The cognitive triad network - oscillation - behaviour links individual differences in EEG theta frequency with task performance and effective connectivity.

We reconcile two significant lines of Cognitive Neuroscience research: the relationship between the structural and functional architecture of the brain and behaviour on the one hand and the functional significance of oscillatory brain processes to behavioural performance on the other. Network neuroscience proposes that the three elements, behavioural performance, EEG oscillation frequency, and network connectivity should be tightly connected at the individual level. Young and old healthy adults were recruited as a proxy for performance variation. An auditory inhibitory control task was used to demonstrate that task performance correlates with the individual EEG frontal theta frequency. Older adults had a significantly slower theta frequency, and both theta frequency and task performance correlated with the strengths of two network connections that involve the main areas of inhibitory control and speech processing. The results suggest that both the recruited functional network and the oscillation frequency induced by the task are specific to the task, are inseparable, and mark individual differences that directly link structure and function to behaviour in health and disease.

Elevating understanding: Linking high-altitude hypoxia to brain aging through EEG functional connectivity and spectral analyses.

High-altitude hypoxia triggers brain function changes reminiscent of those in healthy aging and Alzheimer's disease, compromising cognition and executive functions. Our study sought to validate high-altitude hypoxia as a model for assessing brain activity disruptions akin to aging. We collected EEG data from 16 healthy volunteers during acute high-altitude hypoxia (at 4,000 masl) and at sea level, focusing on relative changes in power and aperiodic slope of the EEG spectrum due to hypoxia. Additionally, we examined functional connectivity using wPLI, and functional segregation and integration using graph theory tools. High altitude led to slower brain oscillations, that is, increased δ and reduced α power, and flattened the 1/f aperiodic slope, indicating higher electrophysiological noise, akin to healthy aging. Notably, functional integration strengthened in the θ band, exhibiting unique topographical patterns at the subnetwork level, including increased frontocentral and reduced occipitoparietal integration. Moreover, we discovered significant correlations between subjects' age, 1/f slope, θ band integration, and observed robust effects of hypoxia after adjusting for age. Our findings shed light on how reduced oxygen levels at high altitudes influence brain activity patterns resembling those in neurodegenerative disorders and aging, making high-altitude hypoxia a promising model for comprehending the brain in health and disease.

Exposure to high-altitude hypoxia, with reduced oxygen levels, can replicate brain function changes akin to aging and Alzheimer's disease. In our work, we propose high-altitude hypoxia as a possible reversible model of human brain aging. We gathered EEG data at high altitude and sea level, investigating the impact of hypoxia on brainwave patterns and connectivity. Our findings revealed that high-altitude exposure led to slower and noisier brain oscillations and produced altered brain connectivity, resembling some remarkable changes seen in the aging process. Intriguingly, these changes were linked to age, even when hypoxia's effects were considered. Our research unveils how high-altitude conditions emulate brain patterns associated with aging and neurodegenerative conditions, providing valuable insights into the understanding of both normal and impaired brain function.

A High-Resolution LED Stimulator for Steady-State Visual Stimulation: Customizable, Affordable, and Open Source.

Visually evoked steady-state potentials (SSVEPs) are neural responses elicited by visual stimuli oscillating at specific frequencies. In this study, we introduce a novel LED stimulator system explicitly designed for steady-state visual stimulation, offering precise control over visual stimulus parameters, including frequency resolution, luminance, and the ability to control the phase at the end of the stimulation. The LED stimulator provides a personalized, modular, and affordable option for experimental setups. Based on the Teensy 3.2 board, the stimulator utilizes direct digital synthesis and pulse width modulation techniques to control the LEDs. We validated its performance through four experiments: the first two measured LED light intensities directly, while the last two assessed the stimulator's impact on EEG recordings. The results demonstrate that the stimulator can deliver a stimulus suitable for generating SSVEPs with the desired frequency and phase resolution. As an open source resource, we provide comprehensive documentation, including all necessary codes and electrical diagrams, which facilitates the system's replication and adaptation for specific experimental requirements, enhancing its potential for widespread use in the field of neuroscience setups.

Self-regulation learning as active inference: dynamic causal modeling of an fMRI neurofeedback task.

Introduction: Learning to self-regulate brain activity by neurofeedback has been shown to lead to changes in the brain and behavior, with beneficial clinical and non-clinical outcomes. Neurofeedback uses a brain-computer interface to guide participants to change some feature of their brain activity. However, the neural mechanism of self-regulation learning remains unclear, with only 50% of the participants succeeding in achieving it. To bridge this knowledge gap, our study delves into the neural mechanisms of self-regulation learning via neurofeedback and investigates the brain processes associated with successful brain self-regulation. Methods: We study the neural underpinnings of self-regulation learning by employing dynamical causal modeling (DCM) in conjunction with real-time functional MRI data. The study involved a cohort of 18 participants undergoing neurofeedback training targeting the supplementary motor area. A critical focus was the comparison between top-down hierarchical connectivity models proposed by Active Inference and alternative bottom-up connectivity models like reinforcement learning. Results: Our analysis revealed a crucial distinction in brain connectivity patterns between successful and non-successful learners. Particularly, successful learners evinced a significantly stronger top-down effective connectivity towards the target area implicated in self-regulation. This heightened top-down network engagement closely resembles the patterns observed in goal-oriented and cognitive control studies, shedding light on the intricate cognitive processes intertwined with self-regulation learning. Discussion: The findings from our investigation underscore the significance of cognitive mechanisms in the process of self-regulation learning through neurofeedback. The observed stronger top-down effective connectivity in successful learners indicates the involvement of hierarchical cognitive control, which aligns with the tenets of Active Inference. This study contributes to a deeper understanding of the neural dynamics behind successful self-regulation learning and provides insights into the potential cognitive architecture underpinning this process.

Modelling neural entrainment and its persistence: influence of frequency of stimulation and phase at the stimulus offset.

Neural entrainment, the synchronization of brain oscillations to the frequency of an external stimuli, is a key mechanism that shapes perceptual and cognitive processes.Objective.Using simulations, we investigated the dynamics of neural entrainment, particularly the period following the end of the stimulation, since the persistence (reverberation) of neural entrainment may condition future sensory representations based on predictions about stimulus rhythmicity.Methods.Neural entrainment was assessed using a modified Jansen-Rit neural mass model (NMM) of coupled cortical columns, in which the spectral features of the output resembled that of the electroencephalogram (EEG). We evaluated spectro-temporal features of entrainment as a function of the stimulation frequency, the resonant frequency of the neural populations comprising the NMM, and the coupling strength between cortical columns. Furthermore, we tested if the entrainment persistence depended on the phase of the EEG-like oscillation at the time the stimulus ended.Main Results.The entrainment of the column that received the stimulation was maximum when the frequency of the entrainer was within a narrow range around the resonant frequency of the column. When this occurred, entrainment persisted for several cycles after the stimulus terminated, and the propagation of the entrainment to other columns was facilitated. Propagation also depended on the resonant frequency of the second column, and the coupling strength between columns. The duration of the persistence of the entrainment depended on the phase of the neural oscillation at the time the entrainer terminated, such that falling phases (fromπ/2 to 3π/2 in a sine function) led to longer persistence than rising phases (from 0 toπ/2 and 3π/2 to 2π).Significance.The study bridges between models of neural oscillations and empirical electrophysiology, providing insights to the mechanisms underlying neural entrainment and the use of rhythmic sensory stimulation for neuroenhancement.

Cortical ignition dynamics is tightly linked to the core organisation of the human connectome.

The capability of cortical regions to flexibly sustain an "ignited" state of activity has been discussed in relation to conscious perception or hierarchical information processing. Here, we investigate how the intrinsic propensity of different regions to get ignited is determined by the specific topological organisation of the structural connectome. More specifically, we simulated the resting-state dynamics of mean-field whole-brain models and assessed how dynamic multistability and ignition differ between a reference model embedding a realistic human connectome, and alternative models based on a variety of randomised connectome ensembles. We found that the strength of global excitation needed to first trigger ignition in a subset of regions is substantially smaller for the model embedding the empirical human connectome. Furthermore, when increasing the strength of excitation, the propagation of ignition outside of this initial core-which is able to self-sustain its high activity-is way more gradual than for any of the randomised connectomes, allowing for graded control of the number of ignited regions. We explain both these assets in terms of the exceptional weighted core-shell organisation of the empirical connectome, speculating that this topology of human structural connectivity may be attuned to support enhanced ignition dynamics.

Persistence of EEG Alpha Entrainment Depends on Stimulus Phase at Offset.

Neural entrainment is the synchronization of neural activity to the frequency of repetitive external stimuli, which can be observed as an increase in the electroencephalogram (EEG) power spectrum at the driving frequency, -also known as the steady-state response. Although it has been systematically reported that the entrained EEG oscillation persists for approximately three cycles after stimulus offset, the neural mechanisms underpinning it remain unknown. Focusing on alpha oscillations, we adopt the dynamical excitation/inhibition framework, which suggests that phases of entrained EEG signals correspond to alternating excitatory/inhibitory states of the neural circuitry. We hypothesize that the duration of the persistence of entrainment is determined by the specific functional state of the entrained neural network at the time the stimulus ends. Steady-state visually evoked potentials (SSVEP) were elicited in 19 healthy volunteers at the participants' individual alpha peaks. Visual stimulation consisted of a sinusoidally-varying light terminating at one of four phases: 0, π/2, π, and 3π/2. The persistence duration of the oscillatory activity was analyzed as a function of the terminating phase of the stimulus. Phases of the SSVEP at the stimulus termination were distributed within a constant range of values relative to the phase of the stimulus. Longer persistence durations were obtained when visual stimulation terminated towards the troughs of the alpha oscillations, while shorter persistence durations occurred when stimuli terminated near the peaks. Source localization analysis suggests that the persistence of entrainment reflects the functioning of fronto-occipital neuronal circuits, which might prime the sensory representation of incoming visual stimuli based on predictions about stimulus rhythmicity. Consequently, different states of the network at the end of the stimulation, corresponding to different states of intrinsic neuronal coupling, may determine the time windows over which coding of incoming sensory stimulation is modulated by the preceding oscillatory activity.

Improving EEG Muscle Artifact Removal With an EMG Array.

Removal of artifacts induced by muscle activity is crucial for analysis of the electroencephalogram (EEG), and continues to be a challenge in experiments where the subject may speak, change facial expressions, or move. Ensemble empirical mode decomposition with canonical correlation analysis (EEMD-CCA) has been proven to be an efficient method for denoising of EEG contaminated with muscle artifacts. EEMD-CCA, likewise the majority of algorithms, does not incorporate any statistical information of the artifact, namely, electromyogram (EMG) recorded over the muscles actively contaminating the EEG. In this paper, we propose to extend EEMD-CCA in order to include an EMG array as information to aid the removal of artifacts, assessing the performance gain achieved when the number of EMG channels grow. By filtering adaptively (recursive least squares, EMG array as reference) each component resulting from CCA, we aim to ameliorate the distortion of brain signals induced by artifacts and denoising methods. We simulated several noise scenarios based on a linear contamination model, between real and synthetic EEG and EMG signals, and varied the number of EMG channels available to the filter. Our results exhibit a substantial improvement in the performance as the number of EMG electrodes increase from 2 to 16. Further increasing the number of EMG channels up to 128 did not have a significant impact on the performance. We conclude by recommending the use of EMG electrodes to filter components, as it is a computationally inexpensive enhancement that impacts significantly on performance using only a few electrodes.

Selective modulation of nociceptive processing due to noise distraction.

This study investigates the effects of noise distraction on the different components and sources of laser-evoked potentials (LEPs) whilst attending to either the spatial component (localisation performance task) or the affective component (unpleasantness rating task) of pain. LEPs elicited by CO2 laser stimulation of the right forearm were recorded from 64 electrodes in 18 consenting healthy volunteers. Subjects reported either pain location or unpleasantness, in the presence and absence of distraction by continuous 85 dBa white noise. Distributed sources of the LEP peaks were identified using Low Resolution Electromagnetic Tomography (LORETA). Pain unpleasantness ratings and P2 (430 ms) peak amplitude were significantly reduced by distraction during the unpleasantness task, whereas the localisation ability and the corresponding N1/N2 (310 ms) peak amplitude remained unchanged. Noise distraction (at 310 ms) reduced activation in the anterior cingulate cortex (ACC) and precuneus during attention to localisation and unpleasantness, respectively. This suggests a complimentary role for these two areas in the control of attention to pain. In contrast, activation of the occipital pole and SII were enhanced by noise during the localisation and unpleasantness task, respectively, suggesting that the presence of noise was associated with increased spatial attentional load. This study has shown selective modulation of affective pain processing by noise distraction, indicated by a reduction in the unpleasantness ratings and P2 peak amplitude and associated activity within the medial pain system. These results show that processing of the affective component of pain can be differentially modulated by top-down processes, providing a potential mechanism for therapeutic intervention.

Exploring event-related brain dynamics with tests on complex valued time-frequency representations.

A question subject to intense debate is whether scalp-recorded event-related brain potentials are due to phase resetting of the ongoing electroencephalogram (EEG) or rather to the superimposition of time-locked components on background activity. The two hypotheses are usually assessed by means of statistics in the time-frequency domain, for example, through wavelet transformation of multiple EEG trials that yield for each time and frequency a scatter plot of complex values coefficients. Currently, intertrial phase correlation (phase locking or phase coherence) is taken as evidence for phase resetting at a given frequency and latency. Here we present a formal analysis using a complex t-statistic to illustrate that such measures are, in effect, tests for the mean vector of the repeated trials, and as such on their own are inappropriate measures of phase resetting. We also propose simple t-like statistics for testing changes in (i) the mean (presence of an event-related potential), (ii) the amplitude variance (presence of (de)synchronization) and (iii) the concentration of phases (phase locking). The first two statistics are found to be proper measures of the presence of a non-zero mean activity and induced activity, respectively. In the third case, two different tests are introduced: one based on measuring the alignment of coefficients in the complex plane and another derived from the argument that phase locking persists when the mean of the coefficients is removed. Both statistics gave unambiguous evidence of the presence of phase locking suggesting that they constitute promising tools in the analysis of event-related brain dynamics.