EEG Electric Field Topography is Stable during Moments of High Field Strength

Anthony P. Zanesco

Department of Psychology, University of Miami

Please address correspondence to Anthony Zanesco, Department of Psychology, University

of Miami, 5665 Ponce de Leon Blvd, Coral Gables, FL 33146. Email: apz13@miami.edu.

Phone: 305-284-8148

Author Statement

I thank the Mind-Body-Emotion group at the Max Planck Institute for Human Cognitive and

Brain Sciences who gathered and made these data available. I utilized the freely available

Cartool software toolbox (cartoolcommunity.unige.ch) programmed by Denis Brunet, from the

Functional Brain Mapping Laboratory, Geneva, Switzerland, and supported by the Center for

Biomedical Imaging of Geneva and Lausanne. The data that support the findings of this study are

openly available in the OSF repository and can be found at: https://osf.io/7bcem/
 Abstract

Spontaneous broadband electroencephalography (EEG) demonstrates short moments of

stability in the spatial distribution of the head-surface voltage topography. This phenomenon

underlies the premise behind segmenting multichannel EEG into topographically defined brain

states, known as EEG microstates. Microstate segmentation methods commonly identify

representative topographical configurations based on clustering applied to a subset of voltage

maps selected at the time series points of greatest strength in the neuroelectric field. These

moments are well-reasoned to best represent periods of quasi-stability, and consequently, points

of greatest signal relative to noise. Yet, more direct empirical evidence for these assumptions is

valuable, and the consistency of this phenomenon across individuals has not been characterized.

In the present investigation, the association between electric field strength and topographic

dissimilarity of temporally adjacent samples of EEG was characterized in a large sample of

healthy adults. Individuals’ demonstrated that samples of their EEG time series high in electric

field strength were topographically similar relative to adjacent time series samples. The strong

activity of phase-synchronized neuronal populations therefore coincides with periods of

topographic stability, providing robust empirical support for the basic premise underlying

segmentation of broadband EEG into microstates.

 Scalp recorded broadband electroencephalography (EEG) exhibits moments of

spontaneous topographic stability that appear to be fundamental to the dynamics of the

neuroelectric field. When carefully examined over short time scales, the spatial distribution of

the head-surface voltage topography plotted as a succession of three-dimensional voltage maps

gives the impression of short periods of stability, in which a particular topographic configuration

predominates momentarily (~ 60–120 msec) before quickly transitioning to a different quasi-

stable configuration. Moreover, the same topographic configurations appear to be common to a

large portion of the voltage maps present during these periods of stability. These observations

underlie the premise behind the segmentation of multichannel EEG time series into microstates

based on clustering of topographic patterns to identify the millisecond spatiotemporal dynamics

of coordinated brain states (Lehmann, Ozaki, and Pal, 1987; Wackermann, Lehmann, Michel,

and Strik, 1993).

Spatial decomposition of EEG time series into microstates has consistently identified a

limited set of data-driven clusters of voltage maps that explain a large portion of observed

topographic variance (see for review, Khanna, Pascual-Leone, Michel, and Farzan, 2015; Michel

and Koenig, 2017). Each distinct topographic configuration of voltage distribution implies by

physical laws different distributions of active neural generators in the brain (Vaughan, 1982),

allowing topography to define changes in the activity of whole-brain neuronal networks. That

dozens of studies have consistently identified similar clusters of maps when segmenting

spontaneous EEG into microstates, suggests a common brain network architecture underlying

large sources of spontaneous phase-synchronized activity observed at rest (Michel and Koenig,

2017). Furthermore, the electrical brain sources of microstates have been found to align with

fMRI-derived resting state functional networks (Britz et al., 2010; Custo et al., 2017; Brechet et
al., 2019), making the microstate approach timely given increasing interest in defining the

spontaneous organization of brain activity into coordinated networks.

Methods of segmenting microstates based on topography commonly utilize topographic

clustering methods applied to a subset of voltage maps (e.g., Michel, Koenig, and Brandeis,

2009), selected at the time series points of greatest strength in the electric field. These points of

greatest field strength can be quantified based on the global field power (GFP), which reflects a

reference-independent measure of the strength of synchronized brain response at a moment in

time (Skrandies, 1990). Selecting local maxima in the GFP time series for topographic clustering

has remained a defensible approach because of the observation that moments of strong phase-

synchronized activity generally demonstrate quasi-stability in their voltage topography

(Skrandies, 1990). These moments therefore reflect points of greatest signal relative to noise

(Koenig and Brandeis, 2016), making peaks in the GFP ideal candidates for identifying quasi-

stable topographic configurations present in the EEG time series.

The most striking evidence for these suppositions, however, come from the broad success

of topographic clustering approaches in identifying common microstate configurations across

studies, and the utility of segmentation procedures in defining microstate sequences (Michel and

Koenig, 2017). Large proportions of topographic variance in the continuous EEG can be

explained by a few cluster centroids derived from clustering of voltage maps at GFP peaks (e.g.,

Seitzman et al., 2017). Yet, more direct empirical evidence for the basic association between

GFP and topographic stability has not been systematically demonstrated in spontaneous EEG

collected at rest. Koenig and Brandeis (2016), however, provided one compelling demonstration

of the strong within-person association between GFP and momentary topographic stability in the

EEG time series. By calculating the similarity between temporally adjacent voltage maps, they
quantified the ongoing topographic stability of the EEG time series. They then demonstrated a

strong association between samples with high GFP and topographic similarity in an hour-long

EEG recording acquired from a single individual (Koenig and Brandeis, 2016). This was an

important observation, but the consistency of this phenomenon across a wider range of

individuals has yet to be characterized.

In the present investigation, I revisit the basic premise behind EEG microstate analysis

that periods of strong phase-synchronized neural activity demonstrate topographic stability for

brief moments of time. In line with the approach used by Koenig and Brandeis (2016), I

examined the consistency and variability of associations between GFP and topographic

dissimilarity of adjacent samples of EEG in a large sample of healthy adults. True to the notion

that strong activity of phase-synchronized neuronal populations coincides with periods of

topographic stability, samples of the EEG time series high in GFP ought to be topographically

similar relative to adjacent samples.

16-minutes of scalp recorded multichannel EEG were acquired from 216 healthy adults at

rest by Babayan and colleagues (2019) and made publicly available on a data-sharing repository.

Resting EEG were divided into eyes-closed (8-minute) and eyes-open (8-minute) epochs. Using

these data, I calculated within-person correlations between the GFP and topographic dissimilarity

of temporally adjacent (sample t and sample t -1) voltage maps for all samples of individuals’

EEG recordings (Skrandies, 1990; Murray, Brunet, and Michel, 2008). In addition to supporting

the notion that maps at local maxima of GFP are strong indicators of momentary stability in

topographic configuration, the degree to which topography is stable relative to GFP may reflect

the general prevalence of microstates predominating in the EEG time series across individuals.

To address this second possibility, I segmented resting EEG into microstates based on data-
driven topographic clustering of voltage maps at GFP peaks, and explored the association

between the degree to which topography remains stable relative to GFP and the proportion of

topographic variance explained overall by microstates in the EEG time series.

Methods

Participants

227 participants were recruited to participate as part of the MPI-Leipzig Mind-Brain-

Body study (MPILMBB; Babayan et al., 2019). Participants were recruited as part of two

separate age cohorts. The younger age cohort was between 20–35 years old (N = 153, 45

females, age M = 25.1 years, SD = 3.1) and the older age cohort was between 59–77 years old (N

= 74, 37 females, age M = 67.6 years, SD = 4.7). Participants underwent an extensive medical

and psychological screening procedure before inclusion (see Babayan et al., 2019). Resting EEG

was recorded from 216 of these participants. All participants were tested at the Day Clinic for

Cognitive Neurology of the University Clinic Leipzig and the Max Planck Institute for Human

Cognitive and Brain Sciences (MPI CBS) in Leipzig, Germany. Written informed consent was

obtained prior to participating in the study, and all participants received monetary compensation.

The study was carried out in accordance with the Declaration of Helsinki and the study protocol

was approved by the ethics committee of the University of Leipzig (reference #154/13-ff).

Procedure

16 minutes of resting EEG was acquired from 216 participants in a sound attenuated

chamber prior to the administration of psychological questionnaires and assessments including a

psychiatric interview (SCID; Wittchen, Wunderlich, and Gruschwitz, 1997). Each EEG

recording was divided into 16 contiguous 1-minute blocks, with two conditions interleaved, eyes

closed (EC) and eyes open (EO), beginning with the EC condition. Presentation software
(Neurobehavioral Systems Inc., USA) was used to indicate changes between blocks. Participants

were instructed to fixate on a black cross presented on a white background during the EO blocks.

EEG Data Collection and Processing

Resting EEG was recorded from a 62-channel active electrode cap (ActiCAP, Brain

Products GmbH, Germany), with 61 channels in the international 10-20 system arrangement and

one additional electrode below the right eye recording vertical eye movements. The reference

electrode was located at electrode position FCz and the ground was located at the sternum.

Electrode impedance were kept below 5 kΩ. Data were acquired with a BrainAmp MR plus

amplifier (Brain Products GmbH, Germany) at an amplitude resolution of 0.1 µV and sampling

rate of 2500 Hz. EEG were bandpass filtered online between 0.015 Hz and 1k Hz, subsequently

downsampled offline to 250 Hz, and bandpass filtered between 1–45 Hz (Butterworth filter, filter

order 4). 8-minute EC and EO epochs were separately concatenated.

Preprocessed EEG recordings (Babayan et al., 2019) were made available for use to

interested researchers on a data-sharing repository (https://ftp.gwdg.de/pub/misc/MPI-

Leipzig_Mind-Brain-Body-LEMON/). Portions of these preprocessed data have already been

utilized in two investigations of oscillatory dynamics in resting EEG (i.e., Mahjoory et al., 2019;

Schaworonkow and Nikulin, 2019). As reported by Babayan and colleagues (2019), outlier

channels with poor signal quality, extreme peak-to-peak deflections, or large bursts of high

frequency activity, were excluded based on visual inspection. Principal component analysis

(PCA) was used to reduce the dimensionality of the data by keeping components that explain

95% of the total data variance. Infomax independent component analysis (ICA) was used to

remove components reflecting eye movements, eye blinks, or heartbeat related signal

contaminants. Remaining independent components (M = 21.4 components, range: 14 – 28) were

then reconstructed and projected back to sensor space. 13 participants were excluded due to

missing event information, different sampling rate, or insufficient data quality.

Following collection of the 406 preprocessed EEG recordings from 203 participants from

the data-sharing repository, missing electrodes were interpolated based on spherical spline

interpolation to a 64-channel montage and average-referenced using the Cartool software toolbox

version 3.7 (Brunet, Murray, and Michel, 2011). To focus my analyses on healthy individuals, I

excluded 12 additional participants from further analysis because the psychiatric interview

conducted at the second assessment identified current psychological diagnoses (e.g., substance

abuse or unspecified hallucinations) that were missed during initial screening.

Global Field Power and Topographic Dissimilarity

Global field power (GFP) and topographic dissimilarity (DISS) were calculated for

samples of the EEG time series. GFP is a reference-independent measure of voltage potential

(µV) that quantifies the strength of the scalp electric field at a given sample of the recording and

is equivalent to the standard deviation of amplitude across the average-referenced electrode

montage (Skrandies, 1990; Murray et al., 2008). GFP was calculated for all samples of the EEG

time series for the remaining 191 participants (382 total recordings). Furthermore, the difference

in topography between temporally adjacent voltage maps was calculated based on the measure of

global topographic dissimilarity (Skrandies, 1990; Murray et al., 2008). DISS is quantified as the

square root of the mean of squared differences between GFP normalized electrodes, and reflects

a measure of the overall difference in spatial configuration between two electric fields,

independent of their strength. The DISS was calculated between the voltage map of each time

series sample (sample t) and the map of the preceding sample (sample t - 1). This quantified the

topographic (dis)similarity between voltage maps in subsequent moments. Figure 1 depicts the
calculation of GFP and DISS from 1 second of EEG from a participant chosen at random. High

DISS indicates that the voltage map of the current sample and the preceding sample were

dissimilar in spatial configuration. Values of GFP and DISS were log-transformed and the

correlation between the strength of the electric field and dissimilarity of temporally adjacent

voltage maps was calculated for the EEG time series of each individual recording. Correlations

were compared between EC and EO conditions with a paired t-test after Fisher r to z

transformation.

Topographic Segmentation and Microstate Parameter Estimation

I conducted topographic segmentation of the EEG in the 191 participants (382

recordings) to identify periods of quasi-stable scalp voltage configurations. This was achieved

through an adapted k-means clustering method, implemented in Cartool (Brunet et al., 2011),

that determines the optimal number of clusters (k) that can account for the greatest global

explained variance (GEV) in the time-series using the smallest number of representative

topographical maps (Murray et al., 2008; Michel et al., 2009). First, topographic voltage maps

were generated at the local maxima (peaks) in the GFP time series. This is illustrated in Figure 2,

which depicts the time series of voltage maps from 1 second of EEG with maps at local GFP

maxima indicated. This was done separately for each individual recording. The initial maps were

generated at the GFP peaks in order to maximize signal-to-noise ratio. Because GFP peaks tend

to reflect moments of high global neuronal synchronization, they are thought to provide optimal

representations of the momentary quasi-stable voltage topography (Koenig and Brandeis, 2016).

Clustering of voltage maps. Each iteration of k-means clustering proceeded as follows.

A subset of 1 to 12 maps (k = [1:12]) was randomly selected from the total set of voltage maps

for a given participant and condition (EC or EO recordings), to use as initial centroids for
clustering. The spatial correlation between the k centroid maps and the remaining voltage maps

were then computed. GFP maps were assigned to the centroid with which they had the highest

spatial correlation, creating k clusters of maps. Maps were only assigned to a cluster if the spatial

correlation with the centroid map exceeded 0.5, and correlation values were based on the relative

topographical configuration (but not the polarity) of the maps. After all correlations, new

centroid maps were created by averaging the constituent maps assigned to a given k cluster. The

set of remaining maps not assigned to a cluster were then compared to the recomputed

(averaged) centroids and assigned again based on the correlation criterion. This process

continued iteratively until the global explained variance (GEV) between the average centroids

and the maps converged to a limit.

This entire procedure was repeated 100 times for each k, with a new subset of k centroids

selected for each iteration. After the 100 iterations, the k set of centroids with maximal GEV

were identified. This was repeated for each level of k = [1:12]. Across all levels of k for an

individual within each condition, the optimal number of k clusters from the maximal GEV

centroids was determined using a metacriterion defined by 7 independent optimization criteria

(see Custo et al., 2017; Brechet et al., 2019). Finally, I collected centroids of maps based on the

optimal number of k clusters for all recordings.

Clustering of subject-level centroids. In a second step, I conducted k-means clustering

on the centroids identified through clustering of the subject-level voltage maps. This was done to

identify the optimal global clusters that best explain the subject-level representative cluster

centroids. A set of k = [1:15] maps were randomly selected from the set of subject-level

topographies and used as random centroids for clustering. For each level of k, 200 iterations were

conducted, until the GEV converged to a limit and the k centroids with the maximal GEV were
selected. After all iterations, the optimal number of k global clusters was determined using the

optimization metacriterion, resulting in a set of k global centroids that best represent the subject-

level topographic configurations.

Parameterization of the microstate time series. The global centroids were then fitted

back to the original EEG recordings to derive time series sequences of microstates. All samples

of an individual subject’s continuous EEG were categorized by the global topography that

demonstrated the highest spatial correlation between the sample-wise voltage map and the

centroids of global clusters. EEG samples that had low spatial correlation (< .5) with all global

centroids were left unassigned to a microstate centroid. Polarity was again ignored during

centroid assignment. Global explained variance (GEV) was calculated for the time series

sequence of microstates and is the percentage of observed topographic variance explained by a

particular microstate configuration (Murray et al., 2008). The sum of GEV for all microstate

configurations was calculated as the total GEV for each individual and condition.

Results

Global field power and topographic temporal stability. Within-person correlations

were calculated between the GFP of each sample and the topographic dissimilarity (DISS)

between temporally adjacent (sample t and sample t – 1) voltage maps for the EEG time series of

each recording. Values of GFP and DISS were log-transformed. Figure 3 depicts scatterplots

demonstrating the association between GFP and DISS, and log-transformed GFP and DISS, for 4

eyes-closed individual recordings chosen at random. Figure 4 depicts scatterplots for 4 eyes-open

recordings chosen at random. At points of high GFP, the DISS of adjacent maps is low. The

reverse, however, does not appear to be true. At points of low GFP, DISS demonstrated

considerable variability.
 Strong negative correlations between log-transformed GFP and DISS were observed on

average in both the EC (M = -.658, SD = 0.060, 95% CI [-.667, -.650]) and EO (M = -.663, SD =

.065, 95% CI [-.672, -.654]) conditions.1 One individual was a clear outlier (EC r = -.261 and EO

r = -.311), and there were two other outlier EO recordings with r = -.293 and r = -.332,

respectively. All the remaining correlations were strongly negative and ranged from r = -.453 to -

.766 for the EC condition, and r = -.426 to -.766 for the EO condition. Figure 5 depicts the mean

correlations for each condition and range of correlations among individuals. Fisher z-transformed

correlations did not significantly differ between conditions, t (190) = 1.338, p = .182, but were

positively correlated within-persons across conditions (r = .582, p < .001, 95% CI [.480, .669]).

These findings confirm the robust association between GFP and topographic temporal stability in

a large sample of healthy adults. The basic premise of microstate segmentation that voltage maps

at local maxima of GFP are strong indicators of momentary stability in topographic

configuration is therefore well supported by empirical data.

Topographic segmentation. k-means clustering of resting EEG revealed between 4 and

8 topographies (M = 5.08, SD = 0.94) as the optimal number of k clusters for each individual

EEG recording across both EC and EO conditions (totaling 1940 topographies). The second

round of k-means clustering of the 1940 individual subject centroid topographies identified five

global clusters that together explained 85.03% of the GEV among individual subject centroids

and were selected as the optimal number of global clusters based on the optimization

metacriterion. The five global cluster centroids thus appeared to be a good representation of the

1
The mean correlations were nearly identical when including the 12 individuals originally
excluded from the sample. Correlations for all 203 individuals were large on average in both the
EC (M = -.657, SD = 0.060, 95% CI [-.666, -.649]) and EO (M = -.663, SD = 0.064, 95% CI [-
.672, -.654]) conditions.
most common topographic patterns observed among participants. Figure 6 depicts the five global

cluster centroids which I named microstates A–E and the 1938 (2 went unassigned) individual

subject cluster topographies grouped according to their global cluster membership.

The five global cluster centroids were then fit to the continuous EEG time series and

successfully assigned (correlation > .5) to 85.94% (SD = 11.1%) of EEG samples on average

across all 382 recordings. Four participants were excluded from further analyses because a large

percentage of the voltage maps for the continuous EEG time-series (> 45%) could not be

assigned to a global cluster on average across conditions. One of these individuals was an outlier

in terms of their within-person correlation (r = -.332) between log-transformed GFP and DISS.

Across the 374 remaining EEG recordings, the five global microstate topographies in total

explained 65.03% (SD = 6.13%) of GEV in the voltage maps of the EEG time series of the eyes-

closed condition, and 60.99% (SD = 5.62%) of GEV in the eyes-open condition, on average

across individuals.

The total topographic variance explained (GEV) in an individual’s EEG time series by all

microstate configurations was furthermore associated with the strength of correlation between

log-transformed GFP and topographic dissimilarity. Three observations were removed prior to

examining these associations (correlations between GFP and DISS weaker than r = -.340),

leaving 371 total observations (186 EC and 185 EO). Fisher z-transformed correlations between

GFP and DISS were significantly correlated with total GEV for the EC (r = -.282, p < .001, 95%

CI [-.410, -.144]) and EO (r = -.473, p < .001, 95% CI [ -.578, -.353]) conditions.2 That is, the

more that an individual’s EEG exhibits topographic stability during moments of high GFP, the

2
Including the three outlier observations did not appear to change the mean correlation for the
EC condition (r = -.282, p < .001, 95% CI [-.409, -.144]) but did unduly influence the magnitude
of the correlation in the EO condition (r = -.420, p < .001, 95% CI [ -.531, -.294]).
better microstates explain the EEG time series. Figure 7 depicts the scatterplot demonstrating this

association for each condition.

Interestingly, this association appeared to be stronger in the case of the EO condition.

This was not the case, however, when tested directly in a multi-level mixed effects model

examining the difference between conditions in z-transformed correlations. A random intercept

was included for individuals and the model was estimated with restricted maximum likelihood in

R package lme4. GEV was significantly predicted by the correlation between GFP and DISS,

F(1, 334.11) = 70.911, p < .001, and conditions differed in their total GEV, F(1, 224.01) = 5.052,

p = .026, but there was no significant interaction between condition and the correlation between

GFP and DISS, F(1, 224.14) = 1.388, p = .240. This indicated that there was not enough

evidence to determine whether the strength of the association differed between conditions.

Together, these effects explained 22% of the total variance (R2 = .224) in GEV.

Discussion

The basic premise underlying segmentation of broadband EEG into microstates based on

momentary periods of topographic stability is well supported by empirical data in the present

study. 191 healthy adults demonstrated large and consistent within-person negative correlations

between log-transformed GFP of samples of their EEG time series and the topographic

dissimilarity of temporally adjacent voltage maps. The time series succession of EEG voltage

maps thus exhibits momentary topographic stability when the electric field strength is high.

When the field strength is low, however, topographic stability varies considerably. The

association between GFP and topographic stability was observed across thousands of samples of

the EEG time series in every recording included in the present study. Yet, there was some

variability within-persons in terms of the magnitude of these correlations. Only 34% of the
variance was shared between eyes-closed and eyes-open conditions, suggesting that the

association between GFP and topographic stability appears to quantify aspects of spatiotemporal

dynamics of EEG that vary within individuals.

Clustering of EEG voltage maps at GFP peaks led to the identification of 5 data-driven

microstate configurations that explained more than 60% of the total topographic variance when

fit to individuals’ EEG time series. The total GEV explained by microstates was associated with

the strength of coupling between GFP and topographic stability. Roughly 20% of variance in

GEV was explained by an individuals within-person correlation between GFP and DISS. The

more that an individual’s EEG exhibits stability surrounding periods of high GFP, the better the

topography of the EEG time series was explained in terms of a limited number of exemplar

microstate configurations. Therefore, the association between GFP and topographic stability

appears to quantify the prevalence of microstate-related spatiotemporal dynamics in the ongoing

EEG time series. One intriguing possibility involves quantifying this association for individuals

as a means of screening the quality of data to improve the global fit of microstate-based

clustering solutions.

Microstate clustering is commonly applied to voltage maps generated at the GFP peaks of

the EEG time series because of the well-reasoned assumption that these instances maximize

signal-to-noise ratio and provide optimal representations of the momentary quasi-stable voltage

topography. Furthermore, moments of high GFP reflect instances of strong synchronized activity

of coordinated networks of brain generators. While the present findings provide additional

empirical support for the notion that clustering at GFP peaks produces microstate configurations

that optimally represent moments of topographic stability, this does not imply that clustering at

other moments of the EEG time series will not produce representative cluster solutions. After all,
there was considerable variability in the similarity of adjacent voltage maps at moments when

GFP was lower. Common topographic patterns may therefore still be identified based on

clustering applied to other moments of the EEG time series, but future studies will need to

examine this question directly.

In total, the present findings provide strong empirical support for the proposal that

periods of topographic stability can be identified in the time series succession of voltage maps

corresponding to common microstate configurations, and that moments of high field strength are

optimal representations of quasi-stable electric field topography. The activity of phase-

synchronized neuronal networks thus persists in a coherent configuration of active neural

generators for brief moments before quickly transitioning to different configurations. This agrees

with the premise underlying microstate segmentation (Lehmann et al., 1987) that neurocognitive

networks manifest cognitive function in real time through sequences of coordinated brain states

(Bressler and Tognoli, 2006; Bressler and Kelso, 2016). Topographic stability during moments

of high field strength therefore appears to be a basic property of the spontaneous spatiotemporal

dynamics of the neuroelectric field.

 References

Babayan, A., Erbey, M., Kumral, D., Reinelt, J. D., Reiter, A. M. F., Röbbig, J., … Villringer, A.
(2019). A mind-brain-body dataset of MRI, EEG, cognition, emotion, and peripheral
physiology in young and old adults. Scientific Data, 6, 180308.

Brechet, L., Brunet, D., Birot, G., Gruetter, R., Michel, C. M., and Jorge, J. (2019). Capturing the
spatiotemporal dynamics of self-generated, task-initiated thoughts with EEG and fMRI.
Neuroimage, 194, 82-92.

Bressler, S. L., and Kelso, S. J. A. (2016). Coordination dynamics in cognitive neuroscience.

Frontiers in Human Neuroscience, 10:397.

Bressler, S. L., and Tognoli, E. (2006). Operational principles of neurocognitive networks.

International Journal of Psychophysiology, 60(2), 139-148.

Britz, J., Van De Ville, D., and Michel, C. M. (2010). BOLD correlates of EEG topography
reveal rapid resting-state network dynamics. Neuroimage, 52, 1162-1170.

Brunet, D., Murray, M. M., and Michel, C. M. (2011). Spatiotemporal analysis of multichannel
EEG: CARTOOL. Computational Intelligence and Neuroscience, 2011, 2.

Custo, A., Van De Ville, D., Wells, W. M., Tomescu, M. I., Brunet, D., and Michel, C. M.
(2017). Electroencephalographic resting-state networks: Source localization of
microstates. Brain Connectivity, 7(10), 671-682.

Khanna, A., Pascual-Leone, A., Michel, C. M., and Farzan, F. (2015). Microstates in resting-
state EEG: current status and future directions. Neuroscience Biobehavioral Reviews, 49,
105-113.

Koenig, T., and Brandeis, D. (2016). Inappropriate assumptions about EEG state changes and
their impact on the quantification of EEG state dynamics. Neuroimage, 125, 1104-1106.

Lehmann, D., Ozaki, H., and Pal, I. (1987). EEG alpha map series: brain micro-states by space-
oriented adaptive segmentation. Electroencephalography and Clinical Neurophysiology,
67, 271-288.
Mahjoory, K., Cesnaite, E., Hohlefeld, F. U., Villringer, A., and Nikulin, V. V. (2019). Power and
temporal dynamics of alpha oscillations at rest differentiate cognitive performance
involving sustained and phasic cognitive control. Neuroimage, 188, 135-144.

Lehmann, D., Strik, W. K., Henggeler, B., Koenig, T., and Koukkou , M. (1998). Brain electric
microstates and momentary concious mind states as building blocks of spontaneous
thinking: I Visual imagery and abstract thoughts. International Journal of
Psychophysiology, 29(1), 1-11.

Michel, C. M., and Koenig, T. (2018). EEG microstates as a tool for studying the temporal
dynamics of whole-brain neuronal networks: A review. Neuroimage, 180, 577-593.

Michel, C. M., Koenig, T., and Brandeis, D. (2009). Electric neuroimaging in the time domain.
In C. M. Michel et al. (Eds.), Electrical Neuroimaging (pp. 111–144). Cambridge, UK:
Cambridge University Press.

Murray, M. M., Brunet, D., and Michel, C. M. (2008). Topographic ERP analyses: A step-by-
step tutorial review. Brain Topography, 4, 249-264.

Seitzman, B. A., Abell, M., Bartley, S. C., Erickson, M. A., Bolbecker, A. R., and Hetrick, W. P.
(2017). Cognitive manipulation of brain electric microstates. Neuroimage, 146, 533-543.

Schaworonkow, N., and Nikulin, V. V. (2019). Spatial neuronal synchronization and the
waveform of oscillations: Implications for EEG and MEG. PloS Computational Biology,
15(5): e1007055.

Skrandies, W. (1990). Global field power and topographic similarity. Brain Topography, 3(1),
137-141.

Vaughan, H. G. (1982). The neural origins of human event-related potentials. Annals of the New
York Academy of Sciences, 388(1), 125-138.

Wackermann, J., Lehmann, D., Michel, C. M., and Strik, W. K. (1993). Adaptive segmentation
of spontaneous EEG map series into spatially defined microstates. International Journal
of Psychophysiology, 14, 269-283.
Wittchen, H.-U., Zaudig, M., and Fydrich, T. (1997). SKID. Strukturiertes klinisches interview
für DSM-IV. Achse I und II. Handanweisung. Göttingen: Hogrefe.
Figure 1: One second of 64-channel eyes-closed resting EEG (sampled at 250 Hz) from a
recording chosen at random. The global field power (GFP) is calculated from the multichannel
EEG and reflects a measure of the ongoing strength of the electric field. The periodicity of peaks
in GFP coincide with oscillations at the dominant EEG frequency. Topographic dissimilarity
(DISS) is also calculated and reflects a measure of the difference in spatial configuration
between electric fields of temporally adjacent (sample t and sample t – 1) voltage maps,
independent of their strength. GFP appears inversely correlated with topographic dissimilarity.
Figure 2: Rows depict the time series succession of voltage maps from left to right of 1 second of
64-channel eyes-closed resting EEG (sampled at 250 Hz) from a recording chosen at random.
Voltage maps are 2D isometric projections with nasion upwards. Voltage maps are highlighted at
the local maxima in the global field power (GFP). The topography generally appears quasi-stable
for several samples surrounding GFP peaks.
Figure 3: Scatterplots depicting samples of global field power (GFP) by topographic dissimilarity
between temporally adjacent (sample t and sample t -1) voltage maps for four eyes-closed EEG
recordings selected at random. The left column depicts raw values and the right column depicts
the corresponding log-transformed values and within-person correlation coefficients.
Figure 4: Scatterplots depicting samples of global field power (GFP) by topographic dissimilarity
between temporally adjacent (sample t and sample t -1) voltage maps for four eyes-open EEG
recordings selected at random. The left column depicts raw values and the right column depicts
the corresponding log-transformed values and within-person correlation coefficients.
Figure 5: Means are plotted for within-person correlations between log-transformed global field
power (GFP) and topographic dissimilarity for eyes-closed and eyes-open conditions. Observed
participant (n = 191) correlations are plotted as dots. Errors bars are 95% CI of the mean.
Figure 6: Five global cluster centroids were identified from k-means clustering during 8-min of
eyes-closed (EC) and 8-min of eyes-open (EO) rest. 1938 cluster centroids derived from k-means
clustering of voltage maps at GFP peaks from 382 individual subject recordings are shown
grouped according to their global cluster membership. Voltage topographies are 2D isometric
projections with nasion upwards. Each global topography is the centroid of respective clusters of
maps.
Figure 7: Scatterplot depicts the total global explained variance (GEV) by the Fisher z-
transformed correlation between log GFP and DISS for the eyes-closed (EC; n = 186, r = -0.282)
and eyes-open (EO; n = 185, r = -0.473) conditions.