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Nikulin, V.

Publications and source records attributed to Nikulin, V..

34 records · Page 2Linked to original sources

Frequency-specific changes in prefrontal activity associated with maladaptive belief updating in volatile environments in euthymic bipolar disorder

Bipolar disorder (BD) involves altered reward processing and decision-making, with inconsistencies across studies. Here, we integrated hierarchical Bayesian modelling with magnetoencephalography (MEG) to characterise maladaptive belief updating in this condition. First, we determined if previously reported increased learning rates in BD stem from a heightened expectation of environmental changes. Additionally, we examined if this increased expectation speeds up belief updating in decision-making, associated with modulation of rhythmic neural activity within the prefrontal, orbitofrontal, and anterior cingulate cortex (PFC, OFC, ACC). Twenty-two 22 euthymic BD and 27 healthy control (HC) participants completed a reward-based motor decision-making task in a volatile setting. Hierarchical Bayesian modelling revealed BD participants anticipated greater environmental volatility, resulting in a more stochastic mapping from beliefs to actions and paralleled by lower win rates and a reduced tendency to repeat rewarded actions than HC. Despite this, BD individuals adjusted their expectations of action-outcome contingencies more slowly, but both groups invigorated their actions similarly. On a neural level, while healthy individuals exhibited an alpha-beta suppression and gamma increase during belief updating, BD participants showed dampened effects, extending across the PFC, OFC, and ACC regions. This was accompanied by an abnormally increased beta-band directed information flow in BD. Overall, the results suggest euthymic BD individuals anticipate environmental change without adequately learning from it, contributing to maladaptive belief updating. Alterations in frequency-domain amplitude and functional connectivity within the PFC, OFC, and ACC during belief updating underlie the computational effects and could serve as potential indicators for predicting relapse in future research.

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Linking brain-heart interactions to emotional arousal in immersive virtual reality

The subjective experience of emotions is linked to the contextualized perception and appraisal of changes in bodily (e.g., heart) activity. Increased emotional arousal has been related to attenuated high-frequency heart rate variability (HF-HRV), lower EEG parieto-occipital alpha power, and higher heartbeat-evoked potential (HEP) amplitudes. We studied emotional arousal-related brain- heart interactions using immersive virtual reality (VR) for naturalistic yet controlled emotion induction. 29 healthy adults (13 women, age: 26{+/-}3) completed a VR experience that included rollercoasters while EEG and ECG were recorded. Continuous emotional arousal ratings were collected during a video replay immediately after. We analyzed emotional arousal-related changes in HF-HRV as well as in BHIs using HEPs. Additionally, we used the oscillatory information in the ECG and the EEG to model the directional information flows between the brain and heart activity. We found that higher emotional arousal was associated with lower HEP amplitudes in a left fronto- central electrode cluster. While parasympathetic modulation of the heart (HF-HRV) and parieto- occipital EEG alpha power were reduced during higher emotional arousal, there was no evidence for the hypothesized emotional arousal-related changes in bidirectional information flow between them. Whole-brain exploratory analyses in additional EEG (delta, theta, alpha, beta and gamma) and HRV (low-frequency, LF, and HF) frequency bands revealed a temporo-occipital cluster, in which higher emotional arousal was linked to decreased brain-to-heart (i.e., gamma[->]HF-HRV) and increased heart-to-brain (i.e., LF-HRV[->]gamma) information flow. Our results confirm previous findings from less naturalistic experiments and suggest a link between emotional arousal and brain-heart interactions in temporo-occipital gamma power.

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Distinguishing between- from within-site phase-amplitude coupling using antisymmetrized bispectra

Phase-amplitude coupling (PAC) is a form of cross-frequency coupling in which the amplitude of a fast oscillation is locked to the phase of a slow oscillation. PAC has been proposed as a mechanism for integrating slow large-scale networks with fast-oscillating local processes in the brain. On a signal level, PAC can be observed in single time series, reflecting local dynamics, or between two time series, potentially reflecting a functional interaction between distinct brain sites. To investigate the role of PAC as a mechanism of brain signalling, it is important to distinguish these two cases. However, when mixtures of underlying signals are observed, between-site PAC can spuriously emerge even if the true interaction is only local (within-site). This problem arises in electrophysiological recordings where mixing occurs due to volume conduction or the presence of a shared electrical reference. To address this problem, we propose to estimate PAC using the anti-symmetrized bispectrum (ASB-PAC). It has previously been shown that the cross-bispectrum can be used to measure PAC while efficiently sup-pressing Gaussian noise, and that the anti-symmetrized bispectra vanish for mixtures of independent sources. However, ASB-PAC has so far not been used to assess the presence of genuine between-site PAC. Using simulations, we here investigate the performance of different algorithms to detect PAC in a mixed signal setting as well as the performance of the same methods to distinguish genuine between-site PAC from within-site PAC. This is done in a minimal two-channel setup as well as in a more complex setting that assesses PAC on reconstructions of simulated EEG sources. We observe that bispectral PAC methods are considerably better at detecting simulated PAC in the volume conduction setting than three conventional PAC estimators. ASB-PAC achieves the highest performance in detecting genuine between-site PAC interactions while detecting the fewest spurious interactions. Using the ASB-PAC could therefore greatly facilitate the interpretation of future PAC studies when discriminating local from remote effects. We demonstrate the application of ASB-PAC on EEG data from a motor imagery experiment. Additionally, we present an upgraded version of the free ROIconnect plugin for the EEGLAB toolbox, which includes PAC and ASB-PAC metriscs based on bispectra.

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Sensorimotor brain-computer interface performance depends on signal-to-noise ratio but not connectivity of the mu rhythm in a multiverse analysis of longitudinal data

ObjectiveServing as a channel for communication with locked-in patients or control of prostheses, sensorimotor brain-computer interfaces (BCIs) decode imaginary movements from the recorded activity of the users brain. However, many individuals remain unable to control the BCI, and the underlying mechanisms are unclear. The users BCI performance was previously shown to correlate with the resting-state signal-to-noise ratio (SNR) of the mu rhythm and the phase synchronization (PS) of the mu rhythm between sensorimotor areas. Yet, these predictors of performance were primarily evaluated in a single BCI session, while the longitudinal aspect remains rather uninvestigated. In addition, different analysis pipelines were used to estimate PS in source space, potentially hindering the reproducibility of the results. ApproachTo systematically address these issues, we performed an extensive validation of the relationship between pre-stimulus SNR, PS, and session-wise BCI performance using a publicly available dataset of 62 human participants performing up to 11 sessions of BCI training. We performed the analysis in sensor space using the surface Laplacian and in source space by combining 24 processing pipelines in a multiverse analysis. This way, we could investigate how robust the observed effects were to the selection of the pipeline. Main resultsOur results show that SNR had both between- and within-subject effects on BCI performance for the majority of the pipelines. In contrast, the effect of PS on BCI performance was less robust to the selection of the pipeline and became non-significant after controlling for SNR. SignificanceTaken together, our results demonstrate that changes in neuronal connectivity within the sensorimotor system are not critical for learning to control a BCI, and interventions that increase the SNR of the mu rhythm might lead to improvements in the users BCI performance.

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Prior movement of one arm facilitates motor adaptation in the other

Many movements in daily life are embedded in motion sequences that involve more than one limb, demanding the motor system to monitor and control different body parts in quick succession. During such movements, systematic changes in the environment or the body might require motor adaptation of specific segments. However, previous motor adaptation research has focused primarily on motion sequences produced by a single limb, or on simultaneous movements of several limbs. For example, adaptation to opposing force fields is possible in unimanual reaching tasks when the direction of a prior or subsequent movement is predictive of force field direction. It is unclear, however, whether multi-limb sequences can support motor adaptation processes in a similar way. In the present study, we investigated whether reaches can be adapted to different force fields in a bimanual motor sequence when the information about the perturbation is associated with the prior movement direction of the other arm. In addition, we examined whether prior perceptual (visual or proprioceptive) feedback of the opposite arm contributes to force field-specific motor adaptation. Our key finding is that only active participation in the bimanual sequential task supports pronounced adaptation. This result suggests that active segments in bimanual motion sequences are linked across limbs. If there is a consistent association between movement kinematics of the linked and goal movement, the learning process of the goal movement can be facilitated. More generally, if motion sequences are repeated often, prior segments can evoke specific adjustments of subsequent movements. Significance statementMovements in a limbs motion sequence can be adjusted based on linked movements. A prerequisite is that kinematics of the linked movements correctly predict which adjustments are needed. We show that use of kinematic information to improve performance is even possible when a prior linked movement is performed with a different limb. For example, a skilled juggler might have learned how to correctly adjust his catching movement of the left hand when the right hand performed a throwing action in a specific way. Linkage is possibly a key mechanism of the human motor system for learning complex bimanual skills. Our study emphasizes that learning of specific movements should not be studied in isolation but within their motor sequence context.

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Identifying best practices for detecting inter-regional functional connectivity from EEG

Aggregating voxel-level statistical dependencies between multivariate time series is an important intermediate step when characterising functional connectivity (FC) between larger brain regions. However, there are numerous ways in which voxel-level data can be aggregated into inter-regional FC, and the advantages of each of these approaches are currently unclear. In this study we generate ground-truth data and compare the performances of various pipelines that estimate directed and undirected linear phase-to-phase FC between regions. We test the ability of several existing and novel FC analysis pipelines to identify the true regions within which connectivity was simulated. We test various inverse modelling algorithms, strategies to aggregate time series within regions, and connectivity metrics. Furthermore, we investigate the influence of the number of interactions, the signal-to-noise ratio, the noise mix, the interaction time delay, and the number of active sources per region on the ability of detecting phase-to-phase FC. Throughout all simulated scenarios, lowest performance is obtained with pipelines involving the absolute value of coherency. Further, the combination of dynamic imaging of coherent sources (DICS) beamforming with directed FC metrics that aggregate information across multiple frequencies leads to unsatisfactory results. Pipeline that show promising results with our simulated pseudo-EEG data involve the following steps: (1) Source projection using the linearly-constrained minimum variance (LCMV) beamformer. (2) Principal component analysis (PCA) using the same fixed number of components within every region. (3) Calculation of the multivariate interaction measure (MIM) for every region pair to assess undirected phase-to-phase FC, or calculation of time-reversed Granger Causality (TRGC) to assess directed phase-to-phase FC. We formulate recommendations based on these results that may increase the validity of future experimental connectivity studies. We further introduce the free ROIconnect plugin for the EEGLAB toolbox that includes the recommended methods and pipelines that are presented here. We show an exemplary application of the best performing pipeline to the analysis EEG data recorded during motor imagery.

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Changes in oscillations in anterior cingulate and medial prefrontal cortex are associated with altered signatures of Bayesian predictive coding in trait anxiety

Recent advances in the computational understanding of decision-making processes have led to proposals that anxiety biases how individuals form beliefs and estimate uncertainty. The anxiety and decision-making circuitry broadly overlap in regions such as the medial prefrontal cortex (mPFC), anterior cingulate cortex (ACC), and orbitofrontal cortex (OFC). Changes in activity across these brain areas could help explain how misestimation of uncertainty and altered belief updating can lead to impaired learning in anxiety. To test this prediction, this study built on recent progress in rhythm-based formulations of Bayesian predictive coding to identify sources of oscillatory modulations across the ACC, mPFC, and OFC that are associated with altered learning in subclinical trait anxiety. In a magnetoencephalography (MEG) experiment, two groups of human participants pre-screened with high and low trait anxiety (HTA, LTA: 39) performed a volatile probabilistic reward-based learning task. We modelled behaviour using a hierarchical Bayesian learning model. Furthermore, we quantified the parametric effects of trial-wise estimates of unsigned precision-weighted prediction errors (pwPEs) and, separately, precision weights and surprise on source-reconstructed MEG time-frequency responses using convolution modelling. We showed that HTA interferes with overall reward-based learning performance associated with more stochastic decisions and more pronounced lose-shift tendencies. These behavioural effects were explained by an overestimation of volatility and faster belief updating in HTA when compared to LTA. On a neural level, we observed enhanced gamma responses and decreased alpha/beta activity in HTA during the encoding of unsigned pwPEs about about stimulus outcomes relative to LTA. These effects emerged primarily in the ACC and dorsomedial PFC (dmPFC), and they were accompanied by additional ACC alpha/beta modulations representing differential encoding of precision weights in each anxiety group. Our study supports the association between subclinical trait anxiety and faster updating of beliefs in a volatile environment through gamma and alpha/beta activity changes in the ACC and dmPFC.

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Separating neural oscillations from aperiodic 1/f activity: challenges and recommendations

Electrophysiological power spectra typically consist of two components: An aperiodic part usually following an 1/f power law P{propto}1/f{beta}and periodic components appearing as spectral peaks. While the investigation of the periodic parts, commonly referred to as neural oscillations, has received considerable attention, the study of the aperiodic part has only recently gained more interest. The periodic part is usually quantified by center frequencies, powers, and bandwidths, while the aperiodic part is parameterized by the y-intercept and the 1/f exponent {beta}. For investigation of either part, however, it is essential to separate the two components. In this article, we scrutinize two frequently used methods, FOOOF (Fitting Oscillations & One-Over-F) and IRASA (Irregular Resampling Auto-Spectral Analysis), that are commonly used to separate the periodic from the aperiodic component. We evaluate these methods using diverse spectra obtained with electroencephalography (EEG), magnetoencephalography (MEG), and local field potential (LFP) recordings relating to three independent research datasets. Each method and each dataset poses distinct challenges for the extraction of both spectral parts. The specific spectral features hindering the periodic and aperiodic separation are highlighted by simulations of power spectra emphasizing these features. Through comparison with the simulation parameters defined a priori, the parameterization error of each method is quantified. Based on the real and simulated power spectra, we evaluate the advantages of both methods, discuss common challenges, note which spectral features impede the separation, assess the computational costs, and propose recommendations on how to use them.

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Harmoni: a Method for Eliminating Spurious Interactions due to the Harmonic Components in Neuronal Data

Cross-frequency synchronization (CFS) has been proposed as a mechanism for integrating spatially and spectrally distributed information in the brain. However, investigating CFS in Magneto- and Electroencephalography (MEG/EEG) is hampered by the presence of spurious neuronal interactions due to the non-sinusoidal waveshape of brain oscillations. Such waveshape gives rise to the presence of oscillatory harmonics mimicking genuine neuronal oscillations. Until recently, however, there has been no methodology for removing these harmonics from neuronal data. In order to address this long-standing challenge, we introduce a novel method (called HARMOnic miNImization - Harmoni) that removes the signal components which can be harmonics of a non-sinusoidal signal. Harmonis working principle is based on the presence of CFS between harmonic components and the fundamental component of a non-sinusoidal signal. We extensively tested Harmoni in realistic EEG simulations. The simulated couplings between the source signals represented genuine and spurious CFS and within-frequency phase synchronization. Using diverse evaluation criteria, including ROC analyses, we showed that the within- and cross-frequency spurious interactions are suppressed significantly, while the genuine activities are not affected. Additionally, we applied Harmoni to real resting-state EEG data revealing intricate remote connectivity patterns which are usually masked by the spurious connections. Given the ubiquity of non-sinusoidal neuronal oscillations in electrophysiological recordings, Harmoni is expected to facilitate novel insights into genuine neuronal interactions in various research fields, and can also serve as a steppingstone towards the development of further signal processing methods aiming at refining within- and cross-frequency synchronization in electrophysiological recordings.

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Alterations in rhythmic and non-rhythmic resting-state EEG activity and their link to cognition in older age

While many structural and biochemical changes in the brain have been previously associated with aging, the findings concerning electrophysiological signatures, reflecting functional properties of neuronal networks, remain rather controversial. To try resolve this issue, we took advantage of a large population study (N=1703) and comprehensively investigated the association of multiple EEG biomarkers (power of alpha and theta oscillations, individual alpha peak frequency (IAF), the slope of 1/f power spectral decay), aging, and aging and cognitive performance. Cognitive performance was captured with three factors representing processing speed, episodic memory, and interference resolution. Our results show that not only did IAF decline with age but it was also associated with interference resolution over multiple cortical areas. To a weaker extent, 1/f slope of the PSD showed age-related reductions, mostly in frontal brain regions. Finally, alpha power was negatively associated with the speed of processing in the right frontal lobe, despite the absence of age-related alterations. Our results thus demonstrate that multiple electrophysiological features, as well as their interplay, should be considered when investigating the association between age, neuronal activity, and cognitive performance.

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A spatio-spectral approach for movement decoding from cortical and subcortical recordings in patients with Parkinson's disease

The application of machine learning to intracranial signal analysis has the potential to revolutionize deep brain stimulation (DBS) by personalizing therapy to dynamic brain states, specific to symptoms and behaviors. Most decoding pipelines for movement decoding in the context of adaptive DBS are based on single channel frequency domain features, neglecting spatial information available in multichannel recordings. Such features are extracted either from DBS lead recordings in the subcortical target and/or from electrocorticography (ECoG). To optimize the simultaneous use of both types of signals, we developed a supervised online-compatible movement decoding pipeline based on multichannel and multiple site recordings. We found that adding spatial information to the model has the potential to improve decoding. In addition, we demonstrate movement decoding from spatio-spectral features derived from cortical and subcortical oscillations. We demonstrate between-patients variability of the spatial neural maps and its relationship to feature decoding performance. This application of spatial filters to decode movement from combined cortical and subcortical recordings is an important step in developing machine learning approaches for intelligent DBS systems.

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Relationship between Regional White Matter Hyperintensities and Alpha Oscillations in Older Adults

Aging is associated with increased white matter hyperintensities (WMHs) and with the alterations of alpha oscillations (7-13 Hz). However, a crucial question remains, whether changes in alpha oscillations relate to aging per se or whether this relationship is mediated by age-related neuropathology like WMHs. Using a large cohort of cognitively healthy older adults (N=907, 60-80 years), we assessed relative alpha power, alpha peak frequency, and long-range temporal correlations (LRTC) from resting-state EEG. We further associated these parameters with voxel-wise WMHs from 3T MRI. We found that a higher prevalence of WMHs in the superior and posterior corona radiata as well as in the thalamic radiation was related to elevated alpha power, with the strongest association in the bilateral occipital cortex. In contrast, we observed no significant relation of the WMHs probability with alpha peak frequency and LRTC. Finally, higher age was associated with elevated alpha power via total WMH volume. Although an increase in alpha oscillations due to WMH can have a compensatory nature, we rather suggest that an elevated alpha power is a consequence of WMH affecting a spatial organization of alpha sources.

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Sensorimotor functional connectivity: a neurophysiological factor related to BCI performance

Brain-Computer Interfaces (BCIs) are systems that allow users to control devices using brain activity alone. However, the ability of participants to command BCIs varies from subject to subject. For BCIs based on the modulation of sensorimotor rhythms as measured by means of electroen-cephalography (EEG), about 20% of potential users do not obtain enough accuracy to gain reliable control of the system. This lack of efficiency of BCI systems to decode users intentions requires the identification of neuro-physiological factors determining good and poor BCI performers. Given that the neuronal oscillations, used in BCI, demonstrate rich a repertoire of spatial interactions, we hypothesized that neuronal activity in sensorimotor areas would define some aspects of BCI performance. Analyses for this study were performed on a large dataset of 80 inexperienced participants. They took part in calibration and an online feedback session in the same day. Undirected functional connectivity was computed over sensorimotor areas by means of the imaginary part of coherency. The results show that post-as well as pre-stimulus connectivity in the calibration recordings is significantly correlated to online feedback performance in and feedback frequency bands. Importantly, the significance of the correlation between connectivity and BCI feedback accuracy was not due to the signal-to-noise ratio of the oscillations in the corresponding post and pre-stimulus intervals. Thus, this study shows that BCI performance is not only dependent on the amplitude of sensorimotor oscillations as shown previously, but that it also relates to sensorimotor connectivity measured during the preceding training session. The presence of such connectivity between motor and somatosensory systems is likely to facilitate motor imagery, which in turn is associated with the generation of a more pronounced modulation of sen-sorimotor oscillations (manifested in ERD/ERS) required for the adequate BCI performance. We also discuss strategies for the up-regulation of such connectivity in order to enhance BCI performance.

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Modulation of neural activity in frontopolar cortex drives reward-based motor learning

Decision-making is increasingly being recognised to play a role in learning motor skills. Understanding the neural processes regulating motor decision-making is therefore essential to identify mechanisms that contribute to motor skill learning. In decision-making tasks, the frontopolar cortex (FPC) is involved in tracking the reward of different alternative choices, as well as their reliability. Whether this FPC function extends to reward landscapes associated with a continuous movement dimension remains unknown. Here we used anodal transcranial direct current stimulation (tDCS) over the right FPC to investigate its role in reward-based motor learning. Nineteen healthy human participants completed a motor sequence learning task using trial-wise reward feedback to discover a hidden performance goal along a continuous dimension: timing. As a control condition, we modulated contralateral motor cortex (left M1) activity with tDCS, which has been shown to benefit motor skill learning but less consistently reward-based motor learning. Each active tDCS condition was contrasted to sham stimulation. Right FPC-tDCS led to faster learning primarily through a regulation of exploration, without concurrent modulation of motor noise. A Bayesian computational model revealed that following rFPC-tDCS, participants had a higher expectation of reward, consistent with their faster learning. These higher reward estimates were inferred to be less volatile, and thus participants under rFPC-tDCS deemed the mapping between movement and reward to be more stable. Relative to sham, lM1-tDCS did not significantly modulate main behavioral outcomes. The results indicate that brain regions previously linked to decision-making, such as the FPC, are relevant for motor skill learning.

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Transient amplitude modulation of alpha-band oscillations by short-time intermittent closed-loop tACS

Non-invasive brain stimulation techniques such as transcranial alternating current stimulation (tACS) have recently become extensively utilized due to their potential to modulate ongoing neuronal oscillatory activity and consequently to induce cortical plasticity relevant for various cognitive functions. However, the neurophysiological basis for stimulation effects as well as their inter-individual differences are not yet understood. In the present study we used a closed-loop EEG transcranial alternating current stimulation protocol (EEG-tACS) to examine the modulation of alpha oscillations generated in occipito-parietal areas. In particular, we investigated the effects of a repeated short-time intermittent stimulation protocol (1 s in every trial) applied over the visual cortex (Cz and Oz) and adjusted according to the phase and frequency of visual alpha oscillations on the amplitude of these oscillations. Based on previous findings, we expected higher increases in alpha amplitudes for tACS applied in-phase with ongoing oscillations as compared to an application in anti-phase and this modulation to be present in low-alpha amplitude states of the visual system (eyes opened) but not high (eyes closed). Contrary to our expectations, we found a transient suppression of alpha power in inter-individually derived spatially specific parieto-occipital components obtained via the estimation of spatial filters by using the common spatial patterns approach. The amplitude modulation was independent of the phase relationship between tACS signal and alpha oscillations, and the state of the visual system manipulated via closed- and open-eye conditions. It was also absent in conventionally analyzed single-channel and multi-channel data from an average parieto-occipital region. The fact that the tACS modulation of oscillations was phase-independent suggests that mechanisms driving the effects of tACS may not be explained by entrainment alone, but rather require neuroplastic changes or transient disruption of neural oscillations. Our study also supports the notion that the response to tACS is subject specific, where the modulatory effects are shaped by the interplay between the stimulation and different alpha generators. This favors stimulation protocols as well as analysis regimes exploiting inter-individual differences, such as spatial filters to reveal otherwise hidden stimulation effects and, thereby, comprehensively induce and study the effects and underlying mechanisms of tACS.

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Hemispheric asymmetries in resting-state EEG and fMRI are related to approach and avoidance behaviour, but not to eating behaviour or BMI

Much of our behaviour is driven by two motivational dimensions - approach and avoidance. These have been related to frontal hemispheric asymmetries in clinical and resting-state EEG studies: approach was linked to higher activity of the left relative to the right hemisphere, while avoidance was related to the opposite pattern. Increased approach behaviour, specifically towards unhealthy foods, is also observed in obesity and has been linked to asymmetry in the framework of the right-brain hypothesis of obesity. Here, we aimed to replicate previous EEG findings of hemispheric asymmetries for self-reported approach/avoidance behaviour and to relate them to eating behaviour. Further, we assessed whether resting fMRI hemispheric asymmetries can be detected and whether they are related to approach/avoidance, eating behaviour, and BMI. We analysed 3 samples: Sample 1 (n=117) containing EEG and fMRI data from lean participants, and Samples 2 (n=89) and 3 (n=152) containing fMRI data from lean, overweight, and obese participants. While in Sample 1 approach in women was related to EEG and fMRI hemispheric asymmetries, in Samples 2 and 3 this effect was not significant. Here, hemispheric asymmetries were neither related to BMI nor eating behaviour. Our study partly replicates previous EEG findings regarding hemispheric asymmetries and indicates that this relationship could also be captured using fMRI. Our findings suggest that eating behaviour and obesity are likely to be mediated by mechanisms not directly relating to frontal asymmetries in neuronal activation quantified with EEG and fMRI.

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