Functional Roles of Alpha-Band Phase Synchronization in Local and Large-Scale Cortical Networks

Alpha-frequency band (8–14 Hz) oscillations are among the most salient phenomena in human electroencephalography (EEG) recordings and yet their functional roles have remained unclear. Much of research on alpha oscillations in human EEG has focused on peri-stimulus amplitude dynamics, which phenomenologically support an idea of alpha oscillations being negatively correlated with local cortical excitability and having a role in the suppression of task-irrelevant neuronal processing. This kind of an inhibitory role for alpha oscillations is also supported by several functional magnetic resonance imaging and trans-cranial magnetic stimulation studies. Nevertheless, investigations of local and inter-areal alpha phase dynamics suggest that the alpha-frequency band rhythmicity may play a role also in active task-relevant neuronal processing. These data imply that inter-areal alpha phase synchronization could support attentional, executive, and contextual functions. In this review, we outline evidence supporting different views on the roles of alpha oscillations in cortical networks and unresolved issues that should be addressed to resolve or reconcile these apparently contrasting hypotheses

Neuronal correlates of full and partial visual conscious perception

Stimuli may induce only partial consciousness—an intermediate between null and full consciousness—where the presence but not identity of an object can be reported. The differences in the neuronal basis of full and partial consciousness are poorly understood. We investigated if evoked and oscillatory activity could dissociate full from partial conscious perception. We recorded human cortical activity with magnetoencephalography (MEG) during a visual perception task in which stimulus could be either partially or fully perceived. Partial consciousness was associated with an early increase in evoked activity and theta/low-alpha-band oscillations while full consciousness was also associated with late evoked activity and beta-band oscillations. Full from partial consciousness was dissociated by stronger evoked activity and late increase in theta oscillations that were localized to higher-order visual regions and posterior parietal and prefrontal cortices. Our results reveal both evoked activity and theta oscillations dissociate partial and full consciousness.Peer reviewe

Spectral and Anatomical Patterns of Large-Scale Synchronization Predict Human Attentional Capacity

The capacity of visual attention determines how many visual objects may be perceived at any moment. This capacity can be investigated with multiple object tracking (MOT) tasks, which have shown that it varies greatly between individuals. The neuronal mechanisms underlying capacity limits have remained poorly understood. Phase synchronization of cortical oscillations coordinates neuronal communication within the fronto-parietal attention network and between the visual regions during endogenous visual attention. We tested a hypothesis that attentional capacity is predicted by the strength of pretarget synchronization within attention-related cortical regions. We recorded cortical activity with magneto- and electroencephalography (M/EEG) while measuring attentional capacity with MOT tasks and identified large-scale synchronized networks from source-reconstructed M/EEG data. Individual attentional capacity was correlated with load-dependent strengthening of theta (3-8 Hz), alpha (8-10 Hz), and gamma-band (30-120 Hz) synchronization that connected the visual cortex with posterior parietal and prefrontal cortices. Individual memory capacity was also preceded by crossfrequency phase-phase and phase-amplitude coupling of alpha oscillation phase with beta and gamma oscillations. Our results show that good attentional capacity is preceded by efficient dynamic functional coupling and decoupling within brain regions and across frequencies, which may enable efficient communication and routing of information between sensory and attentional systems.Peer reviewe

Cross-frequency synchronization connects networks of fast and slow oscillations during visual working memory maintenance

Neuronal activity in sensory and fronto-parietal (FP) areas underlies the representation and attentional control, respectively, of sensory information maintained in visual working memory (VWM). Within these regions, beta/gamma phase-synchronization supports the integration of sensory functions, while synchronization in theta/alpha bands supports the regulation of attentional functions. A key challenge is to understand which mechanisms integrate neuronal processing across these distinct frequencies and thereby the sensory and attentional functions. We investigated whether such integration could be achieved by cross-frequency phase synchrony (CFS). Using concurrent magneto- and electroencephalography, we found that CFS was load-dependently enhanced between theta and alpha–gamma and between alpha and beta-gamma oscillations during VWM maintenance among visual, FP, and dorsal attention (DA) systems. CFS also connected the hubs of within-frequency-synchronized networks and its strength predicted individual VWM capacity. We propose that CFS integrates processing among synchronized neuronal networks from theta to gamma frequencies to link sensory and attentional functions

Roles of Brain Criticality and Multiscale Oscillations in Temporal Predictions for Sensorimotor Processing

Sensorimotor predictions are essential for adaptive behavior. In natural environments, events that demand sensorimotor predictions unfold across many timescales, and corresponding temporal predictions (either explicit or implicit) should therefore emerge in brain dynamics. Neuronal oscillations are scale-specific processes found in several frequency bands. They underlie periodicity in sensorimotor processing and can represent temporal predictions via their phase dynamics. These processes build upon endogenous neural rhythmicity and adapt in response to exogenous timing demands. While much of the research on periodicity in neural processing has focused on subsecond oscillations, these fast-scale rhythms are in fact paralleled by critical-like, scale-free dynamics and fluctuations of brain activity at various timescales, ranging from seconds to hundreds of seconds. In this review, we put forth a framework positing that critical brain dynamics are essential for the role of neuronal oscillations in timing and that cross-frequency coupling flexibly organizes neuronal processing across multiple frequencies.Peer reviewe

Hyperedge bundling : A practical solution to spurious interactions in MEG/EEG source connectivity analyses

Inter-areal functional connectivity (FC), neuronal synchronization in particular, is thought to constitute a key systems-level mechanism for coordination of neuronal processing and communication between brain regions. Evidence to support this hypothesis has been gained largely using invasive electrophysiological approaches. In humans, neuronal activity can be non-invasively recorded only with magneto-and electroencephalography (MEG/EEG), which have been used to assess FC networks with high temporal resolution and whole-scalp coverage. However, even in source-reconstructed MEG/EEG data, signal mixing, or "source leakage", is a significant confounder for FC analyses and network localization. Signal mixing leads to two distinct kinds of false-positive observations: artificial interactions (AI) caused directly by mixing and spurious interactions (SI) arising indirectly from the spread of signals from true interacting sources to nearby false loci. To date, several interaction metrics have been developed to solve the AI problem, but the SI problem has remained largely intractable in MEG/EEG all-to-all source connectivity studies. Here, we advance a novel approach for correcting SIs in FC analyses using source-reconstructed MEG/EEG data. Our approach is to bundle observed FC connections into hyperedges by their adjacency in signal mixing. Using realistic simulations, we show here that bundling yields hyperedges with good separability of true positives and little loss in the true positive rate. Hyperedge bundling thus significantly decreases graph noise by minimizing the false-positive to true-positive ratio. Finally, we demonstrate the advantage of edge bundling in the visualization of large-scale cortical networks with real MEG data. We propose that hypergraphs yielded by bundling represent well the set of true cortical interactions that are detectable and dissociable in MEG/EEG connectivity analysis.Peer reviewe

Comparison of methods to identify modules in noisy or incomplete brain networks

open6siCommunity structure, or "modularity," is a fundamentally important aspect in the organization of structural and functional brain networks, but their identification with community detection methods is confounded by noisy or missing connections. Although several methods have been used to account for missing data, the performance of these methods has not been compared quantitatively so far. In this study, we compared four different approaches to account for missing connections when identifying modules in binary and weighted networks using both Louvain and Infomap community detection algorithms. The four methods are "zeros," "row-column mean," "common neighbors," and "consensus clustering." Using Lancichinetti-Fortunato-Radicchi benchmark-simulated binary and weighted networks, we find that "zeros," "row-column mean," and "common neighbors" approaches perform well with both Louvain and Infomap, whereas "consensus clustering" performs well with Louvain but not Infomap. A similar pattern of results was observed with empirical networks from stereotactical electroencephalography data, except that "consensus clustering" outperforms other approaches on weighted networks with Louvain. Based on these results, we recommend any of the four methods when using Louvain on binary networks, whereas "consensus clustering" is superior with Louvain clustering of weighted networks. When using Infomap, "zeros" or "common neighbors" should be used for both binary and weighted networks. These findings provide a basis to accounting for noisy or missing connections when identifying modules in brain networks.openWilliams N.; Arnulfo G.; Wang S.H.; Nobili L.; Palva S.; Palva J.M.Williams, N.; Arnulfo, G.; Wang, S. H.; Nobili, L.; Palva, S.; Palva, J. M

Analysis of infant cortical synchrony is constrained by the number of recording electrodes and the recording montage

Objective: To assess how the recording montage in the neonatal EEG influences the detection of cortical source signals and their phase interactions. Methods: Scalp EEG was simulated by forward modeling 20-200 simultaneously active sources covering the cortical surface of a realistic neonatal head model. We assessed systematically how the number of scalp electrodes (11-85), analysis montage, or the size of cortical sources affect the detection of cortical phase synchrony. Statistical metrics were developed for quantifying the resolution and reliability of the montages. Results: The findings converge to show that an increase in the number of recording electrodes leads to a systematic improvement in the detection of true cortical phase synchrony. While there is always a ceiling effect with respect to discernible cortical details, we show that the average and Laplacian montages exhibit superior specificity and sensitivity as compared to other conventional montages. Conclusions: Reliability in assessing true neonatal cortical synchrony is directly related to the choice of EEG recording and analysis configurations. Because of the high conductivity of the neonatal skull, the conventional neonatal EEG recordings are spatially far too sparse for pertinent studies, and this loss of information cannot be recovered by re-montaging during analysis. Significance: Future neonatal EEG studies will need prospective planning of recording configuration to allow analysis of spatial details required by each study question. Our findings also advice about the level of details in brain synchrony that can be studied with existing datasets or by using conventional EEG recordings. (C) 2015 International Federation of Clinical Neurophysiology. Published by Elsevier Ireland Ltd. All rights reserved.Peer reviewe

Genuine cross-frequency coupling networks in human resting-state electrophysiological recordings

cited By 0Phase synchronization of neuronal oscillations in specific frequency bands coordinates anatomically distributed neuronal processing and communication. Typically, oscillations and synchronization take place concurrently in many distinct frequencies, which serve separate computational roles in cognitive functions. While within-frequency phase synchronization has been studied extensively, less is known about the mechanisms that govern neuronal processing distributed across frequencies and brain regions. Such integration of processing between frequencies could be achieved via cross-frequency coupling (CFC), either by phase-amplitude coupling (PAC) or by n:m-cross-frequency phase synchrony (CFS). So far, studies have mostly focused on local CFC in individual brain regions, whereas the presence and functional organization of CFC between brain areas have remained largely unknown. We posit that interareal CFC may be essential for large-scale coordination of neuronal activity and investigate here whether genuine CFC networks are present in human resting-state (RS) brain activity. To assess the functional organization of CFC networks, we identified brain-wide CFC networks at mesoscale resolution from stereoelectroencephalography (SEEG) and at macroscale resolution from source-reconstructed magnetoencephalography (MEG) data. We developed a novel, to our knowledge, graph-theoretical method to distinguish genuine CFC from spurious CFC that may arise from nonsinusoidal signals ubiquitous in neuronal activity. We show that genuine interareal CFC is present in human RS activity in both SEEG and MEG data. Both CFS and PAC networks coupled theta and alpha oscillations with higher frequencies in large-scale networks connecting anterior and posterior brain regions. CFS and PAC networks had distinct spectral patterns and opposing distribution of low- and high-frequency network hubs, implying that they constitute distinct CFC mechanisms. The strength of CFS networks was also predictive of cognitive performance in a separate neuropsychological assessment. In conclusion, these results provide evidence for interareal CFS and PAC being 2 distinct mechanisms for coupling oscillations across frequencies in large-scale brain networks.Peer reviewe