresponding neural signals are spread out over time as they arrive at different multisensory brain sites. How subjective timing relates to such neural timing remains a fundamental neuroscientific and philosophical puzzle. A dominant assumption is that temporal coherence is achieved by sensory resynchronisation or recalibration across asynchronous brain events. This assumption is easily confirmed by estimating subjective audiovisual timing for groups of subjects, which is on average similar across different measures and stimuli, and approximately veridical. But few studies have examined normal and pathological individual differences in such measures. Case PH, with lesions in pons and basal ganglia, hears people speak before seeing their lips move. Temporal order judgements (TOJs) confirmed this: voices had to lag lip-movements (by ∼200 msec) to seem synchronous to PH. Curiously, voices had to lead lips (also by ∼200 msec) to maximise the McGurk illusion (a measure of audiovisual speech integration). On average across these measures, PH's timing was therefore still veridical. Age-matched control participants showed similar discrepancies. Indeed, normal individual differences in TOJ and McGurk timing correlated negatively: subjects needing an auditory lag for subjective simultaneity needed an auditory lead for maximal McGurk, and vice versa. This generalised to the Stream-Bounce illusion. Such surprising antagonism seems opposed to good sensory resynchronisation, yet average timing across tasks was still near-veridical. Our findings reveal remarkable disunity of audiovisual timing within and between subjects. To explain this we propose that the timing of audiovisual signals within different brain mechanisms is perceived relative to the average timing across mechanisms. Such renormalisation fully explains the curious antagonistic relationship between disparate timing estimates in PH and healthy participants, and how they can still perceive the timing of external events correctly, on average.

The duration of a co-occurring sound modulates visual detection performance in humans.

BACKGROUND: The duration of sounds can affect the perceived duration of co-occurring visual stimuli. However, it is unclear whether this is limited to amodal processes of duration perception or affects other non-temporal qualities of visual perception. METHODOLOGY/PRINCIPAL FINDINGS: Here, we tested the hypothesis that visual sensitivity--rather than only the perceived duration of visual stimuli--can be affected by the duration of co-occurring sounds. We found that visual detection sensitivity (d') for unimodal stimuli was higher for stimuli of longer duration. Crucially, in a cross-modal condition, we replicated previous unimodal findings, observing that visual sensitivity was shaped by the duration of co-occurring sounds. When short visual stimuli (∼24 ms) were accompanied by sounds of matching duration, visual sensitivity was decreased relative to the unimodal visual condition. However, when the same visual stimuli were accompanied by longer auditory stimuli (∼60-96 ms), visual sensitivity was increased relative to the performance for ∼24 ms auditory stimuli. Across participants, this sensitivity enhancement was observed within a critical time window of ∼60-96 ms. Moreover, the amplitude of this effect correlated with visual sensitivity enhancement found for longer lasting visual stimuli across participants. CONCLUSIONS/SIGNIFICANCE: Our findings show that the duration of co-occurring sounds affects visual perception; it changes visual sensitivity in a similar way as altering the (actual) duration of the visual stimuli does.

Partial recovery of visual extinction by pavlovian conditioning in a patient with hemispatial neglect.

Patients with parietal lesions and unilateral spatial neglect (USN) are unable to detect or respond to information in the contralesional side of space. However, some residual sensory processing may still occur and overcome inattention symptoms when contralesional stimuli are perceptually or biologically salient, as shown for emotional faces or voices. These effects have been attributed to enhanced neural responses of sensory regions to emotional stimuli, presumably driven by feedback signals from limbic regions such as the amygdala. However, because emotional faces and voices also differ from neutral stimuli in terms of physical features, the affective nature of these effects still remains to be confirmed. Here we report data from a right parietal patient in whom left visual extinction was reduced for contralesional visual stimuli following pavlovian aversive conditioning, relative to the same stimulus before conditioning, and relative to similar but non-conditioned stimuli. This reduction of visual extinction was thus mediated by the emotional meaning of stimuli acquired through implicit learning. Functional magnetic resonance imaging also showed that conditioned visual stimuli elicited greater activation in right visual cortex, relative to the non-conditioned stimuli, together with differential activations in amygdala. These results support the hypothesis that emotional appraisal, not only the processing of perceptual features, may partly restore attention to salient information in contralesional space. These findings open new perspectives to improve rehabilitation strategies in neglect, based on affective and motivational signals.

The boundary conditions of priming of visual search: from passive viewing through task-relevant working memory load.

Priming of visual search has a dominating effect upon attentional shifts and is thought to play a decisive role in visual stability. Despite this importance, the nature of the memory underlying priming remains controversial. To understand more fully the necessary conditions for priming, we contrasted passive versus active viewing of visual search arrays. There was no priming from passive viewing of search arrays, while it was strong for active search of the same displays. Displays requiring no search resulted in no priming, again showing that search is needed for priming to occur. Finally, we introduced working memory load during visual search in an effort to disrupt priming. The memorized items had either the same colors as or different colors from the visual search items. Retaining items in working memory inhibited priming of the working memory task-relevant colors, while little interference was observed for unrelated colors. The picture that emerges of priming is that it requires active attentional processing of the search items in addition to the operation of visual working memory, where the task relevance of the working memory load plays a key role.

Visual and audiovisual effects of isochronous timing on visual perception and brain activity.

Understanding how the brain extracts and combines temporal structure (rhythm) information from events presented to different senses remains unresolved. Many neuroimaging beat perception studies have focused on the auditory domain and show the presence of a highly regular beat (isochrony) in "auditory" stimulus streams enhances neural responses in a distributed brain network and affects perceptual performance. Here, we acquired functional magnetic resonance imaging (fMRI) measurements of brain activity while healthy human participants performed a visual task on isochronous versus randomly timed "visual" streams, with or without concurrent task-irrelevant sounds. We found that visual detection of higher intensity oddball targets was better for isochronous than randomly timed streams, extending previous auditory findings to vision. The impact of isochrony on visual target sensitivity correlated positively with fMRI signal changes not only in visual cortex but also in auditory sensory cortex during audiovisual presentations. Visual isochrony activated a similar timing-related brain network to that previously found primarily in auditory beat perception work. Finally, activity in multisensory left posterior superior temporal sulcus increased specifically during concurrent isochronous audiovisual presentations. These results indicate that regular isochronous timing can modulate visual processing and this can also involve multisensory audiovisual brain mechanisms.

Temporal coupling due to illusory movements in bimanual actions: evidence from anosognosia for hemiplegia.

In anosognosia for hemiplegia, patients may claim having performed willed actions with the paralyzed limb despite unambiguous evidence to the contrary. Does this false belief of having moved reflect the functioning of the same mechanisms that govern normal motor performance? Here, we examined whether anosognosics show the same temporal constraints known to exist during bimanual movements in healthy subjects. In these paradigms, when participants simultaneously reach for two targets of different difficulties, the motor programs of one hand affect the execution of the other. In detail, the movement time of the hand going to an easy target (i.e., near and large), while the other is going to a difficult target (i.e., far and small), is slowed with respect to unimanual movements (temporal coupling effect). One right-brain-damaged patient with left hemiplegia and anosognosia, six right-brain-damaged patients with left hemiplegia without anosognosia, and twenty healthy subjects were administered such a bimanual task. We recorded the movement times for easy and difficult targets, both in unimanual (one target) and bimanual (two targets) conditions. We found that, as healthy subjects, the anosognosic patient showed coupling effect. In bimanual asymmetric conditions (when one hand went to the easy target and the other went to the difficult target), the movement time of the non-paralyzed hand going to the easy target was slowed by the 'pretended' movement of the paralyzed hand going to the difficult target. This effect was not present in patients without anosognosia. We concluded that in anosognosic patients, the illusory movements of the paralyzed hand impose to the non-paralyzed hand the same motor constraints that emerge during the actual movements. Our data also support the view that coupling relies on central operations (i.e., activation of intention/programming system), rather than on online information from the periphery.

Preserved local but disrupted contextual figure-ground influences in an individual with abnormal function of intermediate visual areas.

Visual perception depends not only on local stimulus features but also on their relationship to the surrounding stimulus context, as evident in both local and contextual influences on figure-ground segmentation. Intermediate visual areas may play a role in such contextual influences, as we tested here by examining LG, a rare case of developmental visual agnosia. LG has no evident abnormality of brain structure and functional neuroimaging showed relatively normal V1 function, but his intermediate visual areas (V2/V3) function abnormally. We found that contextual influences on figure-ground organization were selectively disrupted in LG, while local sources of figure-ground influences were preserved. Effects of object knowledge and familiarity on figure-ground organization were also significantly diminished. Our results suggest that the mechanisms mediating contextual and familiarity influences on figure-ground organization are dissociable from those mediating local influences on figure-ground assignment. The disruption of contextual processing in intermediate visual areas may play a role in the substantial object recognition difficulties experienced by LG.

Deconstructing the architecture of dorsal and ventral attention systems with dynamic causal modeling.

Attentional orientation to a spatial cue and reorientation-after invalid cueing-are mediated by two distinct networks in the human brain. A bilateral dorsal frontoparietal network, comprising the intraparietal sulcus (IPS) and the frontal eye fields (FEF), controls the voluntary deployment of attention and may modulate visual cortex in preparation for upcoming stimulation. In contrast, reorienting attention to invalidly cued targets engages a right-lateralized ventral frontoparietal network comprising the temporoparietal junction (TPJ) and ventral frontal cortex. The present fMRI study investigated the functional architecture of these two attentional systems by characterizing effective connectivity during lateralized orienting and reorienting of attention, respectively. Subjects performed a modified version of Posner's location-cueing paradigm. Dynamic causal modeling (DCM) of regional responses in the dorsal and ventral network, identified in a conventional (SPM) whole-brain analysis, was used to compare different functional architectures. Bayesian model selection showed that top-down connections from left and right IPS to left and right visual cortex, respectively, were modulated by the direction of attention. Moreover, model evidence was highest for a model with directed influences from bilateral IPS to FEF, and reciprocal coupling between right and left FEF. Invalid cueing enhanced forward connections from visual areas to right TPJ, and directed influences from right TPJ to right IPS and IFG (inferior frontal gyrus). These findings shed further light on the functional organization of the dorsal and ventral attentional network and support a context-sensitive lateralization in the top-down (backward) mediation of attentional orienting and the bottom-up (forward) effects of invalid cueing.

fMRI correlates of subjective reversals in ambiguous structure-from-motion.

We used fMRI to examine the neural correlates of subjective reversals for bistable structure-from-motion. We compared transparent random-dot kinematograms depicting either a cylinder rotating in depth or two flat surfaces translating in opposite directions at apparently different depths. For both such stimuli, the motion of dots on the different apparent depth planes typically appears to reverse direction periodically on prolonged viewing. Yet for cylindrical but not flat stimuli, such subjective reversals also coincide with apparent reversal of 3D rotation direction. We hypothesized that the lateral occipital complex (region LOC), sensitive to 3D form, might show greater event-related activity for subjective reversals of cylindrical than flat stimuli; conversely, motion-sensitive hMT+/V5 should respond in common to subjective reversals for either type of stimuli, as both are perceived as changes in planar motion. We obtained an event-related measure of neural activity associated with subjective reversals after first factoring out block-related differences between cylindrical versus flat stimuli (and thereby the associated low-level blocked stimulus differences). In support of our hypothesis, only the cylindrical stimuli produced reversal-related activity in contralateral human LOC. In contrast, the hMT+/V5 complex was activated alike by subjective reversals for both cylindrical and flat stimuli. Intriguingly, V1 also showed (contralateral) specificity for rotational reversals, suggesting a possible feedback influence from LOC. These results reveal specific neural correlates for subjective switches of 3D rotation versus translation, as distinct from subjective reversals in general.

Rapid enhancement of touch from non-informative vision of the hand.

Processing in one sensory modality may modulate processing in another. Here we investigate how simply viewing the hand can influence the sense of touch. Previous studies showed that non-informative vision of the hand enhances tactile acuity, relative to viewing an object at the same location. However, it remains unclear whether this Visual Enhancement of Touch (VET) involves a phasic enhancement of tactile processing circuits triggered by the visual event of seeing the hand, or more prolonged, tonic neuroplastic changes, such as recruitment of additional cortical areas for tactile processing. We recorded somatosensory evoked potentials (SEPs) evoked by electrical stimulation of the right middle finger, both before and shortly after viewing either the right hand, or a neutral object presented via a mirror. Crucially, and unlike prior studies, our visual exposures were unpredictable and brief, in addition to being non-informative about touch. Viewing the hand, as opposed to viewing an object, enhanced tactile spatial discrimination measured using grating orientation judgements, and also the P50 SEP component, which has been linked to early somatosensory cortical processing. This was a trial-specific, phasic effect, occurring within a few seconds of each visual onset, rather than an accumulating, tonic effect. Thus, somatosensory cortical modulation can be triggered even by a brief, non-informative glimpse of one's hand. Such rapid multisensory modulation reveals novel aspects of the specialised brain systems for functionally representing the body.