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An episode of complete failure to respond during an attentive task accompanied by behavioural signs of sleep is called a behavioural microsleep. We proposed a combination of high-resolution EEG and an advanced method for time-varying effective connectivity estimation for reconstructing the temporal evolution of the causal relations between cortical regions when microsleeps occur during a continuous visuomotor task. We found connectivity patterns involving left-right frontal, left-right parietal, and left-frontal/right-parietal connections commencing in the interval [-500; -250] ms prior to the onset of microsleeps and disappearing at the end of the microsleeps. Our results from global graph indices derived from effective connectivity analysis have revealed EEG-based biomarkers of all stages of microsleeps (preceding, onset, pre-recovery, recovery). In particular, this raises the possibility of being able to predict microsleeps in real-world tasks and initiate a 'wake-up' intervention to avert the microsleeps and, hence, prevent injurious and even multi-fatality accidents., (Copyright © 2015 Elsevier Inc. All rights reserved.)

Partial Directed Coherence (PDC) is a powerful estimator of effective connectivity. In neuroscience it is used in different applications with the aim to investigate the communication between brain regions during the execution of different motor or cognitive tasks. When multiple trials are available, PDC can be computed over multiple realizations, provided that the assumption of stationarity across trials is verified. This allows to improve the amount of data, which is an important constraint for the estimation accuracy. However, the stationarity of the data across trials is not always guaranteed, especially when dealing with patients. In this study we investigated how the inter-trials variability of an EEG dataset affects the PDC accuracy. Effects of density variations and of changes of connectivity values across trials were first investigated with a simulation study and then tested on real EEG data collected from two post-stroke patients during a motor imagery task and characterized by different inter-trials variability. Results showed the effect of different factors on the PDC accuracy and the robustness of such estimator in a range of conditions met in practical applications.

The present research investigates the neurophysiological activity elicited by fast observations of faces of real candidates during simulated political elections. We used simultaneous recording of electroencephalographic (EEG) signals as well as galvanic skin response (GSR) and heart rate (HR) as measurements of central and autonomic nervous systems. Twenty healthy subjects were asked to give judgments on dominance, trustworthiness, and a preference of vote related to the politicians' faces. We used high-resolution EEG techniques to map statistical differences of power spectral density (PSD) cortical activity onto a realistic head model as well as partial directed coherence (PDC) and graph theory metrics to estimate the functional connectivity networks and investigate the role of cortical regions of interest (ROIs). Behavioral results revealed that judgment of dominance trait is the most predictive of the outcome of the simulated elections. Statistical comparisons related to PSD and PDC values highlighted an asymmetry in the activation of frontal cortical areas associated with the valence of the judged trait as well as to the probability to cast the vote. Overall, our results highlight the existence of cortical EEG features which are correlated with the prediction of vote and with the judgment of trustworthy and dominant faces.

Recent findings suggest that the preparation and execution of voluntary self-paced movements are accompanied by the coordination of the oscillatory activities of distributed brain regions. Here, we use electroencephalographic source imaging methods to estimate the cortical movement-related oscillatory activity during finger extension movements. Then, we apply network theory to investigate changes (expressed as differences from the baseline) in the connectivity structure of cortical networks related to the preparation and execution of the movement. We compute the topological accessibility of different cortical areas, measuring how well an area can be reached by the rest of the network. Analysis of cortical networks reveals specific agglomerates of cortical sources that become less accessible during the preparation and the execution of the finger movements. The observed changes neither could be explained by other measures based on geodesics or on multiple paths, nor by power changes in the cortical oscillations.

In the present study, we propose a theoretical graph procedure to investigate multiple pathways in brain functional networks. By taking into account all the possible paths consisting of h links between the nodes pairs of the network, we measured the global network redundancy R(h) as the number of parallel paths and the global network permeability P(h) as the probability to get connected. We used this procedure to investigate the structural and dynamical changes in the cortical networks estimated from a dataset of high-resolution EEG signals in a group of spinal cord injured (SCI) patients during the attempt of foot movement. In the light of a statistical contrast with a healthy population, the permeability index P(h) of the SCI networks increased significantly (P < 0.01) in the Theta frequency band (3-6 Hz) for distances h ranging from 2 to 4. On the contrary, no significant differences were found between the two populations for the redundancy index R(h). The most significant changes in the brain functional network of SCI patients occurred mainly in the lower spectral contents. These changes were related to an improved propagation of communication between the closest cortical areas rather than to a different level of redundancy. This evidence strengthens the hypothesis of the need for a higher functional interaction among the closest ROIs as a mechanism to compensate the lack of feedback from the peripheral nerves to the sensomotor areas.

In this paper we show how the possibility of recording simultaneously the cerebral neuroelectric activity in different subjects (EEG hyperscanning) during the execution of different tasks could return useful information about the "internal" cerebral state of the subjects. We present the results obtained by EEG hyperscannings during ecological task (such as the execution of a card game) as well as that obtained in a series of couples of subjects during the performance of the Prisoner's Dilemma Game. The simultaneous recordings of couples of interacting subjects allows to observe and to model directly the neural signature of human interactions in order to understand the cerebral processes generating and generated by social cooperation or competition. Results obtained in a study of different groups recorded during the card game revealed a larger activity in prefrontal and anterior cingulated cortex in different frequency bands for the player that leads the game when compared to other players. Results collected in a population of 10 subjects during the performance of the Prisoner's Dilemma suggested that the most consistently activated structure is the orbitofrontal region (roughly described by the Brodmann area 10) during the condition of competition in both the tasks. It could be speculated whether the pattern of cortical connectivity between different cortical areas in different subjects could be employed as a tool for assessing the outcome of the task in advance.

The evaluation of the topological properties of brain networks is an emergent research topic, since the estimated cerebral connectivity patterns often have relatively large size and complex structure. Since a graph is a mathematical representation of a network, the use of a theoretical graph approach would describe concisely the topological features of the functional brain connectivity network estimated using neuroimaging techniques. In the present study, we analyze the changes in brain synchronization networks using high-resolution EEG signals obtained during performance of a complex goal-directed visuomotor task. Our results show that the cortical network is more stable when subjects reach the goal than when they fail by hitting an obstacle. These findings suggest the presence of a possible cerebral "marker" for motor actions that result in successful reaching of a target.

The aim of this study was to elucidate if the TV commercials that were remembered by the subjects after their observation within a documentary elicited particular brain activity when compared to the activity generated during the observation of TV commercials that were forgotten. High resolution EEG recordings were performed in a group of 10 healthy subjects with the steady state somatosensory evoked potentials (SSSEPs) technique, in which a series of light electrical stimulation at the left wrist were delivered at the frequency of 20Hz. The brain activity was indexed by the phase delay of the EEG spectral responses at 20Hz with respect to the stimulus delivering and evaluated at the scalp level as well as at the cortical surface using several regions of interest coincident with the Brodmann areas (BAs). Results suggest that the cerebral processes involved during the observation of TV commercials that were remembered by the population examined (RMB dataset) are generated by the posterior parietal cortices and the prefrontal areas, rather bilaterally. These results are compatible with previously results obtained in literature by using MEG and fMRI devices during similar experimental tasks. High resolution EEG is able to summarize, with the use of SSSEPs methodologies, the behavior of the estimated cortical networks subserving the proposed memory tasks. It is likely that such tool could play a role in the next future for the investigation of the neural substrates of the human behavior in decision-making and recognition tasks.

One of the most challenging questions open in Neuroscience today is the characterization of the brain responses during social interaction. A major limitation of the approaches used in most of the studies performed so far is that only one of the participating brains is measured each time. The "interaction" between cooperating, competing or communicating brains is thus not measured directly, but inferred by independent observations aggregated by cognitive models and assumptions that link behavior and neural activation. In this paper, we present the results of the simultaneous neuroelectric recording of 5 couples of subjects engaged in cooperative games (EEG hyperscanning). The simultaneous recordings of couples of interacting subjects allows to observe and model directly the neural signature of human interactions in order to understand the cerebral processes generating and generated by social cooperation or competition. We used a paradigm called Prisoner's dilemma derived from the game theory. Results collected in a population of 10 subjects suggested that the most consistently activated structure in social interaction paradigms is the orbitofrontal region (roughly described by the Brodmann area 10) during the condition of competition.

Brain-computer interface (BCI) applications were initially designed to provide final users with special capabilities, like writing letters on a screen, to communicate with others without muscular effort. In these last few years, the BCI scientific community has been interested in bringing BCI applications outside the scientific laboratories, initially to provide useful applications in everyday life and in future in more complex environments, such as space. Recently, we implemented a control of a domestic environment realized with BCI applications. In the present chapter, we analyze the methodological approach employed to allow the interaction between subjects and domestic devices by use of noninvasive EEG recordings. In particular, we analyze whether the cortical activity estimated from noninvasive EEG recordings could be useful in detecting mental states related to imagined limb movements. We estimate cortical activity from high-resolution EEG recordings in a group of healthy subjects by using realistic head models. Such cortical activity was estimated in a region of interest associated with the subjects' Brodmann areas by use of depth-weighted minimum norm solutions. Results show that the use of the estimated cortical activity instead of unprocessed EEG improves the recognition of the mental states associated with limb-movement imagination in a group of healthy subjects. The BCI methodology here presented has been used in a group of disabled patients to give them suitable control of several electronic devices disposed in a three-room environment devoted to neurorehabilitation. Four of six patients were able to control several electronic devices in the domotic context with the BCI system, with a percentage of correct responses averaging over 63%.

In the present study, we propose a theoretical graph procedure to investigate the communication redundancy in brain networks. By taking into account all the possible paths between pairs of cortical regions, this method captures the network redundancy i.e. a critical resource of the brain enhancing the resilience to neural damages and dysfunctions. As an example for its potential, we apply this procedure to the cortical networks estimated from high-resolution EEG signals in a group of spinal cord injured patients during the attempt of the foot movement. Preliminary results suggest that in the high spectral contents the effects due to the spinal trauma affect the expected redundancy attitude by suppressing mainly the longer alternative pathways between the cortical regions.

Objectives: The main objective is to show current topics and future trends in the field of medical signal processing which are derived from current research concepts. Signal processing as an integrative concept within the scope of medical informatics is demonstrated., Methods: For all examples time-variant multivariate autoregressive models were used. Based on this modeling, the concept of Granger causality in terms of the time-variant Granger causality index and the time-variant partial directed coherence was realized to investigate directed information transfer between different brain regions., Results: Signal informatics encompasses several diverse domains including: processing steps, methodologies, levels and subject fields, and applications. Five trends can be recognized and in order to illustrate these trends, three analysis strategies derived from current neuroscientific studies are presented. These examples comprise high-dimensional fMRI and EEG data. In the first example, the quantification of time-variant-directed information transfer between activated brain regions on the basis of fast-fMRI data is introduced and discussed. The second example deals with the investigation of differences in word processing between dyslexic and normal reading children. Different dynamic neural networks of the directed information transfer are identified on the basis of event-related potentials. The third example shows time-variant cortical connectivity networks derived from a source model., Conclusions: These examples strongly emphasize the integrative nature of signal informatics, encompassing processing steps, methodologies, levels and subject fields, and applications.

We investigated brain activity during the observation of TV commercials by tracking the cortical activity and the functional connectivity changes in normal subjects. The aim was to elucidate if the TV commercials that were remembered by the subjects several days after their first observation elicited particular brain activity and connectivity compared with those generated during the observation of TV commercials that were quickly forgotten. High-resolution electroencephalogram (EEG) recordings were performed in a group of healthy subjects and the cortical activity during the observation of TV commercials was evaluated in several regions of interest coincident with the Brodmann areas (BAs). The patterns of cortical connectivity were obtained in the four principal frequency bands, Theta (3-7 Hz), Alpha (8-12 Hz), Beta (13-30 Hz), Gamma (30-40 Hz) and the directed influences between any given pair of the estimated cortical signals were evaluated by use of a multivariate spectral technique known as partial directed coherence. The topology of the cortical networks has been identified with tools derived from graph theory. Results suggest that the cortical activity and connectivity elicited by the viewing of the TV commercials that were remembered by the experimental subjects are markedly different from the brain activity elicited during the observation of the TV commercials that were forgotten. In particular, during the observation of the TV commercials that were remembered, the amount of cortical spectral activity from the frontal areas (BA 8 and 9) and from the parietal areas (BA 5, 7, and 40) is higher compared with the activity elicited by the observation of TV commercials that were forgotten. In addition, network analysis suggests a clear role of the parietal areas as a target of the incoming flow of information from all the other parts of the cortex during the observation of TV commercials that have been remembered. The techniques presented here shed new light on all the cortical networks and their behavior during the memorization of TV commercials. Such techniques could also be relevant in neuroeconomics and neuromarketing for the investigation of the neural substrates subserving other decision-making and recognition tasks.

Objective: This work intends to evaluate the functional characteristics of the cerebral network during the successful memory encoding of TV commercials., Methods: We estimated the functional networks in the frequency domain from a set of high-resolution EEG data in a group of healthy subjects during the showing of commercial spots within a neutral documentary. We evaluated the differences in the cortical network associated with later remembered and not-remembered commercials by calculating the global-E(g) and local-efficiency E(l) indexes., Results: Successful encoding of TV spots significantly affects the functional communication among cortical areas irrespectively of the frequency band. During the visualization of the video-clips that will be forgotten (FRG), the cortical network exhibited high values of global- and local-efficiency, reflecting a small-world configuration. During the visualization of the video-clips that will be remembered (RMB), the same indexes appeared significantly lower. This difference was especially evident for E(l) in the alpha band and for E(g) in the beta and gamma bands., Conclusions: The presence of attentional and semantic processes during the RMB condition leads to a significant reduction of the network communication efficiency., Significance: Such a reduction could represent a predictive measure of accurate recalls of TV spots.

The present work intends to evaluate the dynamics of the cerebral networks during the preparation and the execution of the foot movement. In order to achieve this objective, we have used mathematical tools capable of estimating the cortical activity via high-resolution EEG techniques. Afterwards we estimated, the instantaneous relationships occurring among the time-series of sixteen regions of interest (ROIs) in the Alpha (7-12 Hz) and Beta (13-29 Hz) band through the adaptive multivariate autoregressive models. Eventually, we evaluated the weighted-topology of the cerebral networks by calculating some theoretical graph indexes. The results show that the main structural changes are encoded in the highest spectral contents (Beta band). In particular, during the execution of the foot movement the cingulate motor areas (CM) work as network "hubs" presenting a large amount of outgoing links to the other ROIs. Moreover, the connectivity pattern changes its structure according to the different temporal stages of the task. In particular, the communication between the ROIs reaches its highest level of efficiency during the preparation of the foot movement, as revealed by the "small-world" property of the network, which is characterized by the presence of abundant clustering connections combined with short average distances between the cortical areas.

The present work intends to evaluate the functional characteristics of the cerebral network during the successful memory encoding of TV commercials. We estimated the functional networks in the frequency domain from a set of high-resolution EEG data. High resolution EEG recordings were performed in a group of healthy subjects and the cortical activity during the observation of TV commercials was evaluated in several regions of interest coincident with the Brodmann areas (BAs). Summarizing the main results of the present study, a sign of the memorization of a particular set of TV commercials have been found in a group of investigated subjects with the aid of advanced modern tools for the acquisition and the processing of EEG data. The cerebral processes involved during the observation of TV commercials that were remembered successively by the population examined (RMB dataset) are generated by the posterior parietal cortices and the prefrontal areas, rather bilaterally and are irrespective of the frequency bands analyzed. Such results are compatible with previously results obtained from EEG recordings with superficial electrodes as well as with the brain activations observed with the use of MEG and fMRI devices.

In the present study, we estimated the cortical networks were from high-resolution EEG recordings in a group of spinal cord injured patients and in a group of healthy subjects, during the preparation of a limb movement. Then, we use the Markov Clustering method to analyse the division of the network into community structures. The results indicate large differences between the injured patients and the healthy subjects. In particular, the networks of spinal cord injured patient exhibited a higher density of clusters. In the Alpha (7-12 Hz) frequency band, the two observed largest communities were mainly composed by the cingulate motor areas with the supplementary motor areas, and by the pre-motor areas with the right primary motor area of the foot. This functional separation could reflect the partial alteration in the primary motor areas because of the effects of the spinal cord injury.

In the present work, we estimated the functional networks in the frequency domain from a set of high-resolution EEG data in a group of healthy subjects during the showing of commercial spots within a neutral documentary. Then, we evaluated the differences in the cortical network associated with later remembered and not-remembered commercials by calculating the global- E(g) and local-efficiency E(l) indexes. During the visualization of the video-clips that will be forgotten (FRG), the cortical network exhibited high values of global- and local-efficiency, reflecting a small-world configuration. During the visualization of the video-clips that will be remembered (RMB), the same indexes appeared significantly lower. Such a difference seems not depending on the spectral content of the cortical activity. This result shows how the network communication efficiency would be affected by the presence of attentional and semantic processes that are behind a successful memory encoding in a lifelike situation.

In this study we estimated time-varying cortical connectivity patterns from a group of Spinal Cord Injured (SCI) patients during the attempt to move a paralyzed limb. This data were compared with the time-varying connectivity patterns estimated in a control group during the effective execution of the movement. Connectivity was estimated from high resolution EEG recordings with the use of realistic head modelling and the linear inverse estimation of the cortical activity. Time-varying PDC was obtained by the adaptive recursive fit of an MVAR model with time-dependent parameters, by means of a generalized recursive least-square (RLS) algorithm, taking into consideration a set of EEG epochs. Such estimator is able to follow rapid changes in the connectivity between cortical areas during an experimental task. The obtained experimental evidences support the conclusion that the SCI population involved a larger cortical network than those generated by the healthy subjects during the task performance. Such network differs for the involvement of the parietal cortices, which increases in strength near to the EMG onset.

The present work aims at analyzing the structure of cortical connectivity during the attempt to move a paralyzed limb by a group of spinal cord injured (SCI) patients. Connectivity patterns were obtained by means of the Directed Transfer Function applied to the cortical signals estimated from high resolution EEG recordings. Electrical activity were estimated in normals (Healthy) and SCI patients on twelve regions of interest (ROIs) coincident with Brodmann areas. Degree distributions showed the presence of few cortical regions with a lot of outgoing connections in all the cortical networks estimated irrespectively of the frequency band investigated. For both of the groups (SCI and Healthy), bilateral cingulate motor area (CMA) acts as hub transmitting information flows. The efficiency index, allowed to assert the ordered properties of such estimated cortical networks in both populations. The comparison of such estimated networks with those obtained from random networks, elicited significant differences (P < 0.05, Bonferroni-corrected for multiple comparisons). A statistical comparison (ANOVA) between SCI patients and healthy subjects showed a significant difference (P < 0.05) between the local efficiency of their respective networks. For three frequency bands (theta 4-7 Hz, alpha 8-12 Hz, and beta 13-29 Hz) the higher value observed in the spinal cord injured population entails a larger level of internal organization and fault tolerance. This fact suggests a sort of compensative mechanism as local response to the alteration in their MIF areas, which is probably due to the indirect effects of the spinal injury., ((copyright) 2007 Wiley-Liss, Inc.)

Assessment of brain connectivity among different brain areas during cognitive or motor tasks is a crucial problem in neuroscience today. Aim of this work is to use a neural mass model to assess the effect of various connectivity patterns in cortical electroencephalogram (EEG) power spectral density, and investigate the possibility to derive connectivity circuits from EEG data. To this end, a model of an individual region of interest (ROI) has been built as the parallel arrangement of three populations, each described as in Wendling et al. (Eur J Neurosci 15:1499-1508, 2002). Connectivity among ROIs includes three parameters, which specify the strength of connection in the different frequency bands. The following main steps have been followed: (1) we analyzed how the power spectral density (PSD) is significantly modified by the kind of coupling hypothesized among the ROIs; (2) with the model, and using an automatic fitting procedure, we looked for a simple connectivity circuit able to reproduce PSD of cortical EEG in three ROIs during a finger-movement task. The estimated parameters represent the strength of connections among the ROIs in the different frequency bands. Cortical EEGs were computed with an inverse propagation algorithm, starting from measurement performed with 96 electrodes on the scalp. The present study suggests that the model can be used as a simulation tool, able to mimic the effect of connectivity on EEG. Moreover, it can be used to look for simple connectivity circuits, able to explain the main features of observed cortical PSD. These results may open new prospectives in the use of neurophysiological models, instead of empirical models, to assess effective connectivity from neuroimaging information.

Static hemodynamic or neuroelectric images of brain regions activated during particular tasks do not convey the information of how these regions communicate to each other. Cortical connectivity estimation aims at describing these interactions as connectivity patterns which hold the direction and strength of the information flow between cortical areas. In this study, we attempted to estimate the causality between distributed cortical systems during a movement volition task in preparation for execution of simple movements by a group of normal healthy subjects and by a group of Spinal Cord Injured (SCI) patients. To estimate the causality between the spatial distributed patterns of cortical activity in the frequency domain, we applied a series of processing steps on the recorded EEG data. From the high-resolution EEG recordings we estimated the cortical waveforms for the regions of interest (ROIs), each representing a selected sensor group population. The solutions of the linear inverse problem returned a series of cortical waveforms for each ROI considered and for each trial analyzed. For each subject, the cortical waveforms were then subjected to Independent Component Analysis (ICA) pre-processing. The independent components obtained by the application of the ThinICA algorithm were further processed by a Partial Directed Coherence algorithm, in order to extract the causality between spatial cortical patterns of the estimated data. The source-target cortical dependencies found in the group of normal subjects were relatively similar in all frequency bands analyzed. For the normal subjects we observed a common source pattern in an ensemble of cortical areas including the right parietal and right lip primary motor areas and bilaterally the primary foot and posterior SMA areas. The target of this cortical network, in the Granger-sense of causality, was shown to be a smaller network composed mostly by the primary foot motor areas and the posterior SMA bilaterally. In the case of the SCI population, both the source and the target cortical patterns had larger sizes than in the normal population. The source cortical areas included always the primary foot and lip motor areas, often bilaterally. In addition, the right parietal area and the bilateral premotor area 6 were also involved. Again, the patterns remained substantially stable across the different frequency bands analyzed. The target cortical patterns observed in the SCI population had larger extensions when compared to the normal ones, since in most cases they involved the bilateral activation of the primary foot movement areas as well as the SMA, the primary lip areas and the parietal cortical areas.

Over the last 20 years, a body of techniques known as high resolution EEG has allowed precise estimation of cortical activity from non-invasive EEG measurements. The availability of cortical waveforms from non-invasive EEG recordings allows to have not only the level of activation within a single region of interest (ROI) during a particular task, but also to estimate the causal relationships among activities of several cortical regions. However, interpreting resulting connectivity patterns is still an open issue, due to the difficulty to provide an objective measure of their properties across different subjects or groups. A novel approach addressed to solve this difficulty consists in manipulating these functional brain networks as graph objects for which a large body of indexes and tools are available in literature and already tested for complex networks at different levels of scale (Social, WorldWide-Web and Proteomics). In the present work, we would like to show the suitability of such approach, showing results obtained comparing separately two groups of subjects during the same motor task and two different motor tasks performed by the same group. In the first experiment two groups of subjects (healthy and spinal cord injured patients) were compared when they moved and attempted to move simultaneously their right foot and lips, respectively. The contrast between the foot-lips movement and the simple foot movement was addressed in the second experiment for the population of the healthy subjects. For both the experiments, the main question is whether the "architecture" of the functional connectivity networks obtained could show properties that are different in the two groups or in the two tasks. All the functional connectivity networks gathered in the two experiments showed ordered properties and significant differences from "random" networks having the same characteristic sizes. The proposed approach, based on the use of indexes derived from graph theory, can apply to cerebral connectivity patterns estimated not only from the EEG signals but also from different brain imaging methods.

The aim of this work is to characterize quantitatively the performance of a body of techniques in the frequency domain for the estimation of cortical connectivity from high-resolution EEG recordings in different operative conditions commonly encountered in practice. Connectivity pattern estimators investigated are the Directed Transfer Function (DTF), its modification known as direct DTF (dDTF) and the Partial Directed Coherence (PDC). Predefined patterns of cortical connectivity were simulated and then retrieved by the application of the DTF, dDTF, and PDC methods. Signal-to-noise ratio (SNR) and length (LENGTH) of EEG epochs were studied as factors affecting the reconstruction of the imposed connectivity patterns. Reconstruction quality and error rate in estimated connectivity patterns were evaluated by means of some indexes of quality for the reconstructed connectivity pattern. The error functions were statistically analyzed with analysis of variance (ANOVA). The whole methodology was then applied to high-resolution EEG data recorded during the well-known Stroop paradigm. Simulations indicated that all three methods correctly estimated the simulated connectivity patterns under reasonable conditions. However, performance of the methods differed somewhat as a function of SNR and LENGTH factors. The methods were generally equivalent when applied to the Stroop data. In general, the amount of available EEG affected the accuracy of connectivity pattern estimations. Analysis of 27 s of nonconsecutive recordings with an SNR of 3 or more ensured that the connectivity pattern could be accurately recovered with an error below 7% for the PDC and 5% for the DTF. In conclusion, functional connectivity patterns of cortical activity can be effectively estimated under general conditions met in most EEG recordings by combining high-resolution EEG techniques, linear inverse estimation of the cortical activity, and frequency domain multivariate methods such as PDC, DTF, and dDTF.

In this work, a novel approach is proposed in order to capture relevant features related to the structure and organization of the functional brain networks estimated in the time-frequency domain. To achieve this, we used a cascade of computational tools able to estimate first the electrical activity of the cortical surface by using high resolution EEG techniques. Then, on the cortical signals from different regions of interests we estimated the time-varying functional connectivity patterns by means of the adaptive Partial Directed Coherence. Such time-varying connectivity estimation returns a series of causality patterns evolving during the examined task which can be summarized and interpreted with the aid of mathematical indexes based on the graph theory. The combination of all these methods is demonstrated on a set of high resolution EEG data recorded from a healthy subject performing a simple foot movement.

In this paper we propose the use of an adaptive multivariate approach to define time-varying multivariate estimators based on the Directed Transfer Function (DTF) and the Partial Directed Coherence (PDC). DTF and PDC are frequency-domain estimators that are able to describe interactions between cortical areas in terms of the concept of Granger causality. Time-varying DTF and PDC were obtained by the adaptive recursive fit of an MVAR model with time-dependent parameters, by means of a generalized recursive least-square (RLS) algorithm, taking into consideration a set of EEG epochs. Such estimators are able to follow rapid changes in the connectivity between cortical areas during an experimental task. We provide an application to the cortical estimations obtained from high resolution EEG data, recorded from a group of healthy subject during a combined foot-lips movement, and present the time-varying connectivity patterns resulting from the application of both DTF and PDC. Two different cortical networks were detected, one constant across the task and the other evolving during the preparation of the joint movement.

A major limitation of the approaches used in most of the studies performed so far for the characterization of the brain responses during social interaction is that only one of the participating brains is measured each time. The "interaction" between cooperating, competing or communicating brains is thus not measured directly, but inferred by independent observations aggregated by cognitive models and assumptions that link behavior and neural activation. In this paper, we use the simultaneous neuroelectric recording of several subjects engaged in cooperative games (EEG hyperscanning). This EEG hyperscanning allow us to observe and model directly the neural signature of human interactions in order to understand the cerebral processes generating and generated by social cooperation or competition. We used a paradigm called Prisoner's dilemma derived from the game theory. Results collected in a population of 22 subjects suggested that the most consistently activated structure in social interaction paradigms is the medial prefrontal cortex, which is found to be active in all the conflict situations analyzed. The role of the anterior cingulated cortex (ACC) assumes a main character being a discriminant factor for the "defect" attitude of the entire population examined. This observation is compatible with the role that the Theory of Mind assigns to the ACC.

Neural mass models have been used for many years to study the macroscopic dynamics of neural populations in a simple and computationally inexpensive way. In this paper, we modified a model proposed by Wendling et al. [Wendling F, Bartolomei F, Bellanger JJ, Chauvel P. Epileptic fast activity can be explained by a model of impaired GABAergic dendritic inhibition. Eur J Neurosci 2002;15:1499-508] to simulate EEG power spectral density (PSD) in some regions of interest (ROIs) during simple tasks (finger movement or working memory tests). The work consists of two subsequent stages: (1) in the first we evaluated the role of some model parameters (i.e., average gain of synapses and their time constants) in affecting power spectral density. This analysis confirmed the possibility to simulate various EEG rhythms (in the alpha, beta and gamma frequency ranges) by modifying just the time constants of the synapses. The position of the individual rhythms (i.e., the corresponding peaks in the PSD) can be finely tuned acting on the average gain of fast inhibitory synapses. This analysis suggested that a single neural mass model produces a unimodal spectrum, which can be finely adjusted, but cannot mimic the overall complexity of EEG in an entire cortical area. (2) Hence, in the second stage we built a model of a ROI by combining three neural mass models arranged in parallel. With this model, and using an automatic fitting procedure, we carefully reproduced the PSD of cortical EEG in several ROIs during finger movement, and their temporal changes during a working memory task, by estimating nine parameters. The estimated parameters represent the excitation of each population (mean value and variance of exogenous input noise) and the average gain of fast inhibitory synapses. Cortical EEGs were computed with an inverse propagation algorithm, starting from measurement performed with a high number of electrodes on the scalp (46-96). Results show that the proposed model is able to mimic PSD of cortical activity acting on a few parameters, which represent external activation and short-time synaptic changes. This information may be exploited to reach a quantitative summary of electrical activity in ROIs during a task, and to derive information on connectivity, starting from non-invasive EEG measurements.

The aim of this paper is to test a technique called partial directed coherence (PDC) and its modification (squared PDC; sPDC) for the estimation of human cortical connectivity by means of simulation study, in which both PDC and sPDC were studied by analysis of variance. The statistical analysis performed returned that both PDC and sPDC are able to estimate correctly the imposed connectivity patterns when data exhibit a signal-to-noise ratio of at least 3 and a length of at least 27 s of nonconsecutive recordings at 250 Hz of sampling rate, equivalent, more generally, to 6750 data samples.

The Directed Transfer Function (DTF) and the Partial Directed Coherence (PDC) are frequency-domain estimators, based on the multivariate autoregressive modelling (MVAR) of time series, that are able to describe interactions between cortical areas in terms of the concept of Granger causality. However, the classical estimation of these methods requires the stationary of the signals. In this way, transient pathways of information transfer remains hidden. The objective of this study is to test a time-varying multivariate method for the estimation of rapidly changing connectivity relationships between cortical areas of the human brain, based on DTF/PDC and on the use of adaptive MVAR modelling (AMVAR). This approach will allow the observation of transient influences between the cortical areas during the execution of a task. Time-varying DTF and PDC were obtained by the adaptive recursive fit of an MVAR model with time-dependent parameters, by means of a generalized recursive least-square (RLS) algorithm, taking into consideration a set of EEG epochs. Simulations were performed under different levels of Signal to Noise Ratio (SNR), number of trials (TRIALS) and frequency bands (BAND), and of different values of the RLS adaptation factor adopted (factor C). The results indicated that time-varying DTF and PDC are able to estimate correctly the imposed connectivity patterns under reasonable operative conditions of SNR ad number of trials. Moreover, the capability of follow the rapid changes in connectivity is highly increased by the number of trials at disposal, and by the right choice of the value adopted for the adaptation factor C. The results of the simulation study indicate that DTF and PDC computed on adaptive MVAR can be effectively used to estimate time-varying patterns of functional connectivity between cortical activations, under general conditions met in practical EEG recordings.

Assessment of brain connectivity among different brain areas during cognitive or motor tasks is a crucial problem in neuroscience today. Aim of this work is to use a neural mass model to assess the effect of various connectivity patterns in the power spectral density (PSD) of cortical EEG, and investigate the possibility to derive connectivity circuits from real EEG data. To this end, a model of an individual region of interest (ROI) has been built as the parallel arrangement of three populations. The present study suggests that the model can be used as a simulation tool, able to produce reliable intracortical EEG signals. Moreover, it can be used to look for simple connectivity circuits, able to explain the main features of observed cortical PSD. These results may open new prospective in the use of neurophysiological models, instead of empirical models, to assess effective connectivity from neuroimaging information.

Today, the concept of brain connectivity plays a central role in the neuroscience. While functional connectivity is defined as the temporal coherence between the activities of different brain areas, the effective connectivity is defined as the simplest brain circuit that would produce the same temporal relationship as observed experimentally between cortical sites. The most used method to estimate effective connectivity in neuroscience is the structural equation modeling (SEM), typically used on data related to the brain hemodynamic behavior. However, the use of hemodynamic measures limits the temporal resolution on which the brain process can be followed. The present research proposes the use of the SEM approach on the cortical waveforms estimated from the high-resolution EEG data, which exhibits a good spatial resolution and a higher temporal resolution than hemodynamic measures. We performed a simulation study, in which different main factors were systematically manipulated in the generation of test signals, and the errors in the estimated connectivity were evaluated by the analysis of variance (ANOVA). Such factors were the signal-to-noise ratio and the duration of the simulated cortical activity. Since SEM technique is based on the use of a model formulated on the basis of anatomical and physiological constraints, different experimental conditions were analyzed, in order to evaluate the effect of errors made in the a priori model formulation on its performances. The feasibility of the proposed approach has been shown in a human study using high-resolution EEG recordings related to finger tapping movements.

Objective: To test a technique called Directed Transfer Function (DTF) for the estimation of human cortical connectivity, by means of simulation study and human study, using high resolution EEG recordings related to finger movements., Methods: The method of the Directed Transfer Function (DTF) is a frequency-domain approach, based on a multivariate autoregressive modeling of time series and on the concept of Granger causality. Since the spreading of the potential from the cortex to the sensors makes it difficult to infer the relation between the spatial patterns on the sensor space and those on the cortical sites, we propose the use of the DTF method on cortical signals estimated from high resolution EEG recordings, which exhibit a higher spatial resolution than conventional cerebral electromagnetic measures. The simulation study was followed by an analysis of variance (ANOVA) of the results obtained for different levels of Signal to Noise Ratio (SNR) and temporal length, as they have been systematically imposed on simulated signals. The whole methodology was then applied to high resolution EEG data recorded during a visually paced finger movement., Results: The statistical analysis performed returns that during simulations, DTF is able to estimate correctly the imposed connectivity patterns under reasonable operative conditions, i.e. when data exhibit a SNR of at least 3 and a length of at least 75 s of non-consecutive recordings at 64 Hz of sampling rate, equivalent, more generally, to 4800 data samples., Conclusions: Functional connectivity patterns of cortical activity can be effectively estimated under general conditions met in any practical EEG recordings, by combining high resolution EEG techniques, linear inverse estimation and the DTF method., Significance: The estimation of cortical connectivity can be performed not only with hemodynamic measurements, by using functional MRI recordings, but also with modern EEG recordings treated with advanced computational techniques.

Nowadays, several types of brain imaging device are available to provide images of the functional activity of the cerebral cortex based on hemodynamic, metabolic, or electromagnetic measurements. However, static images of brain regions activated during particular tasks do not convey the information of how these regions communicate with each other. In this study, advanced methods for the estimation of cortical connectivity from combined high-resolution electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) data are presented. These methods include a subject's multicompartment head model (scalp, skull, dura mater, cortex) constructed from individual magnetic resonance images, multidipole source model, and regularized linear inverse source estimates of cortical current density. Determination of the priors in the resolution of the linear inverse problem was performed with the use of information from the hemodynamic responses of the cortical areas as revealed by block-designed (strength of activated voxels) fMRI. We estimate functional cortical connectivity by computing the directed transfer function (DTF) on the estimated cortical current density waveforms in regions of interest (ROIs) on the modeled cortical mantle. The proposed method was able to unveil the direction of the information flow between the cortical regions of interest, as it is directional in nature. Furthermore, this method allows to detect changes in the time course of information flow between cortical regions in different frequency bands. The reliability of these techniques was further demonstrated by elaboration of high-resolution EEG and fMRI signals collected during visually triggered finger movements in four healthy subjects. Connectivity patterns estimated for this task reveal an involvement of right parietal and bilateral premotor and prefrontal cortical areas. This cortical region involvement resembles that revealed in previous studies where visually triggered finger movements were analyzed with the use of separate EEG or fMRI measurements.

Different brain imaging devices are presently available to provide images of the human functional cortical activity, based on hemodynamic, metabolic or electromagnetic measurements. However, static images of brain regions activated during particular tasks do not convey the information of how these regions are interconnected. The concept of brain connectivity plays a central role in the neuroscience, and different definitions of connectivity, functional and effective, have been adopted in literature. While the functional connectivity is defined as the temporal coherence among the activities of different brain areas, the effective connectivity is defined as the simplest brain circuit that would produce the same temporal relationship as observed experimentally among cortical sites. The structural equation modeling (SEM) is the most used method to estimate effective connectivity in neuroscience, and its typical application is on data related to brain hemodynamic behavior tested by functional magnetic resonance imaging (fMRI), whereas the directed transfer function (DTF) method is a frequency-domain approach based on both a multivariate autoregressive (MVAR) modeling of time series and on the concept of Granger causality. This study presents advanced methods for the estimation of cortical connectivity by applying SEM and DTF on the cortical signals estimated from high-resolution electroencephalography (EEG) recordings, since these signals exhibit a higher spatial resolution than conventional cerebral electromagnetic measures. To estimate correctly the cortical signals, we used a subject's multicompartment head model (scalp, skull, dura mater, cortex) constructed from individual MRI, a distributed source model and a regularized linear inverse source estimates of cortical current density. Before the application of SEM and DTF methodology to the cortical waveforms estimated from high-resolution EEG data, we performed a simulation study, in which different main factors (signal-to-noise ratio, SNR, and simulated cortical activity duration, LENGTH) were systematically manipulated in the generation of test signals, and the errors in the estimated connectivity were evaluated by the analysis of variance (ANOVA). The statistical analysis returned that during simulations, both SEM and DTF estimators were able to correctly estimate the imposed connectivity patterns under reasonable operative conditions, that is, when data exhibit an SNR of at least 3 and a LENGTH of at least 75 s of nonconsecutive EEG recordings at 64 Hz of sampling rate. Hence, effective and functional connectivity patterns of cortical activity can be effectively estimated under general conditions met in any practical EEG recordings, by combining high-resolution EEG techniques and linear inverse estimation with SEM or DTF methods. We conclude that the estimation of cortical connectivity can be performed not only with hemodynamic measurements, but also with EEG signals treated with advanced computational techniques.

The authors delineated the time evolution of alpha event-related desynchronization over human frontal, parietal, and primary sensorimotor areas during the expectancy of a go/no-go task. The main issue under investigation was whether anticipatory processes impinged upon cortical areas in sequential or parallel mode. Compared with the control condition, in the experimental condition there was an Alpha 1 desynchronization over the central midline, an Alpha 2 desynchronization increasing over primary sensorimotor areas, and an Alpha 3 desynchronization increasing in parallel over bilateral primary sensorimotor areas. These processes had different temporal features. Results disclose an anticipatory activity of central midline areas and primary sensorimotor areas in both parallel and sequential modes. This reflects an adaptive, energy-consuming strategy rather than an economic waiting for the go stimulus.

The intent of this electroencephalography study was to investigate the competition between cortical nociceptive and cognitive-motor processes preceding sensorimotor interactions. Sensorimotor expectancy processes to painful stimulation and motor go/no-go demands were indexed over primary sensorimotor and midline cortical areas by contingent negative variation (CNV). Before the sensorimotor interaction, CNV was observed over midline posterior and bilateral central areas. Early expectancy of painful stimulation and the go/no-go task induced an evident midline posterior CNV. During the late expectancy period. CNV extended to the right central area contralateral to the stimulation. These findings suggest a sequential activation of midline posterior and primary sensorimotor areas contralateral to the painful stimulation as a reflection of the enhanced nociceptive processes preceding painful sensorimotor interactions., (Copyright 2004 APA.)

In this chapter, advanced methods for the modeling of human cortical activity from combined high-resolution electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) data are reviewed. These methods include a subject's multicompartment head model (scalp, skull, dura mater, cortex) constructed from magnetic resonance images, multidipole source model, and regularized linear inverse source estimates. Determination of the priors in the resolution of the linear inverse problem was performed with the use of information from the hemodynamic responses of the cortical areas as revealed by block-designed (strength of activated voxels) and event-related (coupling of activated voxels) fMRI. As an example, these methods were applied to EEG (128 electrodes) and fMRI data, which were recorded in separate sessions while normal subjects executed voluntary right one-digit movements.

Objectives: EEG scalp potential distributions recorded in humans are affected by low spatial resolution and by the dependence on the electrical reference used. High resolution EEG technologies are available to drastically increase the spatial resolution of the raw EEG. Such technologies include the computation of surface Laplacian (SL) of the recorded potentials, as well as the use of realistic head models to estimate the cortical sources via linear inverse procedure (low resolution brain electromagnetic tomography, LORETA). However, these deblurring procedures are generally used in conjunction with EEG recordings with 64-128 scalp electrodes and with realistic head models obtained via sequential magnetic resonance images (MRIs) of the subjects. Such recording setup it is not often available in the clinical context, due to both the unavailability of these technologies and the scarce compliance of the patients with them. In this study we addressed the use of SL and LORETA deblurring techniques to analyze data from a standard 10-20 system (19 electrodes) in a group of Alzheimer disease (AD) patients., Methods: EEG data related to unilateral finger movements were gathered from 10 patients affected by AD. SL and LORETA techniques were applied for source estimation of EEG data. The use of MRIs for the construction of head models was avoided by using the quasi-realistic head model of the Brain Imaging Neurology Institute of Montreal., Results: A similar cortical activity estimated by the SL and LORETA techniques was observed during an identical time period of the acquired EEG data in the examined population., Conclusions: The results of the present study suggest that both SL and LORETA approaches can be usefully applied in the clinical context, by using quasi-realistic head modeling and a standard 10-20 system as electrode montage (19 electrodes). These results represent a reciprocal cross-validation of the two mathematically independent techniques in a clinical environment.

High-resolution event-related potentials (ERPs) were used to model the hemispherical representation of the transient cortical responses relating to the observation of movement during execution (right or left aimless finger extension). Subjects were seated in front of the observed person and looked at both their own and the observer's hand to receive similar visual feedback during the two conditions. In a visual control condition, a diode light moved at the observed person's hand. A first potential accompanying the movement execution peaked at about +110 msec over the contralateral somatomotor areas. It was followed by a potential (P300) peaking at about +350 msec over the central midline. In contrast, the potentials accompanying the movement observation peaked later over parietal-occipital other than somatomotor areas (N200 peak, +200 msec; P300 peak, +400 msec). Notably, the N200 was maximum in left parietal area whereas the P300 was maximum in right parietal area regardless the side of the movement. They markedly differed by the potentials following the displacement of the diode light. These results suggest a rapid time evolution (approximately 200-400 msec) of the cortical responses characterizing the observation of aimless movements (as opposite to grasping or handling). The execution of these movements would mainly involve somatomotor cortical responses and would be scarcely founded on the visual feedback. In contrast, the observation of the same movements carried out by others would require dynamical responses of somatomotor and parietal-occipital areas (especially of the right hemisphere), possibly for a stringent visuospatial analysis of the motor event., (Copyright 2003 Wiley-Liss, Inc.)

Previous simulation studies have stressed the importance of the use of fMRI priors in the estimation of cortical current density. However, no systematic variations of signal-to-noise ratio (SNR) and number of electrodes were explicitly taken into account in the estimation process. In this simulation study we considered the utility of including information as estimated from fMRI. This was done by using as the dependent variable both the correlation coefficient and the relative error between the imposed and the estimated waveforms at the level of cortical region of interests (ROI). A realistic head and cortical surface model was used. Factors used in the simulations were the different values of SNR of the scalp-generated data, the different inverse operators used to estimated the cortical source activity, the strengths of the fMRI priors in the fMRI-based inverse operators, and the number of scalp electrodes used in the analysis. Analysis of variance results suggested that all the considered factors significantly afflict the correlation and the relative error between the estimated and the simulated cortical activity. For the ROIs analyzed with simulated fMRI hot spots, it was observed that the best estimation of cortical source currents was performed with the inverse operators that used fMRI information. When the ROIs analyzed do not present fMRI hot spots, both standard (i.e., minimum norm) and fMRI-based inverse operators returned statistically equivalent correlation and relative error values.

In the present high-resolution electroencephalographic (EEG) study, we computed event-related desynchronization and synchronization (ERD/ERS) of alpha (about 10 Hz) and beta (about 20 Hz) rhythms in association with the execution (with visual feedback) and observation of brisk unilateral right and left aimless finger movements. A first scope was to test the topographical "functional equivalence" of cortical rhythmicity related to movement execution and observation, which would represent an ideal cortical observation/execution matching system. A second scope was to evaluate the hypothesis of a left or right hemisphere prevalence of the cortical rhythmicity related to the movement observation compared to the movement execution. EEG (128 electrodes) was recorded in 10 healthy right-handed volunteers. Surface Laplacian estimation spatially enhanced EEG data over a MRI-constraint head model. Under both conditions, ERD peaked during the movement execution or observation and was replaced by a ERS "rebound" or "recovery," which peaked during the postevent period. Topographical results are in favor of a "functional equivalence" (i.e., similar ERD/ERS values in magnitude and timing) of alpha and beta rhythmicity in central scalp regions overlying premotor/primary sensorimotor cortex. On the contrary, the functional equivalence of alpha rhythmicity was negligible (i.e., different ERD/ERS values in magnitude and timing) in parietal-occipital scalp regions overlying posterior parietal and parieto-occipital cortex, which could be the neural substrate to distinguish among the own motor intensions and others' aimless movements (i.e., visuomotor transformation integrated with sensorimotor, postural, and kinematics representations). Finally, the pattern of hemispherical cortical rhythmicity did not support a "simple concentration" of movement observation functions in the left or right hemisphere.

The main aims of this study were 1) a fine spatial analysis of electroencephalographic (EEG) oscillations after galvanic painful stimulation (nonpainful stimulation as a reference) and 2) a comparative evaluation of phase- and nonphase-locked component of these EEG oscillations. Preliminary surface Laplacian transformation of EEG data (31 channels) reduced head volume conductor effects. EEG phase values were computed by FFT analysis and the statistical evaluation of these values was performed by Rayleigh test (P < 0.05). About 50% of the EEG single trials presented statistically the same FFT phase value of the evoked EEG oscillations (phase-locked single trials), indicating a preponderant phase-locked compared to nonphase-locked component. The remaining single trials showed random FFT phase values (nonphase-locked single trials), indicating a preponderant nonphase-locked compared to phase-locked component. Compared to nonpainful stimulation, painful stimulation increased phase-locked theta to gamma band responses in the contralateral hemisphere and decreased the phase-locked beta band response in the ipsilateral hemisphere. Furthermore, nonphase-locked alpha band response decreased in the ipsilateral fronto-central area. In conclusion, both decreased and increased EEG oscillatory responses to galvanic painful stimulation would occur in parallel in different cortical regions and in the phase- and nonphase-locked EEG data sets. This enriches the actual debate on the mapping of event-related oscillatory activity of human brain., (Copyright 2002 Wiley-Liss, Inc.)

Objectives: In this paper, we employed advanced methods for the modeling of human cortical activity related to voluntary right one-digit movements from combined high-resolution electroencepholography (EEG) and functional magnetic resonance imaging (fMRI)., Methods: Multimodal integration between EEG and fMRI data was performed by using realistic head models, a large number of scalp electrodes (128) and the estimation of current density strengths by linear inverse estimation., Results: Increasing of spatial details of the estimated cortical density distributions has been detected by using the proposed integration method with respect to the estimation using EEG data alone., Conclusion: The proposed method of multimodal EEG-fMRI data is useful to increase spatial resolution of movement-related potentials and can also be applied to other kinds of event-related potentials.

Cortical sources of human movement-related potentials (i.e. unilateral finger extension) were modeled using functional magnetic resonance imaging (fMR) data as a constraint of a linear inverse source estimation from highly sampled (128 channels) EEG data. Remarkably, this estimation was performed within realistic subject's MR-constructed head models by boundary element techniques. An appropriate figure of merit served to set the optimal amount of fMR constraints. With respect to standard linear inverse source estimates, fMR-constrained ones presented increased spatial detail and provided a more reliable timing of activation in bilateral sensorimotor cortical regions of interest.

Electroencephalograph (EEG)-based brain-computer interfaces (BCI's) require on-line detection of mental states from spontaneous EEG signals. In this framework, surface Laplacian (SL) transformation of EEG signals has proved to improve the recognition scores of imagined motor activity. The results we obtained in the first year of an European project named adaptive brain interfaces (ABI) suggest that: 1) the detection of mental imagined activity can be obtained by using the signal space projection (SSP) method as a classifier and 2) a particular type of electrodes can be used in such a BCI device, reconciling the benefits of SL waveforms and the need for the use of few electrodes. Recognition of mental activity was attempted on both raw and SL-transformed EEG data from five healthy people performing two mental tasks, namely imagined right and left hand movements.

Human short-latency cortical responses to median nerve stimulation were investigated with a new high resolution electroencephalography technology that markedly enhanced spatial details of somatosensory-evoked potentials (SEPs). Maximum amplitude potentials were estimated over contralateral and/or frontal-mesial scalp regions about 20, 22, 24, 26, 30, 32 and 45 ms following the stimulation. Frontal-lateral P20-N24-N30-P45 and parietal-lateral N20-P24-P30-N45 showed dipolar patterns, whereas frontal-mesial N24-N30-P45 and central-lateral P22-N26-N32-P45 presented no clearcut dipole counterpart. Plausibly, the spatially enhanced frontal-parietal SEP components were generated (tangential dipoles) within the lateral central sulcus cortex, and anticipated the central-lateral and frontal-mesial components generated (radial dipoles) from the crown of the pre- and/or post-central gyri and the supplementary motor area, respectively.