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Probing brain functions by brain stimulation while simultaneously recording brain activity allows addressing major issues in cognitive neuroscience. We review recent studies where electroencephalography (EEG) has been combined with transcranial magnetic stimulation (TMS) in order to investigate possible neuronal substrates of visual perception and attention. TMS-EEG has been used to study both pre-stimulus brain activity patterns that affect upcoming perception, and also the stimulus-evoked and task-related inter-regional interactions within the extended visual-attentional network from which attention and perception emerge. Local processes in visual areas have been probed by directly stimulating occipital cortex while monitoring EEG activity and perception. Interactions within the attention network have been probed by concurrently stimulating frontal or parietal areas. The use of tasks manipulating implicit and explicit memory has revealed in addition a role for attentional processes in memory. Taken together, these studies helped to reveal that visual selection relies on spontaneous intrinsic activity in visual cortex prior to the incoming stimulus, their control by attention, and post-stimulus processes incorporating a re-entrant bias from frontal and parietal areas that depends on the task., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Although oscillatory activity in the alpha band was traditionally associated with lack of alertness, more recent work has linked it to specific cognitive functions, including visual attention. The emerging method of rhythmic transcranial magnetic stimulation (TMS) allows causal interventional tests for the online impact on performance of TMS administered in short bursts at a particular frequency. TMS bursts at 10 Hz have recently been shown to have an impact on spatial visual attention, but any role in featural attention remains unclear. Here we used rhythmic TMS at 10 Hz to assess the impact on attending to global or local components of a hierarchical Navon-like stimulus (D. Navon (1977) Forest before trees: The precedence of global features in visual perception. Cognit. Psychol., 9, 353), in a paradigm recently used with TMS at other frequencies (V. Romei, J. Driver, P.G. Schyns & G. Thut. (2011) Rhythmic TMS over parietal cortex links distinct brain frequencies to global versus local visual processing. Curr. Biol., 2, 334-337). In separate groups, left or right posterior parietal sites were stimulated at 10 Hz just before presentation of the hierarchical stimulus. Participants had to identify either the local or global component in separate blocks. Right parietal 10 Hz stimulation (vs. sham) significantly impaired global processing without affecting local processing, while left parietal 10 Hz stimulation vs. sham impaired local processing with a minor trend to enhance global processing. These 10 Hz outcomes differed significantly from stimulation at other frequencies (i.e. 5 or 20 Hz) over the same site in other recent work with the same paradigm. These dissociations confirm differential roles of the two hemispheres in local vs. global processing, and reveal a frequency-specific role for stimulation in the alpha band for regulating feature-based visual attention., (© 2012 The Authors. European Journal of Neuroscience © 2012 Federation of European Neuroscience Societies and Blackwell Publishing Ltd.)

Multisensory interactions are a fundamental feature of brain organization. Principles governing multisensory processing have been established by varying stimulus location, timing and efficacy independently. Determining whether and how such principles operate when stimuli vary dynamically in their perceived distance (as when looming/receding) provides an assay for synergy among the above principles and also means for linking multisensory interactions between rudimentary stimuli with higher-order signals used for communication and motor planning. Human participants indicated movement of looming or receding versus static stimuli that were visual, auditory, or multisensory combinations while 160-channel EEG was recorded. Multivariate EEG analyses and distributed source estimations were performed. Nonlinear interactions between looming signals were observed at early poststimulus latencies (∼75 ms) in analyses of voltage waveforms, global field power, and source estimations. These looming-specific interactions positively correlated with reaction time facilitation, providing direct links between neural and performance metrics of multisensory integration. Statistical analyses of source estimations identified looming-specific interactions within the right claustrum/insula extending inferiorly into the amygdala and also within the bilateral cuneus extending into the inferior and lateral occipital cortices. Multisensory effects common to all conditions, regardless of perceived distance and congruity, followed (∼115 ms) and manifested as faster transition between temporally stable brain networks (vs summed responses to unisensory conditions). We demonstrate the early-latency, synergistic interplay between existing principles of multisensory interactions. Such findings change the manner in which to model multisensory interactions at neural and behavioral/perceptual levels. We also provide neurophysiologic backing for the notion that looming signals receive preferential treatment during perception.

The effects of repetitive transcranial magnetic stimulation (rTMS) on cortical excitability are usually inferred from indirect indexes, such as EMG responses. It has now become possible to directly evaluate rTMS impact by means of concurrent EEG recording. The aim of this study was to examine the modulation induced by high frequency rTMS (20 Hz) over left primary motor cortex on the ongoing oscillatory activity. Thirteen subjects underwent two sham and a real rTMS session while acquiring EEG. Event-related desynchronization/synchronization was calculated for the α and β bands. rTMS induced a dose-dependent increase in synchronization in both bands over central and parietal sites. The strongest effect found for the α band outlasted the end of the stimulation. Considering previous studies, our data suggest that α generation may represent an intrinsic induced response and a basic response signature to TMS targeting the human resting motor cortex., (Copyright © 2011 Society for Psychophysiological Research.)

Background: Neuronal elements underlying perception, cognition, and action exhibit distinct oscillatory phenomena, measured in humans by electro- or magnetoencephalography (EEG/MEG). So far, the correlative or causal nature of the link between brain oscillations and functions has remained elusive. A compelling demonstration of causality would primarily generate oscillatory signatures that are known to correlate with particular cognitive functions and then assess the behavioral consequences. Here, we provide the first direct evidence for causal entrainment of brain oscillations by transcranial magnetic stimulation (TMS) using concurrent EEG., Results: We used rhythmic TMS bursts to directly interact with an MEG-identified parietal α-oscillator, activated by attention and linked to perception. With TMS bursts tuned to its preferred α-frequency (α-TMS), we confirmed the three main predictions of entrainment of a natural oscillator: (1) that α-oscillations are induced during α-TMS (reproducing an oscillatory signature of the stimulated parietal cortex), (2) that there is progressive enhancement of this α-activity (synchronizing the targeted, α-generator to the α-TMS train), and (3) that this depends on the pre-TMS phase of the background α-rhythm (entrainment of natural, ongoing α-oscillations). Control conditions testing different TMS burst profiles and TMS-EEG in a phantom head confirmed specificity of α-boosting to the case of synchronization between TMS train and neural oscillator., Conclusions: The periodic electromagnetic force that is generated during rhythmic TMS can cause local entrainment of natural brain oscillations, emulating oscillatory signatures activated by cognitive tasks. This reveals a new mechanism of online TMS action on brain activity and can account for frequency-specific behavioral TMS effects at the level of biologically relevant rhythms., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

The notion of driving brain oscillations by directly stimulating neuronal elements with rhythmic stimulation protocols has become increasingly popular in research on brain rhythms. Induction of brain oscillations in a controlled and functionally meaningful way would likely prove highly beneficial for the study of brain oscillations, and their therapeutic control. We here review conventional and new non-invasive brain stimulation protocols as to their suitability for controlled intervention into human brain oscillations. We focus on one such type of intervention, the direct entrainment of brain oscillations by a periodic external drive. We review highlights of the literature on entraining brain rhythms linked to perception and attention, and point out controversies. Behaviourally, such entrainment seems to alter specific aspects of perception depending on the frequency of stimulation, informing models on the functional role of oscillatory activity. This indicates that human brain oscillations and function may be promoted in a controlled way by focal entrainment, with great potential for probing into brain oscillations and their causal role.

Neural networks underlying visual perception exhibit oscillations at different frequencies (e.g.,). But how these map onto distinct aspects of visual perception remains elusive. Recent electroencephalography data indicate that theta or beta frequencies at parietal sensors increase in amplitude when conscious perception is dominated by global or local features, respectively, of a reversible visual stimulus. But this provides only correlative, noninterventional evidence. Here we show via transcranial magnetic stimulation (TMS) interventions that short rhythmic bursts of right-parietal TMS at theta or beta frequency can causally benefit processing of global or local levels, respectively, for hierarchical visual stimuli, especially in the context of salient incongruent distractors. This double dissociation between theta and beta TMS reveals distinct causal roles for particular frequencies in processing global versus local visual features., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Spatial frequencies in an image influence visual analysis across a distributed, hierarchically organized brain network. Low spatial frequency (LSF) information may rapidly reach high-order areas to allow an initial coarse parsing of the visual scene, which could then be "retroinjected" through feedback into lower level visual areas to guide finer analysis on the basis of high spatial frequency (HSF). To test this "coarse-to-fine" processing scheme and to identify its neural substrates in the human brain, we presented sequences of two spatial-frequency-filtered scenes in rapid succession (LSF followed by HSF or vice versa) during fMRI and ERPs in the same participants. We show that for low-to-high sequences (but not for high-to-low sequences), LSF produces a first increase of activity in prefrontal and temporo-parietal areas, followed by enhanced responses to HSF in primary visual cortex. This pattern is consistent with retroactive influences on low-level areas that process HSF after initial activation of higher order areas by LSF.

Current models of brain organization include multisensory interactions at early processing stages and within low-level, including primary, cortices. Embracing this model with regard to auditory-visual (AV) interactions in humans remains problematic. Controversy surrounds the application of an additive model to the analysis of event-related potentials (ERPs), and conventional ERP analysis methods have yielded discordant latencies of effects and permitted limited neurophysiologic interpretability. While hemodynamic imaging and transcranial magnetic stimulation studies provide general support for the above model, the precise timing, superadditive/subadditive directionality, topographic stability, and sources remain unresolved. We recorded ERPs in humans to attended, but task-irrelevant stimuli that did not require an overt motor response, thereby circumventing paradigmatic caveats. We applied novel ERP signal analysis methods to provide details concerning the likely bases of AV interactions. First, nonlinear interactions occur at 60-95 ms after stimulus and are the consequence of topographic, rather than pure strength, modulations in the ERP. AV stimuli engage distinct configurations of intracranial generators, rather than simply modulating the amplitude of unisensory responses. Second, source estimations (and statistical analyses thereof) identified primary visual, primary auditory, and posterior superior temporal regions as mediating these effects. Finally, scalar values of current densities in all of these regions exhibited functionally coupled, subadditive nonlinear effects, a pattern increasingly consistent with the mounting evidence in nonhuman primates. In these ways, we demonstrate how neurophysiologic bases of multisensory interactions can be noninvasively identified in humans, allowing for a synthesis across imaging methods on the one hand and species on the other.

Transcranial magnetic stimulation (TMS) has developed into a powerful tool for studying human brain physiology and brain-behavior relations. When applied in sessions of repeated stimulation, TMS can lead to changes in neuronal activity/excitability that outlast the stimulation itself. Such aftereffects are at the heart of the offline TMS protocols in cognitive neuroscience and neurotherapeutics. However, whether these aftereffects are of applied interest critically depends on their magnitude and duration, which should fall within an experimentally or clinically useful range without increasing risks and adverse effects. In this short review, we survey combined TMS-EEG studies to characterize the TMS-aftereffects as revealed by EEG to contribute to the characterization of the most effective and promising repetitive TMS-parameters. With one session of conventional repetitive TMS (of fixed pulse frequency), aftereffects were consistently comparable in magnitude to EEG-changes reported after learning or with fatigue, and were short-lived (<70 min). The few studies using recently developed protocols (such as theta burst stimulation) suggest comparable effect-size but longer effect-durations. Based on the reviewed data, it is expected that TMS-efficacy can be further promoted by repeating TMS-sessions, by using EEG-gated TMS to tailor TMS to current neuronal state, or by other, non-conventional TMS-protocols. Newly emerging developments in offline TMS research for cognitive neuroscience and neurotherapeutics are outlined.

The combination of brain stimulation by transcranial magnetic stimulation (TMS) with simultaneous electroencephalographic (EEG) imaging has become feasible due to recent technical developments. The TMS-EEG integration provides real-time information on cortical reactivity and connectivity through the analysis of TMS-evoked potentials (TEPs), and how functional activity links to behavior through the study of TMS-induced modulations thereof. It reveals how these effects vary as a function of neuronal state, differing between individuals and patient groups but also changing rapidly over time during task performance. This review discusses the wide range of possible TMS-EEG applications and what new information may be gained using this technique on the dynamics of brain functions, hierarchical organization, and cortical connectivity, as well as on TMS action per se. An advance in the understanding of these issues is timely and promises to have a substantial impact on many areas of clinical and basic neuroscience.

Repetitive Transcranial Magnetic Stimulation (rTMS) is a non-invasive method for brain stimulation. Group-studies applying rTMS in epilepsy patients aiming to decrease epileptic spike- or seizure-frequency have led to inconsistent results. Here we studied whether therapeutic trains of rTMS have detectable effects on individual spike pattern and/or frequency in patients suffering from focal epilepsy. Five patients with focal epilepsy underwent one session of rTMS online with EEG using a 6 Hz prime/1 Hz rTMS protocol (real and sham). The EEG was recorded continuously throughout the stimulation, and the epileptic spikes recorded immediately before (baseline) and after stimulation (sham and real) were subjected to further analysis. Number of spikes, spike-strength and spike-topography were examined. In two of the five patients, real TMS led to significant changes when compared to baseline and sham (decrease in spike-count in one patient, change in topography of the after-discharge in the other patient). Spike-count and topography remained unchanged the remaining patients. Overall, our results do not indicate a consistent effect of rTMS stimulation on interictal spike discharges, but speak in favor of a rather weak and individually variable immediate effect of rTMS on focal epileptic activity. The individuation of most effective stimulation patterns will be decisive for the future role of rTMS in epilepsies and needs to be determined in larger studies.

Evidence of multisensory interactions within low-level cortices and at early post-stimulus latencies has prompted a paradigm shift in conceptualizations of sensory organization. However, the mechanisms of these interactions and their link to behavior remain largely unknown. One behaviorally salient stimulus is a rapidly approaching (looming) object, which can indicate potential threats. Based on findings from humans and nonhuman primates suggesting there to be selective multisensory (auditory-visual) integration of looming signals, we tested whether looming sounds would selectively modulate the excitability of visual cortex. We combined transcranial magnetic stimulation (TMS) over the occipital pole and psychophysics for "neurometric" and psychometric assays of changes in low-level visual cortex excitability (i.e., phosphene induction) and perception, respectively. Across three experiments we show that structured looming sounds considerably enhance visual cortex excitability relative to other sound categories and white-noise controls. The time course of this effect showed that modulation of visual cortex excitability started to differ between looming and stationary sounds for sound portions of very short duration (80 ms) that were significantly below (by 35 ms) perceptual discrimination threshold. Visual perceptions are thus rapidly and efficiently boosted by sounds through early, preperceptual and stimulus-selective modulation of neuronal excitability within low-level visual cortex.

Procedural skills consist of several components that can be simultaneously acquired. During a motor-learning task we can distinguish between how a "movement" is performed (intrinsic component) and the spatial-related (extrinsic) component of this movement. The intrinsic movement component is thought to be supported by motor loops, including primary motor cortex (M1) as assessed with neuroimaging studies. Here we want to test further whether M1 makes a critical contribution to the movement rather than spatial-related component of skill-learning. To this purpose, we used repetitive Transcranial Magnetic Stimulation (rTMS) and the serial reaction time (SRT) task. Twenty right-handed participants performed the SRT-task starting with their left or right hand. After this learning session, participants switched to the untrained hand by performing original (spatial-related) and mirror-ordered (movement-based) sequences. rTMS was applied to M1 ipsi- or contralateral to the transfer-hand and both sequences were retested. Results revealed rTMS-interference with motor-skill transfer of mirror-ordered but not original sequences, showing that M1 is critically involved in the retrieval/transformation of the intrinsic but not the extrinsic movement coordinates. rTMS-interference in the mirror-condition consisted of both (i) disruption and (ii) release of motor-skill transfer depending on the stimulated hemisphere and on transfer-hand. The pattern of results suggests (i) contralateral (right) M1 involvement in retrieval/transformation of motor information during left-hand reproduction of previously acquired right-hand motor-skills; and (ii) modulatory interactions of inhibitory nature from the dominant (left) to the non-dominant (right) M1 in the same transfer-condition. These results provide further evidence that M1 is essential to intrinsic movement-based skill-learning and novel insight on models of motor-learning and hemispheric specialization, suggesting the involvement of interhemispheric inhibition.

There is renewed interest in the functional role of oscillatory brain activity in specific frequency bands, investigated in humans through electroencephalography (EEG) and magnetoencephalography (MEG) recordings. In parallel, there is a growing body of research on non-invasive direct stimulation of the human brain via repetitive (rhythmic) transcranial magnetic stimulation (TMS), and on those frequencies that have the strongest behavioural impact. There is, therefore, great potential in combining these two lines of research to foster knowledge on brain rhythms, in addition to potential therapeutic applications of rhythmic brain stimulation. Here, we review findings from this rapidly evolving field linking intrinsic brain oscillations to distinct sensory, motor and cognitive operations. The findings emphasize that brain rhythms are causally implicated in cognitive functions.

An object's motion relative to an observer can confer ethologically meaningful information. Approaching or looming stimuli can signal threats/collisions to be avoided or prey to be confronted, whereas receding stimuli can signal successful escape or failed pursuit. Using movement detection and subjective ratings, we investigated the multisensory integration of looming and receding auditory and visual information by humans. While prior research has demonstrated a perceptual bias for unisensory and more recently multisensory looming stimuli, none has investigated whether there is integration of looming signals between modalities. Our findings reveal selective integration of multisensory looming stimuli. Performance was significantly enhanced for looming stimuli over all other multisensory conditions. Contrasts with static multisensory conditions indicate that only multisensory looming stimuli resulted in facilitation beyond that induced by the sheer presence of auditory-visual stimuli. Controlling for variation in physical energy replicated the advantage for multisensory looming stimuli. Finally, only looming stimuli exhibited a negative linear relationship between enhancement indices for detection speed and for subjective ratings. Maximal detection speed was attained when motion perception was already robust under unisensory conditions. The preferential integration of multisensory looming stimuli highlights that complex ethologically salient stimuli likely require synergistic cooperation between existing principles of multisensory integration. A new conceptualization of the neurophysiologic mechanisms mediating real-world multisensory perceptions and action is therefore supported.

Switching from one functional or cognitive operation to another is thought to rely on executive/control processes. The efficacy of these processes may depend on the extent of overlap between neural circuitry mediating the different tasks; more effective task preparation (and by extension smaller switch costs) is achieved when this overlap is small. We investigated the performance costs associated with switching tasks and/or switching sensory modalities. Participants discriminated either the identity or spatial location of objects that were presented either visually or acoustically. Switch costs between tasks were significantly smaller when the sensory modality of the task switched versus when it repeated. This was the case irrespective of whether the pre-trial cue informed participants only of the upcoming task, but not sensory modality (Experiment 1) or whether the pre-trial cue was informative about both the upcoming task and sensory modality (Experiment 2). In addition, in both experiments switch costs between the senses were positively correlated when the sensory modality of the task repeated across trials and not when it switched. The collective evidence supports the independence of control processes mediating task switching and modality switching and also the hypothesis that switch costs reflect competitive interference between neural circuits.

Voluntarily directing visual attention to a cued position in space leads to improved processing of forthcoming visual stimuli at the attended position, and attenuated processing of competing stimuli elsewhere, due to anticipatory tuning of visual cortex activity. In EEG, recent evidence points to a determining role of modulations of posterior alpha-band activity (8-14 Hz) in such anticipatory facilitation (alpha-power decreases) versus inhibition (alpha-power increases). Yet, while such alpha-modulations are a common finding, the direction of modulation varies to a great extent across studies implying dependence on task demands. Here, we reveal opposite modulation of posterior alpha-power with early/initiation versus later/sustained processes of anticipatory attention orienting. Marked alpha-decreases were observed during shifting of attention (initial 700 ms) over occipito-parietal areas processing to-be-attended visual space, while alpha-increases dominated in the subsequent maintenance phase (>700 ms) over occipito-parietal cortex tuned to unattended positions. Notably, the presence of alpha-modulation strongly depended on individual resting alpha-power. Overall, this provides further support to an active facilitative versus inhibitory role of alpha-power decreases and increases and suggests that these attention-related changes are differentially deployed during anticipatory attention orienting to prepare versus maintain the cortex for optimal target processing.

Neural activity fluctuates dynamically with time, and these changes have been reported to be of behavioral significance, despite occurring spontaneously. Through electroencephalography (EEG), fluctuations in alpha-band (8-14 Hz) activity have been identified over posterior sites that covary on a trial-by-trial basis with whether an upcoming visual stimulus will be detected or not. These fluctuations are thought to index the momentary state of visual cortex excitability. Here, we tested this hypothesis by directly exciting human visual cortex via transcranial magnetic stimulation (TMS) to induce illusory visual percepts (phosphenes) in blindfolded participants, while simultaneously recording EEG. We found that identical TMS-stimuli evoked a percept (P-yes) or not (P-no) depending on prestimulus alpha-activity. Low prestimulus alpha-band power resulted in TMS reliably inducing phosphenes (P-yes trials), whereas high prestimulus alpha-values led the same TMS-stimuli failing to evoke a visual percept (P-no trials). Additional analyses indicated that the perceptually relevant fluctuations in alpha-activity/visual cortex excitability were spatially specific and occurred on a subsecond time scale in a recurrent pattern. Our data directly link momentary levels of posterior alpha-band activity to distinct states of visual cortex excitability, and suggest that their spontaneous fluctuation constitutes a visual operation mode that is activated automatically even without retinal input.

Variations of oscillatory brain activity have been related to distinct functional states depending on the frequency of oscillations. In the alpha-band (about 8-14 Hz), decreased oscillatory activity is thought to reflect a state of enhanced cortical excitability, and increased activity to reflect a state of cortical idling or inhibition in which excitability is reduced, but the alpha/excitability link has not been probed directly. Here, we studied the relationship between resting oscillatory activity and visual cortex excitability across participants using electroencephalography and transcranial magnetic stimulation to the occipital pole. We found individual posterior alpha-band power to correlate with the individual threshold for eliciting illusory, transcranial magnetic stimulation-induced visual percepts. This provides direct support for an alpha/excitability link and for baseline states of the visual brain to vary across individuals.

Multisensory interactions occur early in time and in low-level cortical areas, including primary cortices. To test current models of early auditory-visual (AV) convergence in unisensory visual brain areas, we studied the effect of transcranial magnetic stimulation (TMS) of visual cortex on behavioral responses to unisensory (auditory or visual) or multisensory (simultaneous auditory-visual) stimulus presentation. Single-pulse TMS was applied over the occipital pole at short delays (30-150 ms) after external stimulus onset. Relative to TMS over a control site, reactions times (RTs) to unisensory visual stimuli were prolonged by TMS at 60-75 ms poststimulus onset (visual suppression effect), confirming stimulation of functional visual cortex. Conversely, RTs to unisensory auditory stimuli were significantly shortened when visual cortex was stimulated by TMS at the same delays (beneficial interaction effect of auditory stimulation and occipital TMS). No TMS-effect on RTs was observed for AV stimulation. The beneficial interaction effect of combined unisensory auditory and TMS-induced visual cortex stimulation matched and was correlated with the RT-facilitation after external multisensory AV stimulation without TMS, suggestive of multisensory interactions between the stimulus-evoked auditory and TMS-induced visual cortex activities. A follow-up experiment showed that auditory input enhances excitability within visual cortex itself (using phosphene-induction via TMS as a measure) over a similarly early time-window (75-120 ms). The collective data support a mechanism of early auditory-visual interactions that is mediated by auditory-driven sensitivity changes in visual neurons that coincide in time with the initial volleys of visual input.

Invasive recordings of local field potentials (LFPs) have been used to "read the mind" of monkeys in real time. Here we investigated whether noninvasive field potentials estimated from the scalp-recorded electroencephalogram (EEG) using the ELECTRA source localization algorithm could provide real-time decoding of mental states in healthy humans. By means of pattern recognition techniques on 500-ms EEG epochs, we were able to discriminate accurately from single trials which of four categories of visual stimuli the subjects were viewing. Our results show that it is possible to reproduce the decoding accuracy previously obtained in animals with invasive recordings. A comparison between the decoding results and the subjects' behavioral performance indicates that oscillatory activity (OA), elicited in specific brain regions) codes better for the visual stimulus category presented than the subjects' actual response, i.e., is insensitive to voluntary or involuntary errors. The identification of brain regions participating in the decoding process allowed us to construct 3D-functional images of the task-related OA. These images revealed the activation of brain regions known for their involvement in the processing of this type of visual stimuli. Electrical neuroimaging therefore appears to have the potential to establish what the brain is processing while the stimuli are being seen or categorized, i.e., concurrently with sensory-perceptual processes., (Copyright 2007 Wiley-Liss, Inc.)

Electroencephalographic studies in humans have demonstrated that orienting of visual attention induces a decrease in oscillatory alpha-band activity (alpha-desynchronization) over cortical areas tuned to the attended visual space. This is interpreted as reflecting intentionally enhanced excitability of these areas to facilitate upcoming visual processing. However, the inverse mechanism might also apply. Brain areas that process task-irrelevant space might be actively suppressed by increased alpha-activity (alpha-synchronization) to protect against input of distracter information. In the present study, we demonstrate that such suppression mechanisms are highly selective and are taking place even without distracters that need to be ignored. During voluntary orienting of attention, we found alpha-synchronization to dominate over desynchronization, to be topographically specific for each of eight attention positions, and to occur over areas processing unattended space in a retinotopically organized pattern. This indicates that alpha-synchronization is an important component of selective attention, serving active suppression of unattended positions during visual spatial orienting.

Steady-state auditory evoked responses (SSAER) obtained via electroencephalography (EEG) co-vary in amplitude with blood flow changes in the auditory area of the cerebellum. The aim of the present EEG study was to probe the cerebellar role in the control of such SSAER. For this purpose, we investigated changes in SSAERs due to transient disruption of the cerebellar hemisphere by repetitive transcranial magnetic stimulation (rTMS). SSAERs to click-trains of three different frequencies in the gamma-band (32, 40 and 47 Hz) were recorded from 45 scalp electrodes in six healthy volunteers immediately after 1-Hz rTMS and compared to baseline SSAERs assessed prior to magnetic stimulation. Cerebellar rTMS contralateral to the stimulated ear significantly reduced the amplitude of steady-state responses to 40-Hz click-trains and showed a tendency to reduce the amplitude to 32-Hz click-trains. No effects were observed for 47-Hz click-trains, nor for magnetic stimulation of the cerebellum ipsilateral to auditory stimulation or after sham stimulation. Our results suggest that interference with cerebellar output by rTMS modifies functional activity associated with cortical auditory processing. The finding of maximum effects on 40-Hz SSAERs provides support to the notion that the cerebellum is part of a distributed network involved in the regulation of cortical oscillatory activity and points at some frequency-specificity for the control of auditory-driven neuronal oscillations.

Covertly directing visual attention toward a spatial location in the absence of visual stimulation enhances future visual processing at the attended position. The neuronal correlates of these attention shifts involve modulation of neuronal "baseline" activity in early visual areas, presumably through top-down control from higher-order attentional systems. We used electroencephalography to study the largely unknown relationship between these neuronal modulations and behavioral outcome in an attention orienting paradigm. Covert visuospatial attention shifts to either a left or right peripheral position in the absence of visual stimulation resulted in differential modulations of oscillatory alpha-band (8-14 Hz) activity over left versus right posterior sites. These changes were driven by varying degrees of alpha-decreases being maximal contralateral to the attended position. When expressed as a lateralization index, these alpha-changes differed significantly between attention conditions, with negative values (alpha&#95;right < alpha&#95;left) indexing leftward and more positive values (alpha&#95;left < or = alpha&#95;right) indexing rightward attention. Moreover, this index appeared deterministic for processing of forthcoming visual targets. Collapsed over trials, there was an advantage for left target processing in accordance with an overall negative bias in alpha-index values. Across trials, left targets were detected most rapidly when preceded by negative index values. Detection of right targets was fastest in trials with most positive values. Our data indicate that collateral modulations of posterior alpha-activity, the momentary bias of visuospatial attention, and imminent visual processing are linked. They suggest that the momentary direction of attention, predicting spatial biases in imminent visual processing, can be estimated from a lateralization index of posterior alpha-activity.

Gamma band (30-80 Hz) oscillations arising in neuronal ensembles are thought to be a crucial component of the neural code. Recent studies in animals suggest a similar functional role for very high frequency oscillations (VHFO) in the range 80-200 Hz. Since some intracerebral studies in humans link VHFO to epileptogenesis, it remains unclear if VHFO appear in the healthy human brain and if so which is their role. This study uses EEG recordings from twelve healthy volunteers, engaged in a visuo-motor reaction time task, to show that VHFO are not necessarily pathological but rather code information about upcoming movements. Oscillations within the range (30-200 Hz) occurring in the period between stimuli presentation and the fastest hand responses allow highly accurate (>96%) prediction of the laterality of the responding hand in single trials. Our results suggest that VHFO belong in functional terms to the gamma band that must be considerably enlarged to better understand the role of oscillatory activity in brain functioning. This study has therefore important implications for the recording and analysis of electrophysiological data in normal subjects and patients.

Objective: We report on the development of an EEG recording system, comprised of electrodes and amplifiers that are compatible with TMS (single and rapid-rate) in both human and animal studies., Methods: We assembled a versatile multi-channel EEG recording system consisting of: (1) two types of electrodes that are safe during TMS or rTMS. (2) Low slew-rate EEG amplifiers that recover within a few milliseconds after the application of TMS pulses., Results: The two electrode types: (a) a conductive-plastic surface electrode with a conductive-silver epoxy coat and (b) a subdermal silver wire electrode (SWE) are compatible to TMS pulses. The amplifiers recover within 30 ms, so that the EEG can be viewed online, essentially without interruption and/or blocking or excessive artifact., Conclusions: Our TMS compatible electrode and EEG recording system allows safe and online viewing/recording of the subject's (human or animal) EEG/EP during experiments or studies involving TMS or rTMS applications. The TMS compatible electrode/amplifier system can be used with any EEG recording instrument., Significance: A simple recording technique coupled with new electrodes permit safe and readable EEG records during TMS in humans and animals. Such online monitoring of the EEG would allow control of TMS/rTMS parameters based on EEG activity.

Objective: The preliminary results of noninvasive brain stimulation for epilepsy treatment have been encouraging, but mixed. Two important factors may contribute to this heterogeneity: the altered brain physiology of patients with epilepsy and the variable presence of antiepileptic drugs. Therefore, we aimed to study the effects of 1 Hz rTMS on corticospinal excitability in patients with juvenile myoclonic epilepsy (JME) in two different conditions: low- or high-plasma valproate levels., Methods: Fifteen patients with JME and 12 age-matched healthy subjects participated in this study. Corticospinal excitability before and after 1 Hz rTMS was assessed in JME patients with low- and high-plasma valproate levels; and these results were compared with those in healthy subjects., Results: In patients with chronic use of valproate and low-plasma concentrations, 1 Hz rTMS had a similar significant inhibitory effect on corticospinal excitability as in healthy subjects. However, in the same patients when the serum valproate concentration was high, 1 Hz rTMS increased the corticospinal excitability significantly. In addition, there was a significant positive correlation between plasma valproate levels and the motor threshold changes after 1 Hz rTMS., Conclusions: Our findings can be accounted for by mechanisms of homeostatic plasticity and illustrate the dependency of the modulatory effects of rTMS on the physiologic state of the targeted brain cortex., Significance: The therapeutic use of rTMS in epilepsy should take into consideration the interaction between rTMS and drugs that change cortical excitability.

Repetitive transcranial magnetic stimulation (rTMS) holds promise as a therapeutic tool in major depression. However, a means to assess the effects of a single rTMS session on mood to guide subsequent sessions would be desirable. The present study examined the effects of a single rTMS session on an affective go/no-go task known to measure emotional-cognitive deficits associated with major depression. Ten patients with an acute episode of unipolar major depression and eight partially or completely remitted (improved) patients underwent 1 Hz rTMS over the left and right dorsolateral prefrontal cortex prior to task performance. TMS over the mesial occipital cortex was used as a control. We observed significantly improved performance in depressed patients following right prefrontal rTMS. This beneficial effect declined with decreasing depression severity and tended to reverse in the improved group. Left prefrontal rTMS had no significant effect in the depressed group, but it resulted in impaired task performance in the improved group. Our findings indicate that the acute response of depressed patients to rTMS varies with the stimulation site and depression severity. Further studies are needed to determine whether the present paradigm could be used to predict antidepressant treatment success or to individualize stimulation parameters according to specific pathology.

Luria (1947/1970) proposed left-top positions in arm folding (AF) and hand clasping (HC) to be signs of "latent left-handedness". However, research since has revealed that (1) left-top positions are canonical for European right-handers and (2) combined AF/HC measures may provide more information about cerebral laterality than either measure considered alone. We tested whether AF and HC or AF/HC combinations predicted diminished right-handedness for 12 handedness items. Results from 509 healthy participants showed that (1) left-top positions in AF and HC were dominant across participants, as was right-handedness, and a right-top position in HC was associated with attenuated right-handedness, (2) right-hand preference was more frequently associated with congruent AF/HC combinations, especially of the LL type (AF: left-top / HC: left-top), and (3) non-right-hand preference was associated with non-congruent, predominantly LR combinations. We conjecture that the LL type combination indicates left hemispheric dominance for motor actions, whereas the LR combination, in which HC as the distally innervated posture deviates from the canonical pattern, indicates attenuated hemispheric asymmetry. Our data support Luria's proposition that a left-top preference in AF points to "latent" left-handedness, but only if associated with a right-top preference in HC. Consistent left-top preference for the combined AF/HC measure appears to predict right-handedness.

Visuo-spatial attention tends to be prioritized towards emotionally negative stimuli such as fearful faces, as opposed to neutral or positive stimuli. Using a covert orienting task, we previously showed that a lateral occipital P1 component, with extrastriate neural sources, was selectively enhanced to lateralized visual targets replacing a fearful face (fear-valid trial) than the same targets replacing a neutral face (fear-invalid trial), providing evidence for exogenous spatial orienting of attention towards threat cues. Here, we describe a new analysis of these data, using topographic evoked potentials mapping methods combined with a distributed source localization technique. We show that an early field topography (40-80 ms post-target onset) with a centro-parietal negativity and a left posterior parietal source distinguished fear-valid from fear-invalid trials, whereas a distinct activity with anterior cingulate sources was selectively evoked during fear-invalid trials. At the same latency, or later, no difference in field topography was found for valid compared to invalid trials with happy faces. The early parietal map preceded a modulation in amplitude of the field strength (approximately 130 ms), corresponding to the enhanced lateral occipital P1 during valid trials in the fear condition. Furthermore, this early topography at 40-80 ms was positively correlated with the subsequent amplitude modulation of P1 at 130-160 ms in the fear condition, suggesting a possible functional coupling between these two successive events. These data have important implications for models of spatial attention and interactions with emotion. They suggest two successive stages of neural activity during exogenous orienting of attention towards visual targets following fearful faces, including an early posterior parietal negativity, followed by gain control mechanisms enhancing visual responses in extrastriate occipital cortex.

Patients with lesions in posterior parietal cortex (PPC) are relatively unimpaired in voluntarily directing visual attention to different spatial locations, while many neuroimaging studies in healthy subjects suggest dorsal PPC involvement in this function. We used an offline repetitive transcranial magnetic stimulation (rTMS) protocol to study this issue further. Ten healthy participants performed a cue-target paradigm. Cues prompted covert orienting of spatial attention under voluntary control to either a left or right visual field position. Targets were flashed subsequently at the cued or uncued location, or bilaterally. Following rTMS over right dorsal PPC, (i) the benefit for target detection at cued versus uncued positions was preserved irrespective of cueing direction (left- or rightward), but (ii) leftward cueing was associated with a global impairment in target detection, at all target locations. This reveals that leftward orienting was still possible after right dorsal PPC stimulation, albeit at an increased overall cost for target detection. In addition, rTMS (iii) impaired left, but (iv) enhanced right target detection after rightward cueing. The finding of a global drop in target detection during leftward orienting with a spared, relative detection benefit at the cued (left) location (i-ii) suggests that right dorsal PPC plays a subsidiary rather than pivotal role in voluntary spatial orienting. This finding reconciles seemingly conflicting results from patients and neuroimaging studies. The finding of attentional inhibition and enhancement occurring contra- and ipsilaterally to the stimulation site (iii-iv) supports the view that spatial attention bias can be selectively modulated through rTMS, which has proven useful to transiently reduce visual hemispatial neglect.

Functional neuroimaging studies have associated affective go/no-go function with lateral prefrontal activation, but they have not established a causal role and have not determined whether one hemisphere is predominantly engaged. In the present study, 11 normal volunteers underwent slow repetitive transcranial magnetic stimulation of the left and right dorsolateral prefrontal cortex, and the occipital cortex prior to performance of a picture-based affective go/no-go task. We found an interfering effect of left prefrontal repetitive transcranial magnetic stimulation compared with both right prefrontal and occipital repetitive transcranial magnetic stimulation. This impairment concerned positive and negative task stimuli to a similar extent, and tended to be greater in shift compared with nonshift blocks. Our findings demonstrate a functionally relevant lateralization of the prefrontal contribution to affective go/no-go tasks.

We describe an electroencephalographic (EEG) device and protocol that allows recording of electrophysiological signals generated by the human brain during transcranial magnetic stimulation (TMS) despite the TMS-induced high-voltage artifacts. The key hardware components include slew-rate limited preamplifiers to prevent saturation of the EEG system due to TMS. The protocol involves artifact subtraction to isolate the electrophysiological signals from residual TMS-induced contaminations. The TMS compatibility of the protocol is illustrated with examples of two data sets demonstrating the feasibility of the approach in the single-pulse TMS design, as well as during repetitive TMS. Our data show that both high-amplitude potentials evoked by visual checkerboard stimulation and low-amplitude steady-state oscillations induced by auditory click-trains can be retrieved with the present protocol. The signals recorded during TMS perfectly matched control EEG responses to the same visual and auditory stimuli. The main field of application of the present protocol is in cognitive neuroscience complementing behavioral studies that use TMS to induce transient, 'virtual lesions'. Combined EEG-TMS techniques provide neuroscientists with a unique method to test hypothesis on functional connectivity, as well as on mechanisms of functional orchestration, reorganization, and plasticity.

The spatial unity of self and body is challenged by various philosophical considerations and several phenomena, perhaps most notoriously the "out-of-body experience" (OBE) during which one's visual perspective and one's self are experienced to have departed from their habitual position within one's body. Although researchers started examining isolated aspects of the self, the neurocognitive processes of OBEs have not been investigated experimentally to further our understanding of the self. With the use of evoked potential mapping, we show the selective activation of the temporoparietal junction (TPJ) at 330-400 ms after stimulus onset when healthy volunteers imagined themselves in the position and visual perspective that generally are reported by people experiencing spontaneous OBEs. Interference with the TPJ by transcranial magnetic stimulation (TMS) at this time impaired mental transformation of one's own body in healthy volunteers relative to TMS over a control site. No such TMS effect was observed for imagined spatial transformations of external objects, suggesting the selective implication of the TPJ in mental imagery of one's own body. Finally, in an epileptic patient with OBEs originating from the TPJ, we show partial activation of the seizure focus during mental transformations of her body and visual perspective mimicking her OBE perceptions. These results suggest that the TPJ is a crucial structure for the conscious experience of the normal self, mediating spatial unity of self and body, and also suggest that impaired processing at the TPJ may lead to pathological selves such as OBEs.

Response to a stimulus is faster when a subject is attending and knows beforehand how to respond. It has been suggested recently that this occurs because ongoing neuronal activity is spatially and temporally structured during states of expectancy preceding a stimulus. This mechanism is believed to mediate top-down processing, facilitating the early grouping and selection of distributed neuronal ensembles implicated in ensuing sensory-motor processing. To validate this model, it must be shown that some features of this early ongoing neural activity are correlated with subsequent perceptual decisions or behavioral events. We investigated this hypothesis in an electrophysiologic study in 12 subjects carrying out a simple visuomotor reaction-time task. Local field potentials (LFP) at each brain voxel were estimated using a linear distributed inverse solution termed "ELECTRA" for each single trial of each subject. The energy of oscillations for different frequency bands was computed for the period between the warning cue and visual stimuli by applying a time-frequency decomposition to the estimated LFP. A nonparametric correlation coefficient was then calculated between energy of oscillations and reaction times for each single sweep. Gamma band oscillatory activity in a frontoparietal network before stimulus onset significantly correlated with reaction time for a significant amount of subjects. These results provide direct evidence for the role of neural oscillations as a top-down attentional control mechanism that mediates the speed of motor actions., ((c) 2004 Wiley-Liss, Inc.)

Studies evaluating the role of the executive motor system in motor imagery came to a general agreement in favour of the activation of the primary motor area (M1) during imagery, although in reduced proportion as compared to motor execution. It is still unclear whether this difference occurs within the preparation period or the execution period of the movement, or both. In the present study, EEG was used to investigate separately the preparation and the execution periods of overt and covert movements in adults. We designed a paradigm that randomly mixed actual and kinaesthetic imagined trials of an externally paced sequence of finger key presses. Sixty channel event-related potentials were recorded to capture the cerebral activations underlying the preparation for motor execution and motor imagery, as well as cerebral activations implied in motor execution and motor imagery. Classical waveform analysis was combined with data-driven spatiotemporal segmentation analysis. In addition, a LAURA source localization algorithm was applied to functionally define brain related motor areas. Our results showed first that the difference between actual and mental motor acts takes place at the late stage of the preparation period and consists of a quantitative modulation of the activity of common structures in M1. Second, they showed that primary motor structures are involved to the same extent in the actual or imagined execution of a motor act. These findings reinforce and refine the functional equivalence hypothesis between actual and imagined motor acts.

We have addressed the role of occipital and somatosensory cortex in a tactile discrimination task. Sight-ed and congenitally blind subjects rated the roughness and distance spacing for a series of raised dot patterns. When judging roughness, intermediate dot spacings were perceived as being the most rough, while distance judgments generated a linear relation. Low-frequency rTMS applied to somatosensory cortex disrupted roughness without affecting distance judgments, while rTMS to occipital cortex disrupted distance but not roughness judgments. We also tested an early blind patient with bilateral occipital cortex damage. Her performance on the roughness determination task was normal; however, she was greatly impaired with distance judgments. The findings suggest a double-dissociation effect in which roughness and distance are primarily processed in somatosensory and occipital cortex, respectively. The differential effect of rTMS on task performance and corroborative clinical evidence suggest that occipital cortex is engaged in tactile tasks requiring fine spatial discrimination.

Objective: To further evaluate the potential of slew-rate limiting amplifiers to record electrophysiological signals in spite of concurrent transcranial magnetic stimulation (TMS), and to explore the effects of single-pulse TMS on electroencephalographic (EEG) correlates of functional brain activity., Methods: Visual-evoked potentials (VEPs) to checkerboards were recorded in 7 right-handed subjects, while single-pulse TMS was applied to the occipital pole either at visual stimulus onset, during the build-up or at the expected peak of the early VEP component P1 (VIS&TMS). Timing of TMS was individually adjusted based on each subject's VEP-latency. A condition of TMS without concurrent visual stimulation (TMS(alone)) served for subtraction purposes (VIS&TMS minus TMS(alone)) to partial out TMS-related contaminations of the EEG signal., Results: When TMS was applied at visual stimulus onset, VEPs (as calculated by subtraction) perfectly matched control VEPs to visual stimulation alone. TMS at around P1, in contrast, modified the targeted (P1) and the subsequent VEP component (N1), independently of whether TMS was given at build-up or peak., Conclusions: The retrieval of regular VEPs with concomitant TMS at visual stimulus onset suggests that the employed EEG system and subtraction procedure are suited for combined EEG-TMS studies. The VEP changes following TMS at around P1 provide direct clues on the temporal dynamics of TMS pulse effects on functional activity in the human brain. Our data suggest effects of relatively long duration (approximately 100 ms) when TMS is applied while functional neuronal activity evolves.

Based on earlier reports of inconsistent postural preferences in psychiatric populations, we assessed arm folding (AF) and hand clasping (HC) postures in 361 healthy right-handers with different degrees of magical ideation (MI). Subjects generally preferred the congruent combination of a left-arm-top (AF) and a left-thumb-top position (HC). Incongruent subjects, in particular those with a right-arm-top/left-thumb-top position, scored higher on MI than did congruent subjects. Our main finding was thus an association between a style of thinking (MI) and lateral preferences for limb postures, inconsistent across distal (primarily contralaterally innervated) and more proximal (more bilaterally innervated) parts of the body. This association is interpreted in the frame of lesser hemispheric asymmetries in subjects bordering to schizotypal personalities.

Objective: To investigate a modulation of the N170 face-sensitive component related to the perception of other-race (OR) and same-race (SR) faces, as well as differences in face and non-face object processing, by combining different methods of event-related potential (ERP) signal analysis., Methods: Sixty-two channel ERPs were recorded in 12 Caucasian subjects presented with Caucasian and Asian faces along with non-face objects. Surface data were submitted to classical waveforms and ERP map topography analysis. Underlying brain sources were estimated with two inverse solutions (BESA and LORETA)., Results: The N170 face component was identical for both race faces. This component and its topography revealed a face specific pattern regardless of race. However, in this time period OR faces evoked significantly stronger medial occipital activity than SR faces. Moreover, in terms of maps, at around 170 ms face-specific activity significantly preceded non-face object activity by 25 ms. These ERP maps were followed by similar activation patterns across conditions around 190-300 ms, most likely reflecting the activation of visually derived semantic information., Conclusions: The N170 was not sensitive to the race of the faces. However, a possible pre-attentive process associated to the relatively stronger unfamiliarity for OR faces was found in medial occipital area. Moreover, our data provide further information on the time-course of face and non-face object processing.

Visual-induced alpha desynchronization (VID) and visual-evoked potentials (VEPs) characterize occipital activation in response to visual stimulation but their exact relationship is unclear. Here, we tested the hypothesis that VID and VEPs reflect different aspects of cortical activation. For this purpose, we determined whether VID and VEPs are differentially modulated by low-frequency repetitive transcranial magnetic stimulation (rTMS) over the occipital pole. Scalp EEG responses to visual stimuli (flashed either to the left or to the right visual field) were recorded for 8 min in six healthy subjects (1) before, (2) immediately following, and (3) 20 min after left occipital rTMS (1 Hz, 10 min). The parameters aimed to reduce cortical excitability beyond the end of the TMS train. In addition, simple reaction times to visual stimulation were recorded (left or right hand in separate blocks). In all subjects, VID was significantly and prominently reduced by rTMS (P = 0.0001). In contrast, rTMS failed to modulate early VEP components (P1/N1). A moderate effect was found on a late VEP component close to manual response onset (P = 0.014) but this effect was in the opposite direction to the VID change. All changes were restricted to the targeted left occipital cortex. The effects were present only after right visual field stimulation when a right hand response was required, were associated with a behavioral effect, and had washed out 20 min after rTMS. We conclude that VID and early VEPs represent different aspects of cortical activation. The findings that rTMS did not change early VEPs and selectively affected VID and late VEPs in conditions where the visual input must be transferred intrahemispherically for visuomotor integration (right visual field/right hand) are suggestive of rTMS interference with higher-order visual functions beyond visual input. This is consistent with the idea that alpha desynchronization serves an integrative role through a corticocortical "gating function."

Recent studies have revealed a distinct cortical network activated during the analysis of sounds' spatial properties. Whether common brain regions in this auditory where pathway are involved in both auditory motion and location processing is unresolved. We investigated this question with multichannel auditory evoked potentials (AEPs) in 11 subjects. Stimuli were binaural 500-ms white noise bursts. Interaural time differences (ITD) created the sensation of moving or stationary sounds within each auditory hemifield, and subjects discriminated either their position or direction of motion in a blocked design. Scalp potential distributions (AEP maps) differentiated electric field configurations across stimulus classes. The initial approximately 250-ms poststimulus yielded common topographies for both stimulus classes and hemifields. After approximately 250-ms, moving and stationary sounds engaged distinct cortical networks at two time periods, again with no differences observed between hemifields. The first ( approximately 250- to 350-ms poststimulus onset) was during stimulus presentation, and the second ( approximately 550- to 900-ms poststimulus onset) occurred after stimulus offset. Distributed linear inverse solutions of the maps over the 250- to 350-ms time period revealed not only bilateral inferior frontal activation for both types of auditory spatial processing, but also strong right inferior parietal activation in the case of auditory motion discrimination. During the later 550-to 900-ms time period, right inferior parietal and bilateral inferior frontal activity was again observed for moving sounds, whereas strong bilateral superior frontal activity was seen in the case of stationary sounds. Collectively, the evidence supports the existence of partly segregated networks within the auditory where pathway for auditory location and auditory motion processing., (2002 Elsevier Science (USA).)

With the purpose of investigating motor and cognitive lateralization profiles associated with long-term motor training, we investigated differences in hemispheric specialization between proficient judo sportsmen and controls through the assessment of a number of handedness and footedness items including postural preferences as well as dichotic listening and lateralized visual field tests. Our data show that: (1) the different handedness and footedness items did differently relate to each other within the athlete and control groups as revealed by a principle component analysis (PCA); (2) stand side correlated differently to these motor profile factors in athletes and controls; (3) athletes preferred more frequently to perform certain movements with the left hand than controls, although overall right-handed; (4) this was especially true for athletes which proved to be most proficient/skilled; and (5) in a lateralized verbal listening task and a lateralized visual field task athletes revealed enhanced right-hemispheric involvement relative to controls. Our results suggest that during motor and postural skill acquisitions (long-term judo training) lateral preferences are modified, probably due to neuroplasticity. Moreover, the present findings support the multidimensional view of handedness by Steenhuis and Bryden [Cortex 25 (1989) 289] and the notion of a right-hemispheric "praxis system" involved in skilled action routines within peripersonal space [Brain and Cognition 23 (1993) 181].

Face perception and recognition is an intriguing ability, already present in neonates. Numerous studies in patients with brain lesions identified the temporo-occipital cortex as the crucial structure for this capacity. Analysis of electrical signals (EEG) inside the brain of patients implanted with intracranial electrodes for diagnostic purposes allows researchers to describe the temporal and spatial organization of responses to various aspects of face processing in human subjects. Several findings have emerged and appear relevant for cerebral organization in general: (1) Selective face responses were obtained from the basal temporo-occipital cortex at around 200 ms (N200); however, other structures such as the lateral temporal lobe and frontal cortex also participate in face recognition and perception tasks. (2) Each structure has a distinct "response profile"; that is, with respect to a given task certain structures respond strongly, others less or not at all. This profile might change with a different task, although the physical parameters of the stimuli remain the same. (3) The right hemispheric predominance of face processing, as suggested by patient data and studies in healthy volunteers, seemed to be restricted to its early stages (i.e., before 100-150 ms). (4) Recognition of faces might be associated with differential intracranial responses, despite an incorrect overt response, reflecting neurophysiological correlates of implicit memory. (5) The more the stimulus resembled a complete human face, the earlier and larger the N200 response was found, in particular over the basal temporobasal cortex. Analysis of electrical signals from intracranial electrodes might help to improve our understanding of the underlying physiological and anatomical constraints of cognitive processes.

We review recent methodological advances in electromagnetic source imaging and present EEG data from our laboratory obtained by application of these methods. There are two principal steps in our analysis of multichannel electromagnetic recordings: (i) the determination of functionally relevant time periods in the ongoing electric activity and (ii) the localization of the sources in the brain that generate these activities recorded on the scalp. We propose a temporal segmentation of the time-varying activity, which is based on determination of changes in the topography of the electric fields, as an approach to the first step, and a distributed linear inverse solution based on realistic head models as an approach to the second step. Data from studies of visual motion perception, visuo-motor transfer, mental imagery, semantic decision, and cognitive interference illustrate that this analysis allows us to define the patterns of electric activity that are present at given time periods after stimulus presentation, as well as those time periods where significantly different patterns appear between different stimuli and tasks. The presented data show rapid and parallel activation of different areas within complex neuronal networks, including early activity of brain regions remote from the primary sensory areas. In addition, the data indicate information exchange between homologous areas of the two hemispheres in cases where unilateral stimulus presentation requires interhemispheric transfer.

Objectives: Characterization of the EEG pattern during the early phase of a seizure is crucial for identifying the epileptic focus. The purpose of the present investigation was to evaluate a method that divides ictal EEG activity into segments of relatively constant surface voltage distribution, and to provide a 3-dimensional localization of the activity during the different segments., Methods: For each timepoint the electrical voltage distribution on the scalp (the voltage map) was determined from the digitized EEG recording. Through a spatial cluster analysis time sequences where the maps did not change much (segments) were identified, and a 3-dimensional source reconstruction of the activity corresponding to the different mean maps was performed using a distributed linear inverse solution algorithm., Results: Segments dominating early in seizure development were identified, and source reconstruction of the EEG activity corresponding to the maps of these segments yielded results which were consistent with the results from invasive recordings. In some cases a sequence of consecutive segments was obtained, which might reflect ictal propagation., Conclusions: Segmentation of ictal EEG with subsequent 3-dimensional source reconstruction is a useful method to non-invasively determine the initiation and perhaps also the spread of epileptiform activity in patients with epileptic seizures.

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

The present study reports on a patient undergoing invasive monitoring for intractable epilepsy who experienced different vestibular sensations after electrical cortical stimulation of the inferior parietal lobule at the anterior part of the intraparietal sulcus. Types of vestibular response ranged from simple to complex sensations and depended on stimulation site and applied current. The findings suggest vestibular topography and hierarchical processing within the parietal vestibular cortex of humans.