Changes in alpha rhythm asymmetry during learning of verbal and visuospatial tasks.

Alpha activity in the two cortical hemispheres was examined during learning and overlearning of a verbal and a visuospatial task. Compared to a pretask baseline condition, the amount of alpha activity during the tasks was decreased. An increase in alpha activity was found from learning to overlearning in both tasks. A greater amount of alpha was found in the right hemisphere for both tasks. In addition, no difference in the degree of alpha asymmetry was found between the tasks. This was discussed in light of prior contrary findings. A significant positive correlation was found between change in learning rate and change in alpha asymmetry during the verbal task. An interpretation of this finding was offered in terms of the level of hemispheric activation reflecting the degree of task engagement.     

Equivalent dipole estimation of spontaneous EEG alpha activity: two-moving dipole approach

A method of estimating equivalent moving and fixed dipoles from the scalp-recorded EEG alpha waves, with the realistic geometry of the head taken into account, is presented. Twenty-one silver electrodes were used to collect spontaneous EEG alpha waves on the scale. Four models, the single-moving dipole model, the single-fixed dipole model, the two-moving dipole model and the two-fixed dipole model were applied to approximate the EEG alpha field on the scalp. The algorithm, based on a least-squares fit for estimating the moving and the fixed dipoles by using a realistically shaped head model, is described. The numerical accuracy of the algorithm is also evaluated by a computer simulation. It is found that the spontaneous EEG alpha activity observed on the scalp can be represented by two equivalent moving dipoles, simultaneously located separately in the occipital regions of the right and the left hemisphere, at a depth of 4–6 cm beneath the scalp, with a goodness-of-fit of up to 97 per cent for all subjects examined. The excellent fit of the two-moving dipole model to the EEG human alpha activity is also compared with the single-dipole fit.     

Long-term enhanced desynchronization of the alpha rhythm following tetanic stimulation of human visual cortex

It had been shown previously that a photic tetanus induces LTP-like changes in the visual cortex, as indexed by an enhancement of the N1b component of the visual evoked potential, recorded non-invasively by electroencephalography. This potentiation was shown to last over 1 h. In the present study, the effect of a photic tetanus on oscillatory activity is investigated. EEGs were collected from eight healthy subjects in three conditions while visual checkerboards were displayed. Following baseline presentations in two conditions a lateralized visual tetanus was given, either to the left or right visual field, and in a third condition no tetanus was given. This was followed by a return to baseline presentations, both immediately after the tetanus/control block, and 1 h later. Enhanced event-related desynchronization (ERD) of the alpha rhythm lasting 1 h was seen following the photic tetanus over occipital electrodes. Because ERD of the alpha rhythm is thought to represent active cortex, these results suggest that the visual tetanus induces long-lasting cortical changes, with stronger neuronal assemblies and increased neuronal output.     

A right‐ear bias of auditory selective attention is evident in alpha oscillations

Auditory selective attention makes it possible to pick out one speech stream that is embedded in a multispeaker environment. We adapted a cued dichotic listening task to examine suppression of a speech stream lateralized to the nonattended ear, and to evaluate the effects of attention on the right ear's well‐known advantage in the perception of linguistic stimuli. After being cued to attend to input from either their left or right ear, participants heard two different four‐word streams presented simultaneously to the separate ears. Following each dichotic presentation, participants judged whether a spoken probe word had been in the attended ear's stream. We used EEG signals to track participants' spatial lateralization of auditory attention, which is marked by interhemispheric differences in EEG alpha (8–14 Hz) power. A right‐ear advantage (REA) was evident in faster response times and greater sensitivity in distinguishing attended from unattended words. Consistent with the REA, we found strongest parietal and right frontotemporal alpha modulation during the attend‐right condition. These findings provide evidence for a link between selective attention and the REA during directed dichotic listening.     

Detection of alpha electro-encephalogram onset following eye closure using four location-based techniques

Detection of alpha activity in the electro-encephalogram (EEG) has been used extensively in neurophysiological studies. Previously applied alpha parameterisation techniques, which utilise the amplitude information from a pair of differential electrodes, are often susceptible to interference from artifact signals. This is an issue if the purpose of detecting the change in alpha wave synchronisation is the basis of an environmental control system (ECS). An alternative approach to alpha activity detection is proposed that utilises the information from an array of electrodes on the scalp to estimate the apparent location of alpha activity in the brain. Four methods are described that successfully detect the onset of alpha EEG increase following eye closure by monitoring the apparent location of alpha activity in the head. The methods use Bartlett beamforming, a four-sphere anatomical head model, the MUSIC algorithm and a new ‘power vector’ technique. Of the methods described, the power vector technique is found to be the most successful. The power vector technique detects the alpha increase associated with eye closure in times that are, on average, 33% lower than previously applied alpha detection methods.     

Neuronal generators of posterior EEG alpha reflect individual differences in prioritizing personal spirituality

Prominent posterior EEG alpha is associated with depression and clinical response to antidepressants. Given that religious belief was protective against depression in a longitudinal study of familial risk, we hypothesized that individuals who differed by strength of spiritual beliefs might also differ in EEG alpha. Clinical evaluations and self-reports of the importance of religion or spirituality (R/S) were obtained from 52 participants, and again at 10-y followup when EEG was measured. EEG alpha was quantified using frequency PCA of current source densities (CSD-fPCA). Participants who rated R/S as highly important at initial assessment showed greater alpha compared to those who did not. Those who rated R/S important in both sessions showed greater alpha than those who changed their ratings. EEG differences were particularly well-defined for participants with lifetime depression. Findings extend the view of alpha as a marker for affective processes, suggesting an association with the ontogenesis of spirituality.     

Lateral inhibition in neural networks and the shape of EEG alpha rhythm waves

Baseline EEG traces were recorded from the right occipital area with the eyes closed in 15 subjects. Rhythmic stimulation with bright, diffuse flashes of light was applied through the closed eyelids at specific points in the alpha-wave phase (trigger photic stimulation). Ten versions of stimulation were used, with application at different phases for 1 min at each phase. Responses occurring in conditions of stimulation in different phases were compared using the mean alpha wave amplitude in each version and the wave shape in terms of the level of asymmetry of the leading and trailing fronts. When flashes were delivered at the middle of the descending front of the potential (positivization), changes in wave shape were most marked and were very different from those seen in the other nine stimulation phases. This effect was most marked in subjects with low-amplitude alpha rhythms. These results suggest that the individual level of the alpha rhythm in a given subject and the dynamics of the wave shape can be explained by the characteristics of the structure of the system of recurrent-lateral inhibition. Translated from Rossiiskii Fiziologicheskii Zhurnal imeni I. M. Sechenova, Vol. 94, No. 2, pp. 152–162, February, 2008.     

Interhemispheric changes in alpha rhythm related to time perception

Each cerebral hemisphere processes environmental information in a different but complementary manner. Structures located in the left hemisphere are assumed to participate in symbolic-logic thinking. Time perception may be considered among such thinking processes. The present study evaluates bilateral occipito-central EEG activity in healthy, right-handed subjects which was produced while they performed a visuomotor monitoring task. The task consisted of two stages. The first stage involved the subject's learning a fixed time interval (10 sec) and the measurement of their reaction time. Subjects responded to an isolated light stimulus by pressing a button with the dominant hand. In the second stage, the subjects accuracy in estimating interval-length was evaluated. Two forms of EEG analysis were used, frequency and alpha ratio, each of which was measured both prior to and subsequent to the motor response. A reversal group was used to carry out a complementary test. Subjects responded in the first block of experiments with the non-dominant (left) hand and with the dominant hand in the second. Results showed that left hemisphere activity was continuous during the interval-learning stage and with optimal reaction times and remained continuous when estimation values approximated the real interval. In addition, in optimal reaction time and near to optimal time estimation responses, the left side showed lower frequency and alpha ratio than did the right. Finally a progressive enhancement in both parameters from the right hemisphere was related to deterioration in test performance. Results from the reversal group did not differ from those of the first group. As evaluated by gross measurements of the EEG, a predominant participation of the left hemisphere in time processing is concluded.     

Partitioning of deep versus superficial intracranial sources using current source densities is not valid

Summary An analytic method has recently been proposed for partitioning scalp-recorded EEG and evoked potential (EP) data into parts arising from deep (i.e., subcortical) vs. superficial (i.e., cortical) sources. The method is based on the observation that the current source density (CSD) is selectively sensitive to electrical activity arising from superficial sources, and the conjecture that the residual potential which remains after subtracting the CSD from the scalp potentials, represents activity from deep sources. We investigated the validity of this procedure by simulating scalp potential data for superficial and deep dipole sources with known locations and orientations. Our single-dipole simulations demonstrated that, when the actual location of the source was superficial, the partitioning procedure erroneously attributed a sizeable proportion of the total topographic variance to the activity of deeper sources. This produced a consistent bias in the simulations with two dipoles, when both superficial and deep sources were present. In such cases, the relative contribution of the deeper source was consistently overestimated, and the scalp topography of the deep source activity was profoundly misrepresented by the residual which results from subtracting the CSD from the scalp potential. We conclude that the proposed method for partitioning EEG and EP data into components arising from deep vs. superficial intracranial sources is not valid.Acknowledgements: This work was supported by general medical research funds from the Department of Veterans Affairs, by a VA psychiatry research training award to Dr. Turetsky and by NIDCD grant #R01 DC00328.     

Dipole Source Analysis May Differentiate Benign Focal Epilepsy of Childhood with Occipital Paroxysms from Symptomatic Occipital Lobe Epilepsy

The aim of the study was to distinguish Benign Focal Epilepsy of Childhood with Occipital Paroxysms (BEOP) from its symptomatic counterpart on the basis of the location of the sources of the interictal EEG spikes. Patients were classified into two groups: idiopathic BEOP and symptomatic occipital lobe epilepsy. Source analysis of the averaged occipital spikes was performed using a homogeneously conducting sphere as the volume conductor model. Results showed a statistically significant difference in the eccentricity, i.e., the distance of the occipital spike focus from the centre of the head. The dipole sources of the occipital spikes in the BEOP group were found to be located more superficially than in the symptomatic group, corresponding in six of the nine cases with a source position estimated to be within the cortical layer just below the skull. The eccentricity of the symptomatic occipital spikes suggests a location deeper than the cortical layer. The results were validated in two patients from the symptomatic group. In one patient the estimated deeper dipole source location corresponded with a deeper location of spike activity observed during ECoG; in the other patient's ECoG, spike activity was observed superficially but over an extended area. The discrepancy between estimated and real location may be explained by the method of dipole source analysis used. It is concluded that the finding of a superficial dipole source location of the occipital spikes provides an indication for the diagnosis BEOP (sensitivity: 67%; specificity: 74%).     

Finite-element model of the human head: scalp potentials due to dipole sources

Three-dimensional finite-element models provide a method to study the relationship between human scalp potentials and neural current sources inside the brain. A new formulation of dipole-like current sources is developed here. Finiteelement analyses based on this formulation are carried out for both a threeconcentric-spheres model and a human-head model. Differences in calculated scalp potentials between these two models are studied in the context of the forward and inverse problems in EEG. The effects of the eye orbit structure on surface potential distribution are also studied.     

Mapping EEG-potentials on the surface of the brain: A strategy for uncovering cortical sources

Summary This paper describes a uniform method for calculating the interpolation of scalp EEG potential distribution, the current source density (CSD), the cortical potential distribution (cortical mapping) and the CSD of the cortical potential distribution. It will be shown that interpolation and deblurring methods such as CSD or cortical mapping are not independent of the inverse problem in potential theory. Not only the resolution but also the accuracy of these techniques, especially those of deblurring, depend greatly on the spatial sampling rate (i.e., the number of electrodes). Using examples from simulated and real (64 channels) data it can be shown that the application of more than 100 EEG channels is not only favourable but necessary to guarantee a reasonable accuracy in the calculations of CSD or cortical mapping. Likewise, it can be shown that using more than 250 electrodes does not improve the resolution.This study was supported by grants from the Deutsche Forschungsgemeinschaft to Thomas Elbert and Brigitte Rockstroh. The authors would like to thank Dr. Dorothy Charbonnier.     

Confidence Interval of Single Dipole Locations Based on EEG Data

Noise in EEG and MEG measurements leads to inaccurate localizations of the sources. A confidence volume is used to describe the amount of localization error. Previous methods to estimate the confidence volume proved insufficient. Thus a new procedure was introduced and compared with previous ones. As one procedure, Monte Carlo simulations (MCS) were performed. The confidence volume was also estimated using two methods with different assumptions about a linear transfer function between source location and the distribution of the potential. One method used variable (LVM) and the other fixed dipole orientations (LFM). Finally, the confidence volume was estimated through a procedure in which there was no linearization of the transfer function. This procedure scans the confidence volume by varying the dipole location in multiple directions. Confidence volumes were calculated for simulated distributions of the electrical potential and for experimental data including somatosensory evoked responses to stimulation of lower lip, thumb, and little finger. Results from simulated data indicated that confidence volumes calculated with the MCS method were largest, and those calculated with the LFM method were smallest. For dipole locations close to the brain surface, the confidence volume was smaller than for a central deeper source. An increase in electrode density resulted in smaller confidence volumes. When the noise was correlated, only the method using the MCS produced acceptable results. Since the noise in experimental data is highly correlated, only the MCS method would appear to be useful in estimating the size of the confidence volume of the dipole locations. Thus, using real data with the MCS method, we easily distinguished separate and distinct representations of the thumb, little finger, and lower lip in the somatosensory cortex (SI). It was concluded that adequate estimation of confidence volumes is useful for localizing neural activity. On a practical level, this information can be used prior to an experiment for determining the conditions necessary to distinguish between different dipole sources, including the required signal to noise ratio and the minimum electrode density.     

Bilateral alpha distribution and anatomic brain asymmetries

Summary The parieto-occipital (PO) index of the VEP alpha-afterdischarge (AD) was assessed bilaterally in normal subjects in three successive sessions separated by one-three weeks with two within-session retests. Correlations between PO values and asymmetries of occipital and parietal width, mastoid area, basal angle, cephalic, and plagiocephaly indices were established in simple correlation and stepwise multiple regression analysis. Occipital width, notably in the right hemisphere correlated reliably with the PO index in relaxed wakefulness and the vigilance task. The population having normal and reversed occipital brain width is predicted to have different parieto-occipital AD distribution. Some errors of lateral EEG measurements introduced by slant cranial deformity are discussed in order to call attention to the fact that anatomy-related lateral EEG changes may masquerade as cognition-related asymmetries.The authors gratefully acknowledge the expert technical assistance of D.Shaefer, N.Schul, and D.Zuker. R.Rosenfield and A.Lefler assisted in 3-D reconstruction of images.     

Interpretation of high-resolution current source density profiles: a simulation of sublaminar contributions to the visual evoked potential

Current source density (CSD) analysis provides an index of the location, direction, and density of transmembrane currents that arise with synchronous activation of neural tissue and that generate an evoked potential profile in the extracellular medium. In neocortex and other laminated structures, a simplified, one-dimensional CSD analysis can be computed by differentiation of voltages sampled at discrete points in a linear array. One-dimensional CSD analysis is a practical and accurate method for defining both regional activity patterns and neural generators of surface-recorded evoked and event-related potentials. In computing the CSD, common practices of differentiating across spatial grids of 200 m or more and use of spatial smoothing routines help to reduce noise, but severely limit the spatial resolution available to the analysis. High-resolution CSD procedures (i.e., 3 point differentiation using a spatial grid of 100 m or less) are more suited to identification of processes within individual cortical laminae or sublaminae, but can magnify the contributions of computational artifacts. Despite the inclusion of independent indices of cellular activity (e.g., multiunit activity), both high- and lowresolution analyses may indicate current source and sink configurations for which there is more than one plausible physiological interpretation. In the present study we examined the resolving capacity and pitfalls of common CSD procedures using simulated ensembles of current dipoles. These were positioned and oriented to model the depolarization of lamina 4C stellate cells and thalamocortical afferents in macaque striate cortex. Empirically, the surface N40 appears in association with a CSD configuration which includes current sinks within the thalamorecipient (stellate) subdivisions of lamina 4C and a large current source extending considerably below 4C. Dipole ensemble contributions to the CSD profile were computed and compared to physiological data from this region. Small asymmetries in activation of model stellate laminae were sufficient to produce substantial open field contributions. However, the best fit with empirical CSD profile was found when the simulation included contributions from thalamocortical axons, along with both open and closed field contributions from dual stellate cell sublaminae. High-resolution CSD profiles were shown to be interpretable when computational artifacts characteristic of closed and open fields were identified using a series of differentiation grids.     

Reliability of quantitative EEG (qEEG) measures and LORETA current source density at 30 days

There is a growing interest for using quantitative EEG and LORETA current source density in clinical and research settings. Importantly, if these indices are to be employed in clinical settings then the reliability of these measures is of great concern. Neuroguide (Applied Neurosciences) is sophisticated software developed for the analyses of power, and connectivity measures of the EEG as well as LORETA current source density. To date there are relatively few data evaluating topographical EEG reliability contrasts for all 19 channels and no studies have evaluated reliability for LORETA calculations. We obtained 4 min eyes-closed and eyes-opened EEG recordings at 30-day intervals. The EEG was analyzed in Neuroguide and FFT power, coherence and phase was computed for traditional frequency bands (delta, theta, alpha and beta) and LORETA current source density was calculated in 1 Hz increments and summed for total power in eight regions of interest (ROI). In order to obtain a robust measure of reliability we utilized a random effects model with an absolute agreement definition. The results show very good reproducibility for total absolute power and coherence. Phase shows lower reliability coefficients. LORETA current source density shows very good reliability with an average 0.81 for ECB and 0.82 for EOB. Similarly, the eight regions of interest show good to very good agreement across time. Implications for future directions and use of qEEG and LORETA in clinical populations are discussed.     

Sources of attention-sensitive visual event-related potentials

Summary In a study of the neural processes that mediate visual attention in humans, 32-channel recordings of event-related potentials were obtained from 14 normal subjects while they performed a spatial attention task. The generator locations of the early C1, P1, and Nl components of the visual evoked response were estimated by means of topographic maps of voltage and current source density in conjunction with dipole modelling. The topography of the C1 component (ca. 85 ms post-stimulus) was consistent with a generator in striate cortex, and this component was unaffected by attention. In contrast, the P1 and Nl components (ca. 95 and 170ms) exhibited current density foci at scalp sites overlying lateral extrastriate cortex and were larger for attended stimuli than for unattended stimuli. The voltage topographies in the 75–175 ms latency range were modeled with a 5-dipole configuration consisting of a single striate dipole and left-right pairs of dipoles located in lateral extrastriate and inferior occipito-temporal areas. This model was found to account for the voltage topographies produced by both attended and unattended stimuli with low residual variance. These results support the proposal that visual-spatial attention modulates neural activity in extrastriate visual cortex but does not affect the initial evoked response in striate cortex.This study was supported by ONR Contract N00014-89-J-1806, by grants from NIMH (MH-25594), NINCDS (NS 17778), the Human Frontier Science Program, DGICYT (PM92-0128), and by a Fulbright scholarship to the first author.     

Unilateral right-hand contractions cause contralateral alpha power suppression and approach motivational affective experience

Contractions of the left hand induce sadness and bias judgments negatively, whereas contractions of the right hand induce positive affect and assertiveness and bias judgments positively. These results have been explained as resulting from activation of right and left frontal cortices, respectively. However, no research has tested this explanation. The present experiment provided such a test by having participants contract the right or left hand while electroencephalographic activity was recorded. Results indicated that right-hand contractions produced greater left than right frontal activity, whereas left-hand contractions produced greater right than left frontal activity (inverse of alpha power). Similar activations occurred in central regions, perhaps due to mu rhythms. Moreover, as compared to left-hand contractions, right-hand contractions caused greater self-reported approach affect to a mildly positive radio editorial.     

Preventing (impulsive) errors: Electrophysiological evidence for online inhibitory control over incorrect responses

In a rich environment, with multiple action affordances, selective action inhibition is critical in preventing the execution of inappropriate responses. Here, we studied the origin and the dynamics of incorrect response inhibition and how it can be modulated by task demands. We used EEG in a conflict task where the probability of compatible and incompatible trials was varied. This allowed us to modulate the strength of the prepotent response, and hence to increase the risk of errors, while keeping the probability of the two responses equal. The correct response activation and execution was not affected by compatibility or by probability. In contrast, incorrect response inhibition in the primary motor cortex ipsilateral to the correct response was more pronounced on incompatible trials, especially in the condition where most of the trials were compatible, indicating a modulation of inhibitory strength within the course of the action. Two prefrontal activities, one medial and one lateral, were also observed before the response, and their potential links with the observed inhibitory pattern observed are discussed.