Quantifying Neural Oscillatory Synchronization: A Comparison between Spectral Coherence and Phase-Locking Value Approaches

Synchronization or phase-locking between oscillating neuronal groups is considered to be important for coordination of information among cortical networks. Spectral coherence is a commonly used approach to quantify phase locking between neural signals. We systematically explored the validity of spectral coherence measures for quantifying synchronization among neural oscillators. To that aim, we simulated coupled oscillatory signals that exhibited synchronization dynamics using an abstract phase-oscillator model as well as interacting gamma-generating spiking neural networks. We found that, within a large parameter range, the spectral coherence measure deviated substantially from the expected phase-locking. Moreover, spectral coherence did not converge to the expected value with increasing signal-to-noise ratio. We found that spectral coherence particularly failed when oscillators were in the partially (intermittent) synchronized state, which we expect to be the most likely state for neural synchronization. The failure was due to the fast frequency and amplitude changes induced by synchronization forces. We then investigated whether spectral coherence reflected the information flow among networks measured by transfer entropy (TE) of spike trains. We found that spectral coherence failed to robustly reflect changes in synchrony-mediated information flow between neural networks in many instances. As an alternative approach we explored a phase-locking value (PLV) method based on the reconstruction of the instantaneous phase. As one approach for reconstructing instantaneous phase, we used the Hilbert Transform (HT) preceded by Singular Spectrum Decomposition (SSD) of the signal. PLV estimates have broad applicability as they do not rely on stationarity, and, unlike spectral coherence, they enable more accurate estimations of oscillatory synchronization across a wide range of different synchronization regimes, and better tracking of synchronization-mediated information flow among networks.     

Hyper-Brain Networks Support Romantic Kissing in Humans

Coordinated social interaction is associated with, and presumably dependent on, oscillatory couplings within and between brains, which, in turn, consist of an interplay across different frequencies. Here, we introduce a method of network construction based on the cross-frequency coupling (CFC) and examine whether coordinated social interaction is associated with CFC within and between brains. Specifically, we compare the electroencephalograms (EEG) of 15 heterosexual couples during romantic kissing to kissing one’s own hand, and to kissing one another while performing silent arithmetic. Using graph-theory methods, we identify theta–alpha hyper-brain networks, with alpha serving a cleaving or pacemaker function. Network strengths were higher and characteristic path lengths shorter when individuals were kissing each other than when they were kissing their own hand. In both partner-oriented kissing conditions, greater strength and shorter path length for 5-Hz oscillation nodes correlated reliably with greater partner-oriented kissing satisfaction. This correlation was especially strong for inter-brain connections in both partner-oriented kissing conditions but not during kissing one’s own hand. Kissing quality assessed after the kissing with silent arithmetic correlated reliably with intra-brain strength of 10-Hz oscillation nodes during both romantic kissing and kissing with silent arithmetic. We conclude that hyper-brain networks based on CFC may capture neural mechanisms that support interpersonally coordinated voluntary action and bonding behavior.     

Oscillatory synchronization in large-scale cortical networks predicts perception

Normal brain function requires the dynamic interaction of functionally specialized but widely distributed cortical regions. Long-range synchronization of oscillatory signals has been suggested to mediate these interactions within large-scale cortical networks, but direct evidence is sparse. Here we show that oscillatory synchronization is organized in such large-scale networks. We implemented an analysis approach that allows for imaging synchronized cortical networks and applied this technique to EEG recordings in humans. We identified two networks: beta-band synchronization (~20 Hz) in a fronto-parieto-occipital network and gamma-band synchronization (~80 Hz) in a centro-temporal network. Strong perceptual correlates support their functional relevance: the strength of synchronization within these networks predicted the subjects' perception of an ambiguous audiovisual stimulus as well as the integration of auditory and visual information. Our results provide evidence that oscillatory neuronal synchronization mediates neuronal communication within frequency-specific, large-scale cortical networks.     

Dynamic processes in the human EEG

The possible mechanisms which determine the temporal dynamics of discrete narrow-band spectral components of human EEG recorded by a single electrode in the state of rest were analyzed. The dynamics of short-segment spectra was observed by application of Fast Fourier Transform (FFT) to 5-s EEG epochs successively shifted by 0.32 s. For each subject the matrices were formed and presented in a graphic mode. Matrix rows represented the number of points in each short-segment spectrum, and the columns represented the number of short-segment spectra. The columns reflect the amplitude dynamics of a given frequency, and power transition between the columns reflects the frequency dynamics. The most common type of the amplitude dynamics consisted in short (2-8 s) periods of stable activity of the discrete spectral components replaced by symmetrical bifurcation or confluence of spectral peaks. The obtained results suggest by the presence of both additive and multiplicative mechanisms of oscillatory interactions in the EEG. More detailed analysis of the amplitude-modulated EEG processes is provided by application of some additive features of the FFT to both EEG and computer-simulated signals.     

A Canonical Circuit for Generating Phase-Amplitude Coupling

‘Phase amplitude coupling’ (PAC) in oscillatory neural activity describes a phenomenon whereby the amplitude of higher frequency activity is modulated by the phase of lower frequency activity. Such coupled oscillatory activity – also referred to as ‘cross-frequency coupling’ or ‘nested rhythms’ – has been shown to occur in a number of brain regions and at behaviorally relevant time points during cognitive tasks; this suggests functional relevance, but the circuit mechanisms of PAC generation remain unclear. In this paper we present a model of a canonical circuit for generating PAC activity, showing how interconnected excitatory and inhibitory neural populations can be periodically shifted in to and out of oscillatory firing patterns by afferent drive, hence generating higher frequency oscillations phase-locked to a lower frequency, oscillating input signal. Since many brain regions contain mutually connected excitatory-inhibitory populations receiving oscillatory input, the simplicity of the mechanism generating PAC in such networks may explain the ubiquity of PAC across diverse neural systems and behaviors. Analytic treatment of this circuit as a nonlinear dynamical system demonstrates how connection strengths and inputs to the populations can be varied in order to change the extent and nature of PAC activity, importantly which phase of the lower frequency rhythm the higher frequency activity is locked to. Consequently, this model can inform attempts to associate distinct types of PAC with different network topologies and physiologies in real data.     

Multimodal functional network connectivity: an EEG-fMRI fusion in network space

EEG and fMRI recordings measure the functional activity of multiple coherent networks distributed in the cerebral cortex. Identifying network interaction from the complementary neuroelectric and hemodynamic signals may help to explain the complex relationships between different brain regions. In this paper, multimodal functional network connectivity (mFNC) is proposed for the fusion of EEG and fMRI in network space. First, functional networks (FNs) are extracted using spatial independent component analysis (ICA) in each modality separately. Then the interactions among FNs in each modality are explored by Granger causality analysis (GCA). Finally, fMRI FNs are matched to EEG FNs in the spatial domain using network-based source imaging (NESOI). Investigations of both synthetic and real data demonstrate that mFNC has the potential to reveal the underlying neural networks of each modality separately and in their combination. With mFNC, comprehensive relationships among FNs might be unveiled for the deep exploration of neural activities and metabolic responses in a specific task or neurological state.     

Role of joint language control during cross-language communication: evidence from cross-frequency coupling

How do bilingual interlocutors inhibit interference from the non-target language to achieve brain-to-brain information exchange in a task to simulate a bilingual speaker–listener interaction. In the current study, two electroencephalogram devices were employed to record pairs of participants’ performances in a joint language switching task. Twenty-eight (14 pairs) unbalanced Chinese–English bilinguals (L1 Chinese) were instructed to name pictures in the appropriate language according to the cue. The phase-amplitude coupling analysis was employed to reveal the large-scale brain network responsible for joint language control between interlocutors. We found that (1) speakers and listeners coordinately suppressed cross-language interference through cross-frequency coupling, as shown in the increased delta/theta phase-amplitude and delta/alpha phase-amplitude coupling when switching to L2 than switching to L1; (2) speakers and listeners were both able to simultaneously inhibit cross-person item-level interference which was demonstrated by stronger cross-frequency coupling in the cross person condition compared to the within person condition. These results indicate that current bilingual models (e.g., the inhibitory control model) should incorporate mechanisms that address inhibiting interference sourced in both language and person (i.e., cross-language and cross-person item-level interference) synchronously through joint language control in dynamic cross-language communication.     

Relevance of nonlinear lumped-parameter models in the analysis of depth-EEG epileptic signals

In the field of epilepsy, the analysis of stereoelectroencephalographic (SEEG, intra-cerebral recording) signals with signal processing methods can help to better identify the epileptogenic zone, the area of the brain responsible for triggering seizures, and to better understand its organization. In order to evaluate these methods and to physiologically interpret the results they provide, we developed a model able to produce EEG signals from “organized” networks of neural populations. Starting from a neurophysiologically relevant model initially proposed by Lopes Da Silva et al. [Lopes da Silva FH, Hoek A, Smith H, Zetterberg LH (1974) Kybernetic 15: 27–37] and recently re-designed by Jansen et al. [Jansen BH, Zouridakis G, Brandt ME (1993) Biol Cybern 68: 275–283] the present study demonstrates that this model can be extended to generate spontaneous EEG signals from multiple coupled neural populations. Model parameters related to excitation, inhibition and coupling are then altered to produce epileptiform EEG signals. Results show that the qualitative behavior of the model is realistic; simulated signals resemble those recorded from different brain structures for both interictal and ictal activities. Possible exploitation of simulations in signal processing is illustrated through one example; statistical couplings between both simulated signals and real SEEG signals are estimated using nonlinear regression. Results are compared and show that, through the model, real SEEG signals can be interpreted with the aid of signal processing methods. Received: 3 January 2000 / Accepted: 24 March 2000     

Coupled oscillators for modeling and analysis of EEG/MEG oscillations.

This study presents three EEG/MEG applications in which the modeling of oscillatory signal components offers complementary analysis and an improved explanation of the underlying generator structures. Coupled oscillator networks were used for modeling. Parameters of the corresponding ordinary coupled differential equation (ODE) system are identified using EEG/MEG data and the resulting solution yields the modeled signals. This model-related analysis strategy provides information about the coupling quantity and quality between signal components (example 1, neonatal EEG during quiet sleep), allows identification of the possible contribution of hidden generator structures (example 2, 600-Hz MEG oscillations in somatosensory evoked magnetic fields), and can explain complex signal characteristics such as amplitude-frequency coupling and frequency entrainment (example 3, EEG burst patterns in sedated patients).     

A novel approach to the detection of synchronisation in EEG based on empirical mode decomposition

Transient neural assemblies mediated by synchrony in particular frequency ranges are thought to underlie cognition. We propose a new approach to their detection, using empirical mode decomposition (EMD), a data-driven approach removing the need for arbitrary bandpass filter cut-offs. Phase locking is sought between modes. We explore the features of EMD, including making a quantitative assessment of its ability to preserve phase content of signals, and proceed to develop a statistical framework with which to assess synchrony episodes. Furthermore, we propose a new approach to ensure signal decomposition using EMD. We adapt the Hilbert spectrum to a time-frequency representation of phase locking and are able to locate synchrony successfully in time and frequency between synthetic signals reminiscent of EEG. We compare our approach, which we call EMD phase locking analysis (EMDPL) with existing methods and show it to offer improved time-frequency localisation of synchrony. Action Editor: Carson C. Chow     

Concerning the amplitude modulation of the human EEG

Amplitude-modulated processes can be formally presented as a product of two or more sinusoids. This makes it possible to study them by means of analysis of multiplicative phenomena using the Fast Fourier Transform (FFT). To assess the contribution of amplitude EEG modulation to the dynamic of electrical activity of the human brain, the results of the FFT of simulated signals obtained by multiplication of oscillatory processes with different parameters were compared with the results of the FFT of a single EEG recording from a subject at rest. We studied the temporal dynamics of spectral components calculated with different spectral resolution under similar conditions for real and simulated signals. An attempt was made to analyze and interpret the amplitude-modulated EEG processes using the additive properties of the FTT. It was shown that processes of amplitude modulation are present in electrical brain activity and determine the synchronism of changes in time in the majority of frequency components of the EEG spectrum. The presence of the amplitude modulation in bioelectrical processes is of a fundamental nature, since it is a direct reflection of the control, synchronization, regulation, and intersystem interaction in the nervous and other body systems. The study of this modulation gives a clue to the mechanisms of these processes.     

Assessing Movement Factors in Upper Limb Kinematics Decoding from EEG Signals

The past decades have seen the rapid development of upper limb kinematics decoding techniques by performing intracortical recordings of brain signals. However, the use of non-invasive approaches to perform similar decoding procedures is still in its early stages. Recent studies show that there is a correlation between electroencephalographic (EEG) signals and hand-reaching kinematic parameters. From these studies, it could be concluded that the accuracy of upper limb kinematics decoding depends, at least partially, on the characteristics of the performed movement. In this paper, we have studied upper limb movements with different speeds and trajectories in a controlled environment to analyze the influence of movement variability in the decoding performance. To that end, low frequency components of the EEG signals have been decoded with linear models to obtain the position of the volunteer’s hand during performed trajectories grasping the end effector of a planar manipulandum. The results confirm that it is possible to obtain kinematic information from low frequency EEG signals and show that decoding performance is significantly influenced by movement variability and tracking accuracy as continuous and slower movements improve the accuracy of the decoder. This is a key factor that should be taken into account in future experimental designs.     

EEG Source Reconstruction Reveals Frontal-Parietal Dynamics of Spatial Conflict Processing

Cognitive control requires the suppression of distracting information in order to focus on task-relevant information. We applied EEG source reconstruction via time-frequency linear constrained minimum variance beamforming to help elucidate the neural mechanisms involved in spatial conflict processing. Human subjects performed a Simon task, in which conflict was induced by incongruence between spatial location and response hand. We found an early (∼200 ms post-stimulus) conflict modulation in stimulus-contralateral parietal gamma (30–50 Hz), followed by a later alpha-band (8–12 Hz) conflict modulation, suggesting an early detection of spatial conflict and inhibition of spatial location processing. Inter-regional connectivity analyses assessed via cross-frequency coupling of theta (4–8 Hz), alpha, and gamma power revealed conflict-induced shifts in cortical network interactions: Congruent trials (relative to incongruent trials) had stronger coupling between frontal theta and stimulus-contrahemifield parietal alpha/gamma power, whereas incongruent trials had increased theta coupling between medial frontal and lateral frontal regions. These findings shed new light into the large-scale network dynamics of spatial conflict processing, and how those networks are shaped by oscillatory interactions.     

A New Method for Nonlinear Dynamic Analysis of the Multi-kinetics Neural Mass Model

ObjectiveThe aim was to use a new method to analyze the nonlinear dynamic characteristics of the multi-kinetics neural mass model. We hope that this new method can be as an auxiliary judgment tool for the diagnosis of brain diseases and the identification of brain activity states.MethodsWe apply the Lorenz plot to analyze the nonlinear dynamic characteristics of electroencephalogram (EEG) signals from the multi-kinetics neural mass models. The standard deviations in two orthogonal directions of the Lorenz plot are further used to quantify the nonlinear dynamic characteristics of EEG signals.ResultsThe results show that the normalized signal frequency power spectrum may not be able to distinguish normal EEG signals and epileptiform spikes, but the Lorenz plot can distinguish the normal EEG signals and epileptiform spikes effectively. For EEG signals with multi-rhythms, the Lorenz plot of all the simulated signals are oval, but the value of SD1/SD2 increases monotonically when the multi-rhythm EEG signals change from low frequency to high frequency.ConclusionThe Lorenz plot of EEG signals with different rhythms presents different distribution. It is an effective nonlinear analysis method for EEG signals.     

Multifractality of decomposed EEG during imaginary and real visual-motor tracking

We test the possible multifractal properties of dominant EEG frequency components, when a subject tracks a path on a map, either only by eyes (imaginary movement – IM) or by visual-motor tracking of discretely moving spot in regular (RM) and Brownian time-step (BM) (real tracking of moving spot). We check the hypotheses that the fractal properties of filtered EEG (1) change with respect to the law of spot movement; (2) differ among filtered EEG components and scalp sites; (3) differ among real and imaginary tracking. Sixteen right-handed subjects begin to perform IM, next – real spot tracking (RM and BM) following a moving spot on streets of a citymap displayed on a computer screen, by push forward/backward a joystick. Multichannel long-lasting EEG is band-pass filtered for theta, alpha, beta and gamma oscillations. The Wavelet-Transform-Modulus-Maxima-Method is applied to reveal multifractality [local fractal dimensions D _max(h)] among task conditions, frequency bands and sites. Non-parametric statistical estimation of the fractal measures h _{D max} is finally applied. Multifractality is established for all experimental conditions, EEG components and sites as follows among filtered components – anticorrelation (h _Dmax < 0.5) in beta and gamma, and long-range correlation (h _Dmax > 0.5) for theta and alpha oscillations; among tasks – for RM and BM, h _Dmax differ significantly whereas IM resembles mostly RM; among sites – no significant difference for local fractal properties is established. The results suggest that for both imaginary and real visual-motor tracking a line, multifractal scaling, specific for lower and higher EEG oscillations, is a very stable intrinsic one for the activity of large brain areas. The external events (task conditions) insert weak effect on the scaling.     

Two generalized algorithms measuring phase–amplitude cross-frequency coupling in neuronal oscillations network

An increasing number of studies pays attention to cross-frequency coupling in neuronal oscillations network, as it is considered to play an important role in exchanging and integrating of information. In this study, two generalized algorithms, phase–amplitude coupling-evolution map approach and phase–amplitude coupling-conditional mutual information which have been developed and applied originally in an identical rhythm, are generalized to measure cross-frequency coupling. The effectiveness of quantitatively distinguishing the changes of coupling strength from the measurement of phase–amplitude coupling (PAC) is demonstrated based on simulation data. The data suggest that the generalized algorithms are able to effectively evaluate the strength of PAC, which are consistent with those traditional approaches, such as PAC-PLV and PAC-MI. Experimental data, which are local field potentials obtained from anaesthetized SD rats, have also been analyzed by these two generalized approaches. The data show that the theta–low gamma PAC in the hippocampal CA3–CA1 network is significantly decreased in the glioma group compared to that in the control group. The results, obtained from either simulation data or real experimental signals, are consistent with that of those traditional approaches PAC-MI and PAC-PLV. It may be considered as a proper indicator for the cross frequency coupling in sub-network, such as the hippocampal CA3 and CA1.     

Individual resistance to factors of monotony reflected in the EEG

Three groups of operators substantially different in their performance quality were examined. Under conditions of monotonous activity, subjects with the highest initial level of activation, minimum EEG total spectral power, and minimum level of EEG coherence in the frontal cortical areas worked most steadily. Under the same conditions, subjects with a rather high spectral power of the theta and beta2 EEG frequency components, highest coherence in the frontal areas, and low coherence in the caudal areas of the cortex worked least steadily. EEG phenomena testify to a rather low level of activation of the frontal cortical areas associated with a facilitation of cortico-subcortical neuronal interactions and an attenuation of the operating neural streams. This results in a decrease in the level of any attention, its involuntary switching, and short-term loss of the control over the current performance.     

Blind source separation,wavelet denoising and discriminant analysis for EEG artefacts and noise cancelling

This paper proposes an automatic method for artefact removal and noise elimination from scalp electroencephalogram recordings (EEG). The method is based on blind source separation (BSS) and supervised classification and proposes a combination of classical and news features and classes to improve artefact elimination (ocular, high frequency muscle and ECG artefacts). The role of a supplementary step of wavelet denoising (WD) is explored and the interactions between BSS, denoising and classification are analyzed. The results are validated on simulated signals by quantitative evaluation criteria and on real EEG by medical expertise. The proposed methodology successfully rejected a good percentage of artefacts and noise, while preserving almost all the cerebral activity. The “denoised artefact-free” EEG presents a very good improvement compared with recorded raw EEG: 96% of the EEGs are easier to interpret.     

Improving the separability of multiple EEG features for a BCI by neural-time-series-prediction-preprocessing

A recently proposed method for EEG preprocessing is extended and analyzed in this work via a range of different tests in combination with various other BCI components. Neural-time-series-prediction-processing (NTSPP) is a predictive approach to EEG preprocessing where prediction models (PMs) are trained to perform one-step-ahead prediction of the EEG times-series which reflect motor imagery induced alterations in neuronal activity. Due to the specialization of distinct PMs, the predicted signals (Ys) and error signals (Es) are distinctly different from the original (Os) signals. The PMs map the Os signals to a higher dimension which, in the majority of cases, produces features that are more separable than those produced by the Os signals. Four feature extraction procedures, ranging in complexity and in terms of the information which is extracted i.e., time domain, frequency domain and time–frequency (t–f) domain, are used to determine the separability enhancements which are verified by comparative statistical tests and brain–computer interface (BCI) tests on six subjects. It is shown that, in the majority of the tests, features extracted from the NTSPP signals are more separable than those extracted from the Os signals, in terms of increased Euclidean distance between class means, reduced inter-class correlations and intra-class variance, and higher classification accuracy (CA), information transfer (IT) rate and mutual information (MI).