Temporal Characteristics of EEG Microstates Mediate Trial-by-Trial Risk Taking.

People seem to have difficulties when perceiving events whose outcome has no influence on the outcome of future events. This illusion that patterns exist where there are none may lead to adverse consequences, such as escalating losses in financial trading or gambling debt. Despite the enormous social consequences of these cognitive biases, however, their neural underpinnings are poorly understood. Attempts to investigate them have so far relied on evoked neural activity, whereas spontaneous brain activity has been treated as noise to be averaged out. Here, we focus on the spontaneous electroencephalographic (EEG) activity during inter-trial-intervals (ITI) in a sequential risky decision-making task. Using multilevel mediation analyses, our results show that the percentage of time covered by two EEG microstates (i.e., functional brain-states of coherent activity) mediate the influence of outcomes of prior decisions on subsequent risk taking on a trial-by-trial basis. The devised multilevel mediation analysis of the temporal characteristics of EEG microstates during ITI provides a new window into the neurobiology of decision making by bringing the spontaneous brain activity to the forefront of the analysis.

Modalities of Thinking: State and Trait Effects on Cross-Frequency Functional Independent Brain Networks.

Functional states of the brain are constituted by the temporally attuned activity of spatially distributed neural networks. Such networks can be identified by independent component analysis (ICA) applied to frequency-dependent source-localized EEG data. This methodology allows the identification of networks at high temporal resolution in frequency bands of established location-specific physiological functions. EEG measurements are sensitive to neural activity changes in cortical areas of modality-specific processing. We tested effects of modality-specific processing on functional brain networks. Phasic modality-specific processing was induced via tasks (state effects) and tonic processing was assessed via modality-specific person parameters (trait effects). Modality-specific person parameters and 64-channel EEG were obtained from 70 male, right-handed students. Person parameters were obtained using cognitive style questionnaires, cognitive tests, and thinking modality self-reports. EEG was recorded during four conditions: spatial visualization, object visualization, verbalization, and resting. Twelve cross-frequency networks were extracted from source-localized EEG across six frequency bands using ICA. RMANOVAs, Pearson correlations, and path modelling examined effects of tasks and person parameters on networks. Results identified distinct state- and trait-dependent functional networks. State-dependent networks were characterized by decreased, trait-dependent networks by increased alpha activity in sub-regions of modality-specific pathways. Pathways of competing modalities showed opposing alpha changes. State- and trait-dependent alpha were associated with inhibitory and automated processing, respectively. Antagonistic alpha modulations in areas of competing modalities likely prevent intruding effects of modality-irrelevant processing. Considerable research suggested alpha modulations related to modality-specific states and traits. This study identified the distinct electrophysiological cortical frequency-dependent networks within which they operate.

Zazen meditation and no-task resting EEG compared with LORETA intracortical source localization.

Meditation is a self-induced and willfully initiated practice that alters the state of consciousness. The meditation practice of Zazen, like many other meditation practices, aims at disregarding intrusive thoughts while controlling body posture. It is an open monitoring meditation characterized by detached moment-to-moment awareness and reduced conceptual thinking and self-reference. Which brain areas differ in electric activity during Zazen compared to task-free resting? Since scalp electroencephalography (EEG) waveforms are reference-dependent, conclusions about the localization of active brain areas are ambiguous. Computing intracerebral source models from the scalp EEG data solves this problem. In the present study, we applied source modeling using low resolution brain electromagnetic tomography (LORETA) to 58-channel scalp EEG data recorded from 15 experienced Zen meditators during Zazen and no-task resting. Zazen compared to no-task resting showed increased alpha-1 and alpha-2 frequency activity in an exclusively right-lateralized cluster extending from prefrontal areas including the insula to parts of the somatosensory and motor cortices and temporal areas. Zazen also showed decreased alpha and beta-2 activity in the left angular gyrus and decreased beta-1 and beta-2 activity in a large bilateral posterior cluster comprising the visual cortex, the posterior cingulate cortex and the parietal cortex. The results include parts of the default mode network and suggest enhanced automatic memory and emotion processing, reduced conceptual thinking and self-reference on a less judgmental, i.e., more detached moment-to-moment basis during Zazen compared to no-task resting.

Brain electrical source imaging in manic and depressive episodes of bipolar disorder.

OBJECTIVE: Bipolar disorder (BD) electroencephalographic (EEG) studies have reported varying results. The present study compared EEG in BD during manic and depressive episodes, using brain electrical source imaging [standardized low-resolution electromagnetic tomography (sLORETA)] to assess the cortical spatial distribution of the sources of EEG oscillation frequencies. METHODS: Two independent datasets (a total of 95 patients with bipolar I disorder, of whom 59 were female) were analyzed. Dataset #1 comprised 14 patients in a manic as well as a depressive episode. Dataset #2 comprised 26 patients in a manic episode and 55 patients in a depressive episode. From the head surface-recorded EEG, sLORETA cortical activity was computed in eight EEG frequency bands, and compared between mood states in both datasets. The results from the two datasets were combined using conjunction analysis. RESULTS: Conjunction analysis yielded significant differences between mood states: In manic compared to depressive states, patients had lesser theta frequency band activity (right-hemispheric lateral lower prefrontal and anterior temporal, mainly Brodmann areas 13, 38, and 47), and greater beta-2 and beta-3 frequency band activity (extended bilateral prefrontal-to-parietal, mainly Brodmann area 6, and the cingulate). CONCLUSIONS: The spatial organization of the brain's electrical oscillations differed in patients with BD between manic and depressive mood states. The brain areas implementing the main functions that show opposing abnormalities during manic and depressive episodes were affected by unduly increased or decreased activity (beta or theta). The discussion considers that facilitating (beta) or inhibiting (theta) electrical activity can in either case result in behavioral facilitation or inhibition, depending on the function of the brain area.

Reduced functional connectivity between cortical sources in five meditation traditions detected with lagged coherence using EEG tomography.

Brain functional states are established by functional connectivities between brain regions. In experienced meditators (13 Tibetan Buddhists, 15 QiGong, 14 Sahaja Yoga, 14 Ananda Marga Yoga, 15 Zen), 19-channel EEG was recorded before, during and after that meditation exercise which their respective tradition regards as route to the most desirable meditative state. The head surface EEG data were recomputed (sLORETA) into 19 cortical regional source model time series. All 171 functional connectivities between regions were computed as 'lagged coherence' for the eight EEG frequency bands (delta through gamma). This analysis removes ambiguities of localization, volume conduction-induced inflation of coherence, and reference-dependence. All significant differences (corrected for multiple testing) between meditation compared to no-task rest before and after meditation showed lower coherence during meditation, in all five traditions and eight (inhibitory as well as excitatory) frequency bands. Conventional coherence between the original head surface EEG time series very predominantly also showed reduced coherence during meditation. The topography of the functional connectivities was examined via PCA-based computation of principal connectivities. When going into and out of meditation, significantly different connectivities revealed clearly different topographies in the delta frequency band and minor differences in the beta-2 band. The globally reduced functional interdependence between brain regions in meditation suggests that interaction between the self process functions is minimized, and that constraints on the self process by other processes are minimized, thereby leading to the subjective experience of non-involvement, detachment and letting go, as well as of all-oneness and dissolution of ego borders during meditation.

EEG microstates during resting represent personality differences.

We investigated the spontaneous brain electric activity of 13 skeptics and 16 believers in paranormal phenomena; they were university students assessed with a self-report scale about paranormal beliefs. 33-channel EEG recordings during no-task resting were processed as sequences of momentary potential distribution maps. Based on the maps at peak times of Global Field Power, the sequences were parsed into segments of quasi-stable potential distribution, the 'microstates'. The microstates were clustered into four classes of map topographies (A-D). Analysis of the microstate parameters time coverage, occurrence frequency and duration as well as the temporal sequence (syntax) of the microstate classes revealed significant differences: Believers had a higher coverage and occurrence of class B, tended to decreased coverage and occurrence of class C, and showed a predominant sequence of microstate concatenations from A to C to B to A that was reversed in skeptics (A to B to C to A). Microstates of different topographies, putative "atoms of thought", are hypothesized to represent different types of information processing.The study demonstrates that personality differences can be detected in resting EEG microstate parameters and microstate syntax. Microstate analysis yielded no conclusive evidence for the hypothesized relation between paranormal belief and schizophrenia.

EEG source imaging during two Qigong meditations.

Experienced Qigong meditators who regularly perform the exercises "Thinking of Nothing" and "Qigong" were studied with multichannel EEG source imaging during their meditations. The intracerebral localization of brain electric activity during the two meditation conditions was compared using sLORETA functional EEG tomography. Differences between conditions were assessed using t statistics (corrected for multiple testing) on the normalized and log-transformed current density values of the sLORETA images. In the EEG alpha-2 frequency, 125 voxels differed significantly; all were more active during "Qigong" than "Thinking of Nothing," forming a single cluster in parietal Brodmann areas 5, 7, 31, and 40, all in the right hemisphere. In the EEG beta-1 frequency, 37 voxels differed significantly; all were more active during "Thinking of Nothing" than "Qigong," forming a single cluster in prefrontal Brodmann areas 6, 8, and 9, all in the left hemisphere. Compared to combined initial-final no-task resting, "Qigong" showed activation in posterior areas whereas "Thinking of Nothing" showed activation in anterior areas. The stronger activity of posterior (right) parietal areas during "Qigong" and anterior (left) prefrontal areas during "Thinking of Nothing" may reflect a predominance of self-reference, attention and input-centered processing in the "Qigong" meditation, and of control-centered processing in the "Thinking of Nothing" meditation.

Core networks for visual-concrete and abstract thought content: a brain electric microstate analysis.

Commonality of activation of spontaneously forming and stimulus-induced mental representations is an often made but rarely tested assumption in neuroscience. In a conjunction analysis of two earlier studies, brain electric activity during visual-concrete and abstract thoughts was studied. The conditions were: in study 1, spontaneous stimulus-independent thinking (post-hoc, visual imagery or abstract thought were identified); in study 2, reading of single nouns ranking high or low on a visual imagery scale. In both studies, subjects' tasks were similar: when prompted, they had to recall the last thought (study 1) or the last word (study 2). In both studies, subjects had no instruction to classify or to visually imagine their thoughts, and accordingly were not aware of the studies' aim. Brain electric data were analyzed into functional topographic brain images (using LORETA) of the last microstate before the prompt (study 1) and of the word-type discriminating event-related microstate after word onset (study 2). Conjunction analysis across the two studies yielded commonality of activation of core networks for abstract thought content in left anterior superior regions, and for visual-concrete thought content in right temporal-posterior inferior regions. The results suggest that two different core networks are automatedly activated when abstract or visual-concrete information, respectively, enters working memory, without a subject task or instruction about the two classes of information, and regardless of internal or external origin, and of input modality. These core machineries of working memory thus are invariant to source or modality of input when treating the two types of information.

Tonic activity level in the right prefrontal cortex predicts individuals' risk taking.

Human risk taking is characterized by a large amount of individual heterogeneity. In this study, we applied resting-state electroencephalography, which captures stable individual differences in neural activity, before subjects performed a risk-taking task. Using a source-localization technique, we found that the baseline cortical activity in the right prefrontal cortex predicts individual risk-taking behavior. Individuals with higher baseline cortical activity in this brain area display more risk aversion than do other individuals. This finding demonstrates that neural characteristics that are stable over time can predict a highly complex behavior such as risk-taking behavior and furthermore suggests that hypoactivity in the right prefrontal cortex might serve as a dispositional indicator of lower regulatory abilities, which is expressed in greater risk-taking behavior.

Meditators and non-meditators: EEG source imaging during resting.

Many meditation exercises aim at increased awareness of ongoing experiences through sustained attention and at detachment, i.e., non-engaging observation of these ongoing experiences by the intent not to analyze, judge or expect anything. Long-term meditation practice is believed to generalize the ability of increased awareness and greater detachment into everyday life. We hypothesized that neuroplasticity effects of meditation (correlates of increased awareness and detachment) would be detectable in a no-task resting state. EEG recorded during resting was compared between Qigong meditators and controls. Using LORETA (low resolution electromagnetic tomography) to compute the intracerebral source locations, differences in brain activations between groups were found in the inhibitory delta EEG frequency band. In the meditators, appraisal systems were inhibited, while brain areas involved in the detection and integration of internal and external sensory information showed increased activation. This suggests that neuroplasticity effects of long-term meditation practice, subjectively described as increased awareness and greater detachment, are carried over into non-meditating states.

Rivastigmine effects on EEG spectra and three-dimensional LORETA functional imaging in Alzheimer's disease.

OBJECTIVE: The objective of the study is to investigate the electrocortical and the global cognitive effects of 3 months rivastigmine medication in a group of mild to moderate Alzheimer's disease patients. MATERIALS AND METHODS: Multichannel EEG and cognitive performances measured with the Mini Mental State Examination in a group of 16 patients with mild to moderate Alzheimer's Disease were collected before and 3 months after the onset of rivastigmine medication. RESULTS: Spectral analysis of the EEG data showed a significant power decrease in the delta and theta frequency bands during rivastigmine medication, i.e., a shift of the power spectrum towards 'normalization'. Three-dimensional low resolution electromagnetic tomography (LORETA) functional imaging localized rivastigmine effects in a network that includes left fronto-parietal regions, posterior cingulate cortex, bilateral parahippocampal regions, and the hippocampus. Moreover, a correlation analysis between differences in the cognitive performances during the two recordings and LORETA-computed intracortical activity showed, in the alpha1 frequency band, better cognitive performance with increased cortical activity in the left insula. CONCLUSION: The results point to a 'normalization' of the EEG power spectrum due to medication, and the intracortical localization of these effects showed an increase of cortical activity in frontal, parietal, and temporal regions that are well-known to be affected in Alzheimer's disease. The topographic convergence of the present results with the memory network proposed by Vincent et al. (J. Neurophysiol. 96:3517-3531, 2006) leads to the speculation that in our group of patients, rivastigmine specifically activates brain regions that are involved in memory functions, notably a key symptom in this degenerative disease.

Correlation between disease severity and brain electric LORETA tomography in Alzheimer's disease.

OBJECTIVE: To compare EEG power spectra and LORETA-computed intracortical activity between Alzheimer's disease (AD) patients and healthy controls, and to correlate the results with cognitive performance in the AD group. METHODS: Nineteen channel resting EEG was recorded in 21 mild to moderate AD patients and in 23 controls. Power spectra and intracortical LORETA tomography were computed in seven frequency bands and compared between groups. In the AD patients, the EEG results were correlated with cognitive performance (Mini Mental State Examination, MMSE). RESULTS: AD patients showed increased power in EEG delta and theta frequency bands, and decreased power in alpha2, beta1, beta2 and beta3. LORETA specified that increases and decreases of power affected different cortical areas while largely sparing prefrontal cortex. Delta power correlated negatively and alpha1 power positively with the AD patients' MMSE scores; LORETA tomography localized these correlations in left temporo-parietal cortex. CONCLUSIONS: The non-invasive EEG method of LORETA localized pathological cortical activity in our mild to moderate AD patients in agreement with the literature, and yielded striking correlations between EEG delta and alpha1 activity and MMSE scores in left temporo-parietal cortex. SIGNIFICANCE: The present data support the hypothesis of an asymmetrical progression of the Alzheimer's disease.

Coherence and phase locking in the scalp EEG and between LORETA model sources, and microstates as putative mechanisms of brain temporo-spatial functional organization.

Brain electric mechanisms of temporary, functional binding between brain regions are studied using computation of scalp EEG coherence and phase locking, sensitive to time differences of few milliseconds. However, such results if computed from scalp data are ambiguous since electric sources are spatially oriented. Non-ambiguous results can be obtained using calculated time series of strength of intracerebral model sources. This is illustrated applying LORETA modeling to EEG during resting and meditation. During meditation, time series of LORETA model sources revealed a tendency to decreased left-right intracerebral coherence in the delta band, and to increased anterior-posterior intracerebral coherence in the theta band. An alternate conceptualization of functional binding is based on the observation that brain electric activity is discontinuous, i.e., that it occurs in chunks of up to about 100 ms duration that are detectable as quasi-stable scalp field configurations of brain electric activity, called microstates. Their functional significance is illustrated in spontaneous and event-related paradigms, where microstates associated with imagery- versus abstract-type mentation, or while reading positive versus negative emotion words showed clearly different regions of cortical activation in LORETA tomography. These data support the concept that complete brain functions of higher order such as a momentary thought might be incorporated in temporal chunks of processing in the range of tens to about 100 ms as quasi-stable brain states; during these time windows, subprocesses would be accepted as members of the ongoing chunk of processing.

Nonsmooth nonnegative matrix factorization (nsNMF).

We propose a novel nonnegative matrix factorization model that aims at finding localized, part-based, representations of nonnegative multivariate data items. Unlike the classical nonnegative matrix factorization (NMF) technique, this new model, denoted "nonsmooth nonnegative matrix factorization" (nsNMF), corresponds to the optimization of an unambiguous cost function designed to explicitly represent sparseness, in the form of nonsmoothness, which is controlled by a single parameter. In general, this method produces a set of basis and encoding vectors that are not only capable of representing the original data, but they also extract highly localized patterns, which generally lend themselves to improved interpretability. The properties of this new method are illustrated with several data sets. Comparisons to previously published methods show that the new nsNMF method has some advantages in keeping faithfulness to the data in the achieving a high degree of sparseness for both the estimated basis and the encoding vectors and in better interpretability of the factors.

Psychobiology of altered states of consciousness.

The article reviews the current knowledge regarding altered states of consciousness (ASC) (a) occurring spontaneously, (b) evoked by physical and physiological stimulation, (c) induced by psychological means, and (d) caused by diseases. The emphasis is laid on psychological and neurobiological approaches. The phenomenological analysis of the multiple ASC resulted in 4 dimensions by which they can be characterized: activation, awareness span, self-awareness, and sensory dynamics. The neurophysiological approach revealed that the different states of consciousness are mainly brought about by a compromised brain structure, transient changes in brain dynamics (disconnectivity), and neurochemical and metabolic processes. Besides these severe alterations, environmental stimuli, mental practices, and techniques of self-control can also temporarily alter brain functioning and conscious experience.

EEG microstate duration and syntax in acute, medication-naive, first-episode schizophrenia: a multi-center study.

In young, first-episode, productive, medication-naive patients with schizophrenia, EEG microstates (building blocks of mentation) tend to be shortened. Koenig et al. [Koenig, T., Lehmann, D., Merlo, M., Kochi, K., Hell, D., Koukkou, M., 1999. A deviant EEG brain microstate in acute, neuroleptic-naive schizophrenics at rest. European Archives of Psychiatry and Clinical Neuroscience 249, 205-211] suggested that shortening concerned specific microstate classes. Sequence rules (microstate concatenations, syntax) conceivably might also be affected. In 27 patients of the above type and 27 controls, from three centers, multichannel resting EEG was analyzed into microstates using k-means clustering of momentary potential topographies into four microstate classes (A-D). In patients, microstates were shortened in classes B and D (from 80 to 70 ms and from 94 to 82 ms, respectively), occurred more frequently in classes A and C, and covered more time in A and less in B. Topography differed only in class B where LORETA tomography predominantly showed stronger left and anterior activity in patients. Microstate concatenation (syntax) generally were disturbed in patients; specifically, the class sequence A-->C-->D-->A predominated in controls, but was reversed in patients (A-->D-->C-->A). In schizophrenia, information processing in certain classes of mental operations might deviate because of precocious termination. The intermittent occurrence might account for Bleuler's "double bookkeeping." The disturbed microstate syntax opens a novel physiological comparison of mental operations between patients and controls.

Brain electrical activity and subjective experience during altered states of consciousness: ganzfeld and hypnagogic states.

Manifestations of experimentally induced altered states of consciousness in the brain's electrical activity as well as in subjective experience were explored via the hypnagogic state at sleep onset, and the state induced by exposure to an unstructured perceptual field (ganzfeld). Twelve female paid volunteers participated in sessions involving sleep onset, ganzfeld, and eyes-closed relaxed waking, and were repeatedly prompted for recall of their momentary mentation, according to a predefined schedule. Nineteen channel EEG, two channels EOG and EMG were recorded simultaneously. The mentation reports were followed by the subjects' ratings of their experience on a number of ordinal scales. Two-hundred and forty-one mentation reports were collected. EEG epochs immediately preceding the mentation reports were FFT-analysed and the spectra compared between states. The ganzfeld EEG spectrum, showing no signs of decreased vigilance, was very similar to the EEG spectrum of waking states, even showed a minor acceleration of alpha activity. The subjective experience data were reduced to four principal components: Factor I represented the subjective vigilance dimension, as confirmed by correlations with EEG spectral indices. Only Factor IV, the 'absorption' dimension, differentiated between the ganzfeld state (more absorption) and other states. In waking states and in ganzfeld, the subjects estimated elapsed time periods significantly shorter than in states at sleep onset. The results did not support the assumption of a hypnagogic nature of the ganzfeld imagery. Dream-like imagery can occur in various global functional states of the brain; hypnagogic and ganzfeld-induced states should be conceived as special cases of a broader class of 'hypnagoid' phenomena.

sLORETA intracortical lagged coherence during breath counting in meditation-naïve participants.

We investigated brain functional connectivity comparing no-task resting to breath counting (a meditation exercise but given as task without referring to meditation). Functional connectivity computed as EEG coherence between head-surface data suffers from localization ambiguity, reference dependence, and overestimation due to volume conduction. Lagged coherence between intracortical model sources addresses these criticisms. With this analysis approach, experienced meditators reportedly showed reduced coherence during meditation, meditation-naïve participants have not yet been investigated. 58-channel EEG from 23 healthy, right-handed, meditation-naïve males during resting [3 runs] and breath counting [2 runs] was computed into sLORETA time series of intracortical electrical activity in 19 regions of interest (ROI) corresponding to the cortex underlying 19 scalp electrode sites, for each of the eight independent EEG frequency bands covering 1.5-44 Hz. Intracortical lagged coherences and head-surface conventional coherences were computed between the 19 regions/sites. During breath counting compared to resting, paired t-tests corrected for multiple testing revealed four significantly lower intracortical lagged coherences, but four significantly higher head-surface conventional coherences. Lowered intracortical lagged coherences involved left BA 10 and right BAs 3, 10, 17, 40. In conclusion, intracortical lagged coherence can yield results that are inverted to those of head-surface conventional coherence. The lowered functional connectivity between cognitive control areas and sensory perception areas during meditation-type breath counting compared to resting conceivably reflects the attention to a bodily percept without cognitive reasoning. The reductions in functional connectivity were similar but not as widespread as the reductions reported during meditation in experienced meditators.

Functionally aberrant electrophysiological cortical connectivities in first episode medication-naive schizophrenics from three psychiatry centers.

Functional dissociation between brain processes is widely hypothesized to account for aberrations of thought and emotions in schizophrenic patients. The typically small groups of analyzed schizophrenic patients yielded different neurophysiological findings, probably because small patient groups are likely to comprise different schizophrenia subtypes. We analyzed multichannel eyes-closed resting EEG from three small groups of acutely ill, first episode productive schizophrenic patients before start of medication (from three centers: Bern N = 9; Osaka N = 9; Berlin N = 12) and their controls. Low resolution brain electromagnetic tomography (LORETA) was used to compute intracortical source model-based lagged functional connectivity not biased by volume conduction effects between 19 cortical regions of interest (ROIs). The connectivities were compared between controls and patients of each group. Conjunction analysis determined six aberrant cortical functional connectivities that were the same in the three patient groups. Four of these six concerned the facilitating EEG alpha-1 frequency activity; they were decreased in the patients. Another two of these six connectivities concerned the inhibiting EEG delta frequency activity; they were increased in the patients. The principal orientation of the six aberrant cortical functional connectivities was sagittal; five of them involved both hemispheres. In sum, activity in the posterior brain areas of preprocessing functions and the anterior brain areas of evaluation and behavior control functions were compromised by either decreased coupled activation or increased coupled inhibition, common across schizophrenia subtypes in the three patient groups. These results of the analyzed three independent groups of schizophrenics support the concept of functional dissociation.