Combined effects of neuroticism and negative emotional context on spontaneous EEG dynamics.

Neuroticism is a personality trait with great clinical relevance, defined as a tendency to experience negative affect, sustained self-generated negative thoughts and impaired emotion regulation. Here, we investigated spontaneous brain dynamics in the aftermath of negative emotional events and their links with neuroticism in order to shed light on the prolonged activity of large-scale brain networks associated with the control of affect. We recorded electroencephalography (EEG) from 36 participants who were asked to rest after watching neutral or fearful video clips. Four topographic maps (i.e. microstates classes A, B, C and D) explained the majority of the variance in spontaneous EEG. Participants showed greater presence of microstate D and lesser presence of microstate C following exposure to fearful stimuli, pointing to changes in attention- and introspection-related networks previously associated with these microstates. These emotional effects were more pronounced for participants with low neuroticism. Moreover, neuroticism scores were positively correlated with microstate C and negatively correlated with microstate D, regardless of previous emotional stimulation. Our results reveal distinctive effects of emotional context on resting-state EEG, consistent with a prolonged impact of negative affect on the brain, and suggest a possible link with neuroticism.

Electrophysiological correlates of distance and direction processing during cognitive map retrieval: A source analysis.

Introduction: The cognitive map is an internal representation of the environment and allows us to navigate through familiar environments. It preserves the distances and directions between landmarks which help us orient ourselves in our surroundings. The aim of our task was to understand the role played by theta waves in the cognitive map and especially how the cognitive map is recalled and how the manipulation of distances and directions occurs within the cognitive map. Method: In order to investigate the neural correlates of the cognitive map, we used the Cognitive Map Recall Test, in which 33 participants had to estimate distances and directions between familiar landmarks tailored to their own knowledge. We examined the role of theta waves in the cognitive map, as well as the brain regions that generated them. To that aim, we performed electroencephalographic source imaging while focusing on frequency spectral analysis. Results: We observed increases of theta amplitude in the frontal, temporal, parahippocampal gyri and temporal poles during the recall of the cognitive map. We also found increases of theta amplitude in the temporal pole and retrosplenial cortex during manipulation of directions. Overall, direction processing induces higher theta amplitude than distance processing, especially in the temporal lobe, and higher theta amplitude during recall compared to manipulation, except in the retrosplenial cortex where this pattern was reversed. Discussion: We reveal the role of theta waves as a marker of directional processing in the retrosplenial cortex and the temporal poles during the manipulation of spatial information. Increases in theta waves in frontal, parahippocampal, temporal and temporal pole regions appear to be markers of working memory and cognitive map recall. Therefore, our Cognitive Map Recall Test could be useful for testing directional difficulties in patients. Our work also shows that there are two distinct parts to the cognitive map test: recall and manipulation of spatial information. This is often considered as two similar processes in the literature, but our work demonstrates that these processes could be different, with theta waves from different brain regions contributing to either recall or manipulation; this should be considered in future studies.

Oscillatory Neural Signatures of Visual Perception Across Developmental Stages in Individuals With 22q11.2 Deletion Syndrome.

BACKGROUND: Numerous behavioral studies have highlighted the contribution of visual perceptual deficits to the nonverbal cognitive profile of individuals with 22q11.2 deletion syndrome. However, the neurobiological processes underlying these widespread behavioral alterations are yet to be fully understood. Thus, in this paper, we investigated the role of neural oscillations toward visuoperceptual deficits to elucidate the neurobiology of sensory impairments in deletion carriers. METHODS: We acquired 125 high-density electroencephalography recordings during a visual grating task in a group of 62 deletion carriers and 63 control subjects. Stimulus-elicited oscillatory responses were analyzed with 1) time-frequency analysis using wavelets decomposition at sensor and source level, 2) intertrial phase coherence, and 3) Granger causality connectivity in source space. Additional analyses examined the development of neural oscillations across age bins. RESULTS: Deletion carriers had decreased theta-band (4-8 Hz) and gamma-band (58-68 Hz) spectral power compared with control subjects in response to the visual stimuli, with an absence of age-related increase of theta- and gamma-band responses. Moreover, adult deletion carriers had decreased gamma- and theta-band responses but increased alpha/beta desynchronization (10-25 Hz) that correlated with behavioral performance. Granger causality estimates reflected an increased frontal-occipital connectivity in the beta range (22-40 Hz). CONCLUSIONS: Deletion carriers exhibited decreased theta- and gamma-band responses to visual stimuli, while alpha/beta desynchronization was preserved. Overall, the lack of age-related changes in deletion carriers implicates developmental impairments in circuit mechanisms underlying neural oscillations. The dissociation between the maturation of theta/gamma- and alpha/beta-band responses may indicate a selective impairment in supragranular cortical layers, leading to compensatory top-down connectivity.

EEG microstate dynamics indicate a U-shaped path to propofol-induced loss of consciousness.

Evidence suggests that the stream of consciousness is parsed into transient brain states manifesting themselves as discrete spatiotemporal patterns of global neuronal activity. Electroencephalographical (EEG) microstates are proposed as the neurophysiological correlates of these transiently stable brain states that last for fractions of seconds. To further understand the link between EEG microstate dynamics and consciousness, we continuously recorded high-density EEG in 23 surgical patients from their awake state to unconsciousness, induced by step-wise increasing concentrations of the intravenous anesthetic propofol. Besides the conventional parameters of microstate dynamics, we introduce a new implementation of a method to estimate the complexity of microstate sequences. The brain activity under the surgical anesthesia showed a decreased sequence complexity of the stereotypical microstates, which became sparser and longer-lasting. However, we observed an initial increase in microstates' temporal dynamics and complexity with increasing depth of sedation leading to a distinctive "U-shape" that may be linked to the paradoxical excitation induced by moderate levels of propofol. Our results support the idea that the brain is in a metastable state under normal conditions, balancing between order and chaos in order to flexibly switch from one state to another. The temporal dynamics of EEG microstates indicate changes of this critical balance between stability and transition that lead to altered states of consciousness.

Aberrant Developmental Patterns of Gamma-Band Response and Long-Range Communication Disruption in Youths With 22q11.2 Deletion Syndrome.

OBJECTIVE: Brain oscillations play a pivotal role in synchronizing responses of local and global ensembles of neurons. Patients with schizophrenia exhibit impairments in oscillatory response, which are thought to stem from abnormal maturation during critical developmental stages. Studying individuals at genetic risk for psychosis, such as 22q11.2 deletion carriers, from childhood to adulthood may provide insights into developmental abnormalities. METHODS: The authors acquired 106 consecutive T1-weighted MR images and 40-Hz auditory steady-state responses (ASSRs) with high-density (256 channel) EEG in a group of 58 22q11.2 deletion carriers and 48 healthy control subjects. ASSRs were analyzed with 1) time-frequency analysis using Morlet wavelet decomposition, 2) intertrial phase coherence (ITPC), and 3) theta-gamma phase-amplitude coupling estimated in the source space between brain regions activated by the ASSRs. Additionally, volumetric analyses were performed with FreeSurfer. Subanalyses were conducted in deletion carriers who endorsed psychotic symptoms and in subgroups with different age bins. RESULTS: Deletion carriers had decreased theta and late-latency 40-Hz ASSRs and phase synchronization compared with control subjects. Deletion carriers with psychotic symptoms displayed a further reduction of gamma-band response, decreased ITPC, and decreased top-down modulation of gamma-band response in the auditory cortex. Reduced gamma-band response was correlated with the atrophy of auditory cortex in individuals with psychotic symptoms. In addition, a linear increase of theta and gamma power from childhood to adulthood was found in control subjects but not in deletion carriers. CONCLUSIONS: The results suggest that while all deletion carriers exhibit decreased gamma-band response, more severe local and long-range communication abnormalities are associated with the emergence of psychotic symptoms and gray matter loss. Additionally, the lack of age-related changes in deletion carriers indexes a potential developmental impairment in circuits underlying the maturation of neural oscillations during adolescence. The progressive disruption of gamma-band response in 22q11.2 deletion syndrome supports a developmental perspective toward understanding and treating psychotic disorders.

Beyond broadband: Towards a spectral decomposition of electroencephalography microstates.

Originally applied to alpha oscillations in the 1970s, microstate (MS) analysis has since been used to decompose mainly broadband electroencephalogram (EEG) signals (e.g., 1-40 Hz). We hypothesised that MS decomposition within separate, narrow frequency bands could provide more fine-grained information for capturing the spatio-temporal complexity of multichannel EEG. In this study, using a large open-access dataset (n = 203), we first filtered EEG recordings into four classical frequency bands (delta, theta, alpha and beta) and thereafter compared their individual MS segmentations using mutual information as well as traditional MS measures (e.g., mean duration and time coverage). Firstly, we confirmed that MS topographies were spatially equivalent across all frequencies, matching the canonical broadband maps (A, B, C, D and C'). Interestingly, however, we observed strong informational independence of MS temporal sequences between spectral bands, together with significant divergence in traditional MS measures. For example, relative to broadband, alpha/beta band dynamics displayed greater time coverage of maps A and B, while map D was more prevalent in delta/theta bands. Moreover, using a frequency-specific MS taxonomy (e.g., Ï´A and αC), we were able to predict the eyes-open versus eyes-closed behavioural state significantly better using alpha-band MS features compared with broadband ones (80 vs. 73% accuracy). Overall, our findings demonstrate the value and validity of spectrally specific MS analyses, which may prove useful for identifying new neural mechanisms in fundamental research and/or for biomarker discovery in clinical populations.

Phase-Amplitude Coupling and Phase Synchronization Between Medial Temporal, Frontal and Posterior Brain Regions Support Episodic Autobiographical Memory Recall.

Episodic autobiographical memory (EAM) is a complex cognitive function that emerges from the coordination of specific and distant brain regions. Specific brain rhythms, namely theta and gamma oscillations and their synchronization, are thought of as putative mechanisms enabling EAM. Yet, the mechanisms of inter-regional interaction in the EAM network remain unclear in humans at the whole brain level. To investigate this, we analyzed EEG recordings of participants instructed to retrieve autobiographical episodes. EEG recordings were projected in the source space, and time-courses of atlas-based brain regions-of-interest (ROIs) were derived. Directed phase synchrony in high theta (7-10 Hz) and gamma (30-80 Hz) bands and high theta-gamma phase-amplitude coupling were computed between each pair of ROIs. Using network-based statistics, a graph-theory method, we found statistically significant networks for each investigated mechanism. In the gamma band, two sub-networks were found, one between the posterior cingulate cortex (PCC) and the medial temporal lobe (MTL) and another within the medial frontal areas. In the high theta band, we found a PCC to ventromedial prefrontal cortex (vmPFC) network. In phase-amplitude coupling, we found the high theta phase of the left MTL biasing the gamma amplitude of posterior regions and the vmPFC. Other regions of the temporal lobe and the insula were also phase biasing the vmPFC. These findings suggest that EAM, rather than emerging from a single mechanism at a single frequency, involves precise spatio-temporal signatures mapping on distinct memory processes. We propose that the MTL orchestrates activity in vmPFC and PCC via precise phase-amplitude coupling, with vmPFC and PCC interaction via high theta phase synchrony and gamma synchronization contributing to bind information within the PCC-MTL sub-network or valuate the candidate memory within the medial frontal sub-network.

Linear distributed inverse solutions for interictal EEG source localisation.

OBJECTIVE: To compare the spatial accuracy of 6 linear distributed inverse solutions for EEG source localisation of interictal epileptic discharges: Minimum Norm, Weighted Minimum Norm, Low-Resolution Electromagnetic Tomography (LORETA), Local Autoregressive Average (LAURA), Standardised LORETA, and Exact LORETA. METHODS: Spatial accuracy was assessed clinically by retrospectively comparing the maximum source of averaged interictal discharges to the resected brain area in 30 patients with successful epilepsy surgery, based on 204-channel EEG. Additionally, localisation errors of the inverse solutions were assessed in computer simulations, with different levels of noise added to the signal in both sensor space and source space. RESULTS: In the clinical evaluations, the source maximum was located inside the resected brain area in 50-57% of patients when using LORETA or LAURA, while all other inverse solutions performed significantly worse (17-30%; corrected p < 0.01). In the simulation studies, when noise levels exceeded 10%, LORETA and LAURA had substantially smaller localisation errors than the other inverse solutions. CONCLUSIONS: LORETA and LAURA provided the highest spatial accuracy both in clinical and simulated data, alongside with a comparably high robustness towards noise. SIGNIFICANCE: Among the different linear inverse solution algorithms tested, LORETA and LAURA might be preferred for interictal EEG source localisation.

Time to move: Brain dynamics underlying natural action and cognition.

Advances in Mobile Brain/Body Imaging (MoBI) technology allows for real-time measurements of human brain dynamics during every day, natural, real-life situations. This special issue Time to Move brings together a collection of experimental papers, targeted reviews and opinion articles that lay out the latest MoBI findings. A wide range of topics across different fields are covered including art, athletics, virtual reality, and mobility. What unites these diverse topics is the common goal to enhance and restore human abilities by reaching a better understanding on how cognition is implemented by the brain-body relationship. The breadth and novelty of paradigms and findings reported here positions MoBI as a new frontier in the field of human cognitive neuroscience.

Enhancing visual motion discrimination by desynchronizing bifocal oscillatory activity.

Visual motion discrimination involves reciprocal interactions in the alpha band between the primary visual cortex (V1) and mediotemporal areas (V5/MT). We investigated whether modulating alpha phase synchronization using individualized multisite transcranial alternating current stimulation (tACS) over V5 and V1 regions would improve motion discrimination. We tested 3 groups of healthy subjects with the following conditions: (1) individualized In-Phase V1alpha-V5alpha tACS (0° lag), (2) individualized Anti-Phase V1alpha-V5alpha tACS (180° lag) and (3) sham tACS. Motion discrimination and EEG activity were recorded before, during and after tACS. Performance significantly improved in the Anti-Phase group compared to the In-Phase group 10 and 30 min after stimulation. This result was explained by decreases in bottom-up alpha-V1 gamma-V5 phase-amplitude coupling. One possible explanation of these results is that Anti-Phase V1alpha-V5alpha tACS might impose an optimal phase lag between stimulation sites due to the inherent speed of wave propagation, hereby supporting optimized neuronal communication.

Synchronous Brain Dynamics Establish Brief States of Communality in Distant Neuronal Populations.

Intrinsic brain dynamics co-fluctuate between distant regions in an organized manner during rest, establishing large-scale functional networks. We investigate these brain dynamics on a millisecond time scale by focusing on electroencephalographic (EEG) source analyses. While synchrony is thought of as a neuronal mechanism grouping distant neuronal populations into assemblies, the relevance of simultaneous zero-lag synchronization between brain areas in humans remains largely unexplored. This negligence is because of the confound of volume conduction, leading inherently to temporal dependencies of source estimates derived from scalp EEG [and magnetoencephalography (MEG)], referred to as spatial leakage. Here, we focus on the analyses of simultaneous, i.e., quasi zero-lag related, synchronization that cannot be explained by spatial leakage phenomenon. In eighteen subjects during rest with eyes closed, we provide evidence that first, simultaneous synchronization is present between distant brain areas and second, that this long-range synchronization is occurring in brief epochs, i.e., 54-80 ms. Simultaneous synchronization might signify the functional convergence of remote neuronal populations. Given the simultaneity of distant regions, these synchronization patterns might relate to the representation and maintenance, rather than processing of information. This long-range synchronization is briefly stable, not persistently, indicating flexible spatial reconfiguration pertaining to the establishment of particular, re-occurring states. Taken together, we suggest that the balance between temporal stability and spatial flexibility of long-range, simultaneous synchronization patterns is characteristic of the dynamic coordination of large-scale functional brain networks. As such, quasi zero-phase related EEG source fluctuations are physiologically meaningful if spatial leakage is considered appropriately.

High-density Electric Source Imaging of interictal epileptic discharges: How many electrodes and which time point?

OBJECTIVE: To assess the value of caudal EEG electrodes over cheeks and neck for high-density electric source imaging (ESI) in presurgical epilepsy evaluation, and to identify the best time point during averaged interictal epileptic discharges (IEDs) for optimal ESI accuracy. METHODS: We retrospectively examined presurgical 257-channel EEG recordings of 45 patients with pharmacoresistant focal epilepsy. By stepwise removal of cheek and neck electrodes, averaged IEDs were downsampled to 219, 204, and 156 EEG channels. Additionally, ESI at the IED's half-rise was compared to other time points. The respective sources of maximum activity were compared to the resected brain area and postsurgical outcome. RESULTS: Caudal channels had disproportionately more artefacts. In 30 patients with favourable outcome, the 204-channel array yielded the most accurate results with ESI maxima < 10 mm from the resection in 67% and inside affected sublobes in 83%. Neither in temporal nor in extratemporal cases did the full 257-channel setup improve ESI accuracy. ESI was most accurate at 50% of the IED's rising phase. CONCLUSION: Information from cheeks and neck electrodes did not improve high-density ESI accuracy, probably due to higher artefact load and suboptimal biophysical modelling. SIGNIFICANCE: Very caudal EEG electrodes should be used for ESI with caution.

Using structural connectivity to augment community structure in EEG functional connectivity.

Recently, EEG recording techniques and source analysis have improved, making it feasible to tap into fast network dynamics. Yet, analyzing whole-cortex EEG signals in source space is not standard, partly because EEG suffers from volume conduction: Functional connectivity (FC) reflecting genuine functional relationships is impossible to disentangle from spurious FC introduced by volume conduction. Here, we investigate the relationship between white matter structural connectivity (SC) and large-scale network structure encoded in EEG-FC. We start by confirming that FC (power envelope correlations) is predicted by SC beyond the impact of Euclidean distance, in line with the assumption that SC mediates genuine FC. We then use information from white matter structural connectivity in order to smooth the EEG signal in the space spanned by graphs derived from SC. Thereby, FC between nearby, structurally connected brain regions increases while FC between nonconnected regions remains unchanged, resulting in an increase in genuine, SC-mediated FC. We analyze the induced changes in FC, assessing the resemblance between EEG-FC and volume-conduction- free fMRI-FC, and find that smoothing increases resemblance in terms of overall correlation and community structure. This result suggests that our method boosts genuine FC, an outcome that is of interest for many EEG network neuroscience questions.

ECoG Beta Suppression and Modulation During Finger Extension and Flexion.

Neural oscillations originate predominantly from interacting cortical neurons and consequently reflect aspects of cortical information processing. However, their functional role is not yet fully understood and their interpretation is debatable. Amplitude modulations (AMs) in alpha (8-12 Hz), beta (13-30 Hz), and high gamma (70-150 Hz) band in invasive electrocorticogram (ECoG) and non-invasive electroencephalogram (EEG) signals change with behavior. Alpha and beta band AMs are typically suppressed (desynchronized) during motor behavior, while high gamma AMs highly correlate with the behavior. These two phenomena are successfully used for functional brain mapping and brain-computer interface (BCI) applications. Recent research found movement-phase related AMs (MPA) also in high beta/low gamma (24-40 Hz) EEG rhythms. These MPAs were found by separating the suppressed AMs into sustained and dynamic components. Sustained AM components are those with frequencies that are lower than the motor behavior. Dynamic components those with frequencies higher than the behavior. In this paper, we study ECoG beta/low gamma band (12-30 Hz/30-42 Hz) AM during repetitive finger movements addressing the question whether or not MPAs can be found in ECoG beta band. Indeed, MPA in the 12-18 Hz and 18-24 Hz band were found. This additional information may lead to further improvements in ECoG-based prediction and reconstruction of motor behavior by combining high gamma AM and beta band MPA.

Subcortical electrophysiological activity is detectable with high-density EEG source imaging.

Subcortical neuronal activity is highly relevant for mediating communication in large-scale brain networks. While electroencephalographic (EEG) recordings provide appropriate temporal resolution and coverage to study whole brain dynamics, the feasibility to detect subcortical signals is a matter of debate. Here, we investigate if scalp EEG can detect and correctly localize signals recorded with intracranial electrodes placed in the centromedial thalamus, and in the nucleus accumbens. Externalization of deep brain stimulation (DBS) electrodes, placed in these regions, provides the unique opportunity to record subcortical activity simultaneously with high-density (256 channel) scalp EEG. In three patients during rest with eyes closed, we found significant correlation between alpha envelopes derived from intracranial and EEG source reconstructed signals. Highest correlation was found for source signals in close proximity to the actual recording sites, given by the DBS electrode locations. Therefore, we present direct evidence that scalp EEG indeed can sense subcortical signals.

EEG Oscillations Are Modulated in Different Behavior-Related Networks during Rhythmic Finger Movements.

Sequencing and timing of body movements are essential to perform motoric tasks. In this study, we investigate the temporal relation between cortical oscillations and human motor behavior (i.e., rhythmic finger movements). High-density EEG recordings were used for source imaging based on individual anatomy. We separated sustained and movement phase-related EEG source amplitudes based on the actual finger movements recorded by a data glove. Sustained amplitude modulations in the contralateral hand area show decrease for α (10-12 Hz) and ß (18-24 Hz), but increase for high γ (60-80 Hz) frequencies during the entire movement period. Additionally, we found movement phase-related amplitudes, which resembled the flexion and extension sequence of the fingers. Especially for faster movement cadences, movement phase-related amplitudes included high ß (24-30 Hz) frequencies in prefrontal areas. Interestingly, the spectral profiles and source patterns of movement phase-related amplitudes differed from sustained activities, suggesting that they represent different frequency-specific large-scale networks. First, networks were signified by the sustained element, which statically modulate their synchrony levels during continuous movements. These networks may upregulate neuronal excitability in brain regions specific to the limb, in this study the right hand area. Second, movement phase-related networks, which modulate their synchrony in relation to the movement sequence. We suggest that these frequency-specific networks are associated with distinct functions, including top-down control, sensorimotor prediction, and integration. The separation of different large-scale networks, we applied in this work, improves the interpretation of EEG sources in relation to human motor behavior. SIGNIFICANCE STATEMENT: EEG recordings provide high temporal resolution suitable to relate cortical oscillations to actual movements. Investigating EEG sources during rhythmic finger movements, we distinguish sustained from movement phase-related amplitude modulations. We separate these two EEG source elements motivated by our previous findings in gait. Here, we found two types of large-scale networks, representing the right fingers in distinction from the time sequence of the movements. These findings suggest that EEG source amplitudes reconstructed in a cortical patch are the superposition of these simultaneously present network activities. Separating these frequency-specific networks is relevant for studying function and possible dysfunction of the cortical sensorimotor system in humans as well as to provide more advanced features for brain-computer interfaces.

Corrigendum: EEG beta suppression and low gamma modulation are different elements of human upright walking.

[This corrects the article on p. 485 in vol. 8, PMID: 25071515.].

High and low gamma EEG oscillations in central sensorimotor areas are conversely modulated during the human gait cycle.

Investigating human brain function is essential to develop models of cortical involvement during walking. Such models could advance the analysis of motor impairments following brain injuries (e.g., stroke) and may lead to novel rehabilitation approaches. In this work, we applied high-density EEG source imaging based on individual anatomy to enable neuroimaging during walking. To minimize the impact of muscular influence on EEG recordings we introduce a novel artifact correction method based on spectral decomposition. High γ oscillations (>60Hz) were previously reported to play an important role in motor control. Here, we investigate high γ amplitudes while focusing on two different aspects of a walking experiment, namely the fact that a person walks and the rhythmicity of walking. We found that high γ amplitudes (60-80Hz), located focally in central sensorimotor areas, were significantly increased during walking compared to standing. Moreover, high γ (70-90Hz) amplitudes in the same areas are modulated in relation to the gait cycle. Since the spectral peaks of high γ amplitude increase and modulation do not match, it is plausible that these two high γ elements represent different frequency-specific network interactions. Interestingly, we found high γ (70-90Hz) amplitudes to be coupled to low γ (24-40Hz) amplitudes, which both are modulated in relation to the gait cycle but conversely to each other. In summary, our work is a further step towards modeling cortical involvement during human upright walking.

EEG beta suppression and low gamma modulation are different elements of human upright walking.

Cortical involvement during upright walking is not well-studied in humans. We analyzed non-invasive electroencephalographic (EEG) recordings from able-bodied volunteers who participated in a robot-assisted gait-training experiment. To enable functional neuroimaging during walking, we applied source modeling to high-density (120 channels) EEG recordings using individual anatomy reconstructed from structural magnetic resonance imaging scans. First, we analyzed amplitude differences between the conditions, walking and upright standing. Second, we investigated amplitude modulations related to the gait phase. During active walking upper µ (10-12 Hz) and ß (18-30 Hz) oscillations were suppressed [event-related desynchronization (ERD)] compared to upright standing. Significant ß ERD activity was located focally in central sensorimotor areas for 9/10 subjects. Additionally, we found that low γ (24-40 Hz) amplitudes were modulated related to the gait phase. Because there is a certain frequency band overlap between sustained ß ERD and gait phase related modulations in the low γ range, these two phenomena are superimposed. Thus, we observe gait phase related amplitude modulations at a certain ERD level. We conclude that sustained µ and ß ERD reflect a movement related state change of cortical excitability while gait phase related modulations in the low γ represent the motion sequence timing during gait. Interestingly, the center frequencies of sustained ß ERD and gait phase modulated amplitudes were identified to be different. They may therefore be caused by different neuronal rhythms, which should be taken under consideration in future studies.