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Why do people sometimes report that they remember dreams, while at other times they recall no experience? Despite the interest in dreams that may happen during the night, it has remained unclear which brain states determine whether these conscious experiences will occur and what prevents us from waking up during these episodes. Here we address this issue by comparing the EEG activity preceding awakenings with recalled vs. no recall of dreams using the EEG microstate approach. This approach characterizes transiently stable brain states of sub-second duration that involve neural networks with nearly synchronous dynamics. We found that two microstates (3 and 4) dominated during NREM sleep compared to resting wake. Further, within NREM sleep, microstate 3 was more expressed during periods followed by dream recall, whereas microstate 4 was less expressed. Source localization showed that microstate 3 encompassed the medial frontal lobe, whereas microstate 4 involved the occipital cortex, as well as thalamic and brainstem structures. Since NREM sleep is characterized by low-frequency synchronization, indicative of neuronal bistability, we interpret the increased presence of the “frontal” microstate 3 as a sign of deeper local deactivation, and the reduced presence of the “occipital” microstate 4 as a sign of local activation. The latter may account for the occurrence of dreaming with rich perceptual content, while the former may account for why the dreaming brain may undergo executive disconnection and remain asleep. This study demonstrates that NREM sleep consists of alternating brain states whose temporal dynamics determine whether conscious experience arises.

Using electroencephalography (EEG) to elucidate the spontaneous activation of brain resting-state networks (RSNs) is nontrivial as the signal of interest is of low amplitude and it is difficult to distinguish the underlying neural sources. Using the principles of electric field topographical analysis, it is possible to estimate the meta-stable states of the brain (i.e., the resting-state topographies, so-called microstates). We estimated seven resting-state topographies explaining the EEG data set with k-means clustering (N = 164, 256 electrodes). Using a method specifically designed to localize the sources of broadband EEG scalp topographies by matching sensor and source space temporal patterns, we demonstrated that we can estimate the EEG RSNs reliably by measuring the reproducibility of our findings. After subtracting their mean from the seven EEG RSNs, we identified seven state-specific networks. The mean map includes regions known to be densely anatomically and functionally connected (superior frontal, superior parietal, insula, and anterior cingulate cortices). While the mean map can be interpreted as a “router,” crosslinking multiple functional networks, the seven state-specific RSNs partly resemble and extend previous functional magnetic resonance imaging-based networks estimated as the hemodynamic correlates of four canonical EEG microstates.

•Microstate sequences are best studied considering high-order temporal characteristics.•We present a method, Microsynt, to study the syntax of microstate sequences.•Microstate sequences are not random but favor low entropy words with binary loops.•During consciousness, externally oriented microstate binary loops are prominent.•Internally generated binary loops become prominent during unconscious states. Microstates represent electroencephalographic (EEG) activity as a sequence of switching, transient, metastable states. Growing evidence suggests the useful information on brain states is to be found in the higher-order temporal structure of these sequences. Instead of focusing on transition probabilities, here we propose “Microsynt”, a method designed to highlight higher-order interactions that form a preliminary step towards understanding the syntax of microstate sequences of any length and complexity. Microsynt extracts an optimal vocabulary of “words” based on the length and complexity of the full sequence of microstates. Words are then sorted into classes of entropy and their representativeness within each class is statistically compared with surrogate and theoretical vocabularies. We applied the method on EEG data previously collected from healthy subjects undergoing propofol anesthesia, and compared their “fully awake” (BASE) and “fully unconscious” (DEEP) conditions. Results show that microstate sequences, even at rest, are not random but tend to behave in a more predictable way, favoring simpler sub-sequences, or “words”. Contrary to high-entropy words, lowest-entropy binary microstate loops are prominent and favored on average 10 times more than what is theoretically expected. Progressing from BASE to DEEP, the representation of low-entropy words increases while that of high-entropy words decreases. During the awake state, sequences of microstates tend to be attracted towards “A – B – C” microstate hubs, and most prominently A – B binary loops. Conversely, with full unconsciousness, sequences of microstates are attracted towards “C – D – E” hubs, and most prominently C – E binary loops, confirming the putative relation of microstates A and B to externally-oriented cognitive processes and microstate C and E to internally-generated mental activity. Microsynt can form a syntactic signature of microstate sequences that can be used to reliably differentiate two or more conditions.

This paper describes methods to analyze the brain's electric fields recorded with multichannel Electroencephalogram (EEG) and demonstrates their implementation in the software CARTOOL. It focuses on the analysis of the spatial properties of these fields and on quantitative assessment of changes of field topographies across time, experimental conditions, or populations. Topographic analyses are advantageous because they are reference independents and thus render statistically unambiguous results. Neurophysiologically, differences in topography directly indicate changes in the configuration of the active neuronal sources in the brain. We describe global measures of field strength and field similarities, temporal segmentation based on topographic variations, topographic analysis in the frequency domain, topographic statistical analysis, and source imaging based on distributed inverse solutions. All analysis methods are implemented in a freely available academic software package called CARTOOL. Besides providing these analysis tools, CARTOOL is particularly designed to visualize the data and the analysis results using 3-dimensional display routines that allow rapid manipulation and animation of 3D images. CARTOOL therefore is a helpful tool for researchers as well as for clinicians to interpret multichannel EEG and evoked potentials in a global, comprehensive, and unambiguous way.

Electroencephalographic source localisation (ESL) of interictal epileptiform discharges is a valuable tool for presurgical evaluation of pharmacoresistant focal epilepsy. Various forward models, inverse solutions algorithms, and software packages have been published. However, clinical validation studies are based on heterogenous end points and study cohorts. To allow comparison of different interictal ESL methods within one standardised dataset, we provide deidentified clinical data of 44 well-characterised patients with pharmacoresistant focal epilepsy and a first resective surgery, validated by 12-month postsurgical outcome. Thirty patients had favourable outcomes, including 28 with complete seizure freedom, indicating that the epileptogenic zone was correctly estimated. For each patient, pre-processed individual structural MRI, 257-channel EEG averages of homologous discharges, postsurgical structural neuroimaging, and detailed clinical and technical information are given. In patients with favourable outcomes, source maxima of averaged discharges were <10 mm remote from the resection in 67% and within a sublobe affected by the surgery in 83%. Future validation studies of new ESL approaches can be compared to this benchmark.

Though commonly held that multisensory experiences enrich our memories and that memories influence ongoing sensory processes, their neural mechanisms remain unresolved. Here, electrical neuroimaging shows that auditory–visual multisensory experiences alter subsequent processing of unisensory visual stimuli during the same block of trials at early stages poststimulus onset and within visual object recognition areas. We show this with a stepwise analysis of scalp-recorded event-related potentials (ERPs) that statistically tested (1) ERP morphology and amplitude, (2) global electric field power, (3) topographic stability of and changes in the electric field configuration, and (4) intracranial distributed linear source estimations. Subjects performed a continuous recognition task, discriminating repeated vs. initial image presentations. Corresponding, but task-irrelevant, sounds accompanied half of the initial presentations during a given block of trials. On repeated presentations within a block of trials, only images appeared, yielding two situations—the image's prior presentation was only visual or with a sound. Image repetitions that had been accompanied by sounds yielded improved memory performance accuracy (old or new discrimination) and were differentiated as early as ∼ 60–136 ms from images that had not been accompanied by sounds through generator changes in areas of the right lateral–occipital complex (LOC). It thus appears that unisensory percepts trigger multisensory representations associated with them. The collective data support the hypothesis that perceptual or memory traces for multisensory auditory–visual events involve a distinct cortical network that is rapidly activated by subsequent repetition of just the unisensory visual component.

The temporal structure of self-generated cognition is a key attribute to the formation of a meaningful stream of consciousness. When at rest, our mind wanders from thought to thought in distinct mental states. Despite the marked importance of ongoing mental processes, it is challenging to capture and relate these states to specific cognitive contents. In this work, we employed ultra-high field functional magnetic resonance imaging (fMRI) and high-density electroencephalography (EEG) to study the ongoing thoughts of participants instructed to retrieve self-relevant past episodes for periods of 22sec. These task-initiated, participant-driven activity patterns were compared to a distinct condition where participants performed serial mental arithmetic operations, thereby shifting from self-related to self-unrelated thoughts. BOLD activity mapping revealed selective enhanced activity in temporal, parietal and occipital areas during the memory compared to the mental arithmetic condition, evincing their role in integrating the re-experienced past events into conscious representations during memory retrieval. Functional connectivity analysis showed that these regions were organized in two major subparts, previously associated to “scene-reconstruction” and “self-experience” subsystems. EEG microstate analysis allowed studying these participant-driven thoughts in the millisecond range by determining the temporal dynamics of brief periods of stable scalp potential fields. This analysis revealed selective modulation of occurrence and duration of specific microstates in the memory and in the mental arithmetic condition, respectively. EEG source analysis revealed similar spatial distributions of the sources of these microstates and the regions identified with fMRI. These findings imply a functional link between BOLD activity changes in regions related to a certain mental activity and the temporal dynamics of mentation, and support growing evidence that specific fMRI networks can be captured with EEG as repeatedly occurring brief periods of integrated coherent neuronal activity, lasting only fractions of seconds.

In this tutorial review, we detail both the rationale for as well as the implementation of a set of analyses of surface-recorded event-related potentials (ERPs) that uses the reference-free spatial (i.e. topographic) information available from high-density electrode montages to render statistical information concerning modulations in response strength, latency, and topography both between and within experimental conditions. In these and other ways these topographic analysis methods allow the experimenter to glean additional information and neurophysiologic interpretability beyond what is available from canonical waveform analyses. In this tutorial we present the example of somatosensory evoked potentials (SEPs) in response to stimulation of each hand to illustrate these points. For each step of these analyses, we provide the reader with both a conceptual and mathematical description of how the analysis is carried out, what it yields, and how to interpret its statistical outcome. We show that these topographic analysis methods are intuitive and easy-to-use approaches that can remove much of the guesswork often confronting ERP researchers and also assist in identifying the information contained within high-density ERP datasets.

Microstate analysis of resting-state EEG is a unique data-driven method for identifying patterns of scalp potential topographies, or microstates, that reflect stable but transient periods of synchronized neural activity evolving dynamically over time. During infancy – a critical period of rapid brain development and plasticity – microstate analysis offers a unique opportunity for characterizing the spatial and temporal dynamics of brain activity. However, whether measurements derived from this approach (e.g., temporal properties, transition probabilities, neural sources) show strong psychometric properties (i.e., reliability) during infancy is unknown and key information for advancing our understanding of how microstates are shaped by early life experiences and whether they relate to individual differences in infant abilities. A lack of methodological resources for performing microstate analysis of infant EEG has further hindered adoption of this cutting-edge approach by infant researchers. As a result, in the current study, we systematically addressed these knowledge gaps and report that most microstate-based measurements of brain organization and functioning except for transition probabilities were stable with four minutes of video-watching resting-state data and highly internally consistent with just one minute. In addition to these results, we provide a step-by-step tutorial, accompanying website, and open-access data for performing microstate analysis using a free, user-friendly software called Cartool. Taken together, the current study supports the reliability and feasibility of using EEG microstate analysis to study infant brain development and increases the accessibility of this approach for the field of developmental neuroscience.

The error-related negativity (ERN) is a negative deflection in the electroencephalography (EEG) waveform at frontal-central scalp sites that occurs after error commission. The relationship between the ERN and broader patterns of brain activity measured across the entire scalp that support error processing during early childhood is unclear. We examined the relationship between the ERN and EEG microstates – whole-brain patterns of dynamically evolving scalp potential topographies that reflect periods of synchronized neural activity – during both a go/no-go task and resting-state in 90, 4-8-year-old children. The mean amplitude of the ERN was quantified during the -64 to 108 millisecond (ms) period of time relative to error commission, which was determined by data-driven microstate segmentation of error-related activity. We found that greater magnitude of the ERN associated with greater global explained variance (GEV; i.e., the percentage of total variance in the data explained by a given microstate) of an error-related microstate observed during the same -64 to 108 ms period (i.e., error-related microstate 3), and to greater anxiety risk as measured by parent-reported behavioral inhibition. During resting-state, six data-driven microstates were identified. Both greater magnitude of the ERN and greater GEV values of error-related microstate 3 associated with greater GEV values of resting-state microstate 4, which showed a frontal-central scalp topography. Source localization results revealed overlap between the underlying neural generators of error-related microstate 3 and resting-state microstate 4 and canonical brain networks (e.g., ventral attention) known to support the higher-order cognitive processes involved in error processing. Taken together, our results clarify how individual differences in error-related and intrinsic brain activity are related and enhance our understanding of developing brain network function and organization supporting error processing during early childhood.