The role of alpha oscillations in free- and goal-directed semantic associations.

Alpha oscillations are known to play a central role in several higher-order cognitive functions, especially selective attention, working memory, semantic memory, and creative thinking. Nonetheless, we still know very little about the role of alpha in the generation of more remote semantic associations, which is key to creative and semantic cognition. Furthermore, it remains unclear how these oscillations are shaped by the intention to "be creative," which is the case in most creativity tasks. We aimed to address these gaps in two experiments. In Experiment 1, we compared alpha oscillatory activity (using a method which distinguishes genuine oscillatory activity from transient events) during the generation of free associations which were more vs. less distant from a given concept. In Experiment 2, we replicated these findings and also compared alpha oscillatory activity when people were generating free associations versus associations with the instruction to be creative (i.e. goal-directed). We found that alpha was consistently higher during the generation of more distant semantic associations, in both experiments. This effect was widespread, involving areas in both left and right hemispheres. Importantly, the instruction to be creative seems to increase alpha phase synchronisation from left to right temporal brain areas, suggesting that intention to be creative changed the flux of information in the brain, likely reflecting an increase in top-down control of semantic search processes. We conclude that goal-directed generation of remote associations relies on top-down mechanisms compared to when associations are freely generated.

Novice Users' Brain Activity and Biomechanics Change After Practicing Walking With an Ankle Exoskeleton.

Walking with an exoskeleton represents a sophisticated interplay between human physiology and mechanical augmentation, yet understanding of cortical responses in this context remains limited. To address this gap, this study aimed to explore cortical responses during walking with an ankle exoskeleton, examining how these responses evolve with familiarity to the augmentation. Healthy participants without prior exoskeleton experience underwent EEG, EMG, and motion capture analysis while walking with exoskeleton assistance at 1.2m/s. Initially, exoskeleton-assisted walking induced significant biomechanical changes accompanied by corresponding cortical alterations, leading to increased cortical involvement. In addition, after a brief familiarization period, significant increases in alpha band cortical power were observed, indicating decreased cortical engagement. These findings hold significance for elucidating the cortical mechanisms involved in exoskeleton-assisted walking and may contribute to the development of more seamlessly integrated augmentation devices.

Neural basis of lower-limb visual feedback therapy: an EEG study in healthy subjects.

BACKGROUND: Video-feedback observational therapy (VOT) is an intensive rehabilitation technique based on movement repetition and visualization that has shown benefits for motor rehabilitation of the upper and lower limbs. Despite an increase in recent literature on the neurophysiological effects of VOT in the upper limb, there is little knowledge about the cortical effects of visual feedback therapies when applied to the lower limbs. The aim of our study was to better understand the neurophysiological effects of VOT. Thus, we identified and compared the EEG biomarkers of healthy subjects undergoing lower limb VOT during three tasks: passive observation, observation and motor imagery, observation and motor execution. METHODS: We recruited 38 healthy volunteers and monitored their EEG activity while they performed a right ankle dorsiflexion task in the VOT. Three graded motor tasks associated with action observation were tested: action observation alone (O), motor imagery with action observation (OI), and motor execution synchronized with action observation (OM). The alpha and beta event-related desynchronization (ERD) and event-related synchronization (or beta rebound, ERS) rhythms were used as biomarkers of cortical activation and compared between conditions with a permutation test. Changes in connectivity during the task were computed with phase locking value (PLV). RESULTS: During the task, in the alpha band, the ERD was comparable between O and OI activities across the precentral, central and parietal electrodes. OM involved the same regions but had greater ERD over the central electrodes. In the beta band, there was a gradation of ERD intensity in O, OI and OM over central electrodes. After the task, the ERS changes were weak during the O task but were strong during the OI and OM (Cz) tasks, with no differences between OI and OM. CONCLUSION: Alpha band ERD results demonstrated the recruitment of mirror neurons during lower limb VOT due to visual feedback. Beta band ERD reflects strong recruitment of the sensorimotor cortex evoked by motor imagery and action execution. These results also emphasize the need for an active motor task, either motor imagery or motor execution task during VOT, to elicit a post-task ERS, which is absent during passive observation. Trial Registration NCT05743647.

Evaluation of Acupuncture Efficacy in Modulating Brain Activity With Periodic-Aperiodic EEG Measurements.

Acupuncture is an important therapeutic method of traditional Chinese medicine and can effectively modulate brain disorders. The therapeutic efficacy of acupuncture is hard to evaluate due to lacking of effective measurements of brain activity. In this work, we design an EEG-based monitoring system to evaluate therapeutic effect of acupuncture on human brain by extracting periodic-aperiodic features. Power spectral density is estimated to compute the adjusted power of periodic oscillatory rhythm in EEG under acupuncture stimulation. It is exhibited that the brain activity in alpha band (8-12 Hz) is significantly enhanced during acupuncture, especially in parietal and occipital lobe regions. To probe the modulatory effect of acupuncture on aperiodic brain activity, we calculate the aperiodic exponent based on the parameterization of EEG power spectra. The aperiodic exponent decreases along with acupuncture process, which is more significant in central and frontal lobe regions. Furthermore, sensitivity of different brain regions to acupuncture is assessed by the integration of adjusted power and aperiodic exponent. Experimental results demonstrate the effectiveness of proposed periodic-aperiodic measurements of EEG signals, by which different effects of four acupuncture manipulations are precisely evaluated and a knowledge graph is established. The monitoring system provides a new perspective to quantitatively evaluate acupuncture effect on human brain and improve its therapeutic efficacy in clinical applications for neural disorders.

Alpha tACS on Parieto-Occipital Cortex Mitigates Motion Sickness Based on Multiple Physiological Observation.

Approximately one third of the population is prone to motion sickness (MS), which is associated with the dysfunction in the integration of sensory inputs. Transcranial alternating current stimulation (tACS) has been widely used to modulate neurological functions by affecting neural oscillation. However, it has not been applied in the treatment of motion sickness. This study aims to investigate changes in brain oscillations during exposure to MS stimuli and to further explore the potential impact of tACS with the corresponding frequency and site on MS symptoms. A total of 19 subjects were recruited to be exposed to Coriolis stimuli to complete an inducing session. After that, they were randomly assigned to tACS stimulation group or sham stimulation group to complete a stimulation session. Electroencephalography (EEG), electrocardiogram, and galvanic skin response were recorded during the experiment. All the subjects suffering from obvious MS symptoms after inducing session were observed that alpha power of four channels of parieto-occipital lobe significantly decreased (P7: t =3.589, p <0.001; P8: t =2.667, p <0.05; O1: t =3.556, p <0.001; O2: t =2.667, p <0.05). Based on this, tACS group received the tACS stimulation at 10Hz from Oz to CPz. Compared to sham group, tACS stimulation significantly improved behavioral performance and entrained the alpha oscillation in individuals whose alpha power decrease during the inducing session. The findings show that parieto-occipital alpha oscillation plays a critical role in the integration of sensory inputs, and alpha tACS on parieto-occipital can become a potential method to mitigate MS symptoms.

Time to get deep.

Asymmetries in the size of structures deep below the cortex explain how alpha oscillations in the brain respond to shifts in attention.

Modulation of alpha oscillations by attention is predicted by hemispheric asymmetry of subcortical regions.

Evidence suggests that subcortical structures play a role in high-level cognitive functions such as the allocation of spatial attention. While there is abundant evidence in humans for posterior alpha band oscillations being modulated by spatial attention, little is known about how subcortical regions contribute to these oscillatory modulations, particularly under varying conditions of cognitive challenge. In this study, we combined MEG and structural MRI data to investigate the role of subcortical structures in controlling the allocation of attentional resources by employing a cued spatial attention paradigm with varying levels of perceptual load. We asked whether hemispheric lateralization of volumetric measures of the thalamus and basal ganglia predicted the hemispheric modulation of alpha-band power. Lateral asymmetry of the globus pallidus, caudate nucleus, and thalamus predicted attention-related modulations of posterior alpha oscillations. When the perceptual load was applied to the target and the distractor was salient caudate nucleus asymmetry predicted alpha-band modulations. Globus pallidus was predictive of alpha-band modulations when either the target had a high load, or the distractor was salient, but not both. Finally, the asymmetry of the thalamus predicted alpha band modulation when neither component of the task was perceptually demanding. In addition to delivering new insight into the subcortical circuity controlling alpha oscillations with spatial attention, our finding might also have clinical applications. We provide a framework that could be followed for detecting how structural changes in subcortical regions that are associated with neurological disorders can be reflected in the modulation of oscillatory brain activity.

Developmental trajectories of EEG aperiodic and periodic components in children 2-44 months of age.

The development of neural circuits has long-lasting effects on brain function, yet our understanding of early circuit development in humans remains limited. Here, periodic EEG power features and aperiodic components were examined from longitudinal EEGs collected from 592 healthy 2-44 month-old infants, revealing age-dependent nonlinear changes suggestive of distinct milestones in early brain maturation. Developmental changes in periodic peaks include (1) the presence and then absence of a 9-10 Hz alpha peak between 2-6 months, (2) nonlinear changes in high beta peaks (20-30 Hz) between 4-18 months, and (3) the emergence of a low beta peak (12-20 Hz) in some infants after six months of age. We hypothesized that the emergence of the low beta peak may reflect maturation of thalamocortical network development. Infant anesthesia studies observe that GABA-modulating anesthetics do not induce thalamocortical mediated frontal alpha coherence until 10-12 months of age. Using a small cohort of infants (n = 23) with EEG before and during GABA-modulating anesthesia, we provide preliminary evidence that infants with a low beta peak have higher anesthesia-induced alpha coherence compared to those without a low beta peak.

Individualized Closed-Loop Acoustic Stimulation Suggests an Alpha Phase Dependence of Sound Evoked and Induced Brain Activity Measured with EEG Recordings.

Following repetitive visual stimulation, post hoc phase analysis finds that visually evoked response magnitudes vary with the cortical alpha oscillation phase that temporally coincides with sensory stimulus. This approach has not successfully revealed an alpha phase dependence for auditory evoked or induced responses. Here, we test the feasibility of tracking alpha with scalp electroencephalogram (EEG) recordings and play sounds phase-locked to individualized alpha phases in real-time using a novel end-point corrected Hilbert transform (ecHT) algorithm implemented on a research device. Based on prior work, we hypothesize that sound-evoked and induced responses vary with the alpha phase at sound onset and the alpha phase that coincides with the early sound-evoked response potential (ERP) measured with EEG. Thus, we use each subject's individualized alpha frequency (IAF) and individual auditory ERP latency to define target trough and peak alpha phases that allow an early component of the auditory ERP to align to the estimated poststimulus peak and trough phases, respectively. With this closed-loop and individualized approach, we find opposing alpha phase-dependent effects on the auditory ERP and alpha oscillations that follow stimulus onset. Trough and peak phase-locked sounds result in distinct evoked and induced post-stimulus alpha level and frequency modulations. Though additional studies are needed to localize the sources underlying these phase-dependent effects, these results suggest a general principle for alpha phase-dependence of sensory processing that includes the auditory system. Moreover, this study demonstrates the feasibility of using individualized neurophysiological indices to deliver automated, closed-loop, phase-locked auditory stimulation.

Predicting brain edema and outcomes after thrombectomy in stroke: Frontal delta/alpha ratio as an optimal quantitative EEG index.

OBJECTIVE: We aimed to determine whether quantitative electroencephalography (QEEG) measures have predictive value for cerebral edema (CED) and clinical outcomes in acute ischemic stroke (AIS) patients with anterior circulation large vessel occlusion who underwent mechanical thrombectomy (MT). METHODS: A total of 105 patients with AIS in the anterior circulation were enrolled in this prospective study. The occurrence and severity of CED were assessed through computed tomography conducted 24 h after MT. Clinical outcomes were evaluated based on early neurological deterioration (END) and 3-month functional status, as measured by the modified Rankin scale (mRS). Electroencephalography (EEG) recordings were performed 24 h after MT, and QEEG indices were calculated from the standard 16 electrodes and 2 frontal channels (F3-C3, F4-C4). The delta/alpha ratio (DAR), the (delta + theta) / (alpha + beta) ratio (DTABR), and relative delta power were averaged over all electrodes (global) and the F3-C3 and F4-C4 channels (frontal). The predictive effect and value of QEEG indices for CED and clinical outcomes were assessed using ordinal and logistic regression models, as well as receiver operating characteristic (ROC) curves. RESULTS: Significantly, both global and frontal DAR were found to be associated with the severity of CED, END, and poor functional outcomes at 90 days, while global and frontal DTABR and relative delta power were not associated with outcomes. In ROC analysis, the best predictive effect was observed in frontal DAR, with an area under the curve of approximately 0.80. It exhibited approximately 75% sensitivity and 71% specificity for radiological and clinical outcomes when a threshold of 3.3 was used. CONCLUSIONS: QEEG techniques may be considered an efficient bedside monitoring method for assessing treatment efficacy, identifying patients at higher risk of severe CED and END, and predicting long-term functional outcomes. SIGNIFICANCE: QEEG can help identify patients at risk of severe neurological complications that can impact long-term functional recovery in AIS patients who underwent MT.

Representations of imaginary scenes and their properties in cortical alpha activity.

Imagining natural scenes enables us to engage with a myriad of simulated environments. How do our brains generate such complex mental images? Recent research suggests that cortical alpha activity carries information about individual objects during visual imagery. However, it remains unclear if more complex imagined contents such as natural scenes are similarly represented in alpha activity. Here, we answer this question by decoding the contents of imagined scenes from rhythmic cortical activity patterns. In an EEG experiment, participants imagined natural scenes based on detailed written descriptions, which conveyed four complementary scene properties: openness, naturalness, clutter level and brightness. By conducting classification analyses on EEG power patterns across neural frequencies, we were able to decode both individual imagined scenes as well as their properties from the alpha band, showing that also the contents of complex visual images are represented in alpha rhythms. A cross-classification analysis between alpha power patterns during the imagery task and during a perception task, in which participants were presented images of the described scenes, showed that scene representations in the alpha band are partly shared between imagery and late stages of perception. This suggests that alpha activity mediates the top-down re-activation of scene-related visual contents during imagery.

Analysis of the alpha activity envelope in electroencephalography in relation to the ratio of excitatory to inhibitory neural activity.

Alpha waves, one of the major components of resting and awake cortical activity in human electroencephalography (EEG), are known to show waxing and waning, but this phenomenon has rarely been analyzed. In the present study, we analyzed this phenomenon from the viewpoint of excitation and inhibition. The alpha wave envelope was subjected to secondary differentiation. This gave the positive (acceleration positive, Ap) and negative (acceleration negative, An) values of acceleration and their ratio (Ap-An ratio) at each sampling point of the envelope signals for 60 seconds. This analysis was performed on 36 participants with Alzheimer's disease (AD), 23 with frontotemporal dementia (FTD) and 29 age-matched healthy participants (NC) whose data were provided as open datasets. The mean values of the Ap-An ratio for 60 seconds at each EEG electrode were compared between the NC and AD/FTD groups. The AD (1.41 ±0.01 (SD)) and FTD (1.40 ±0.02) groups showed a larger Ap-An ratio than the NC group (1.38 ±0.02, p<0.05). A significant correlation between the envelope amplitude of alpha activity and the Ap-An ratio was observed at most electrodes in the NC group (Pearson's correlation coefficient, r = -0.92 ±0.15, mean for all electrodes), whereas the correlation was disrupted in AD (-0.09 ±0.21, p<0.05) and disrupted in the frontal region in the FTD group. The present method analyzed the envelope of alpha waves from a new perspective, that of excitation and inhibition, and it could detect properties of the EEG, Ap-An ratio, that have not been revealed by existing methods. The present study proposed a new method to analyze the alpha activity envelope in electroencephalography, which could be related to excitatory and inhibitory neural activity.

Physical activity and verbal memory performance: Mediating effects of resting-state brain activity.

Verbal short-term and long-term memory are crucial neuropsychological functions involved in core cognitive abilities. They constitute vital components of subjective well-being and academic achievement. To date, there is limited research on the association between regular physical activity and memory abilities during young adulthood. The Individual Alpha Peak Frequency (IAPF) contributes to various cognitive abilities and also appears to be sensitive to physical activity. Consequently, the IAPF has the potential to underlie the association between physical activity and verbal memory. We examined the direct relation of physical activity and verbal memory, and the potential indirect relation via IAPF in young adults. Regular physical activity was assessed via accelerometry on seven consecutive days in 115 participants (N=115, 48% female) aged 18-35 years (M=24.1, SD=3.8). In addition, verbal memory performance was assessed using an immediate and delayed free-recall task. Brain activity during rest was recorded with EEG, and IAPF was extracted for mediation analyses. Path analysis revealed pronounced sex differences in the association between physical activity, IAPF, and verbal memory performance. Exclusively in female participants, higher vigorous physical activity levels were associated with better recall performance. In contrast, no association of physical activity and memory was found in male participants. However, being more physically active was related to a higher IAPF exclusively in male participants. Physical activity shows differential associations between IAPF and verbal memory in male and female participants. However, the lack of a mediating role of IAPF suggests that this neurophysiological marker cannot explain these specific associations in young adults.

A closed-loop auditory stimulation approach selectively modulates alpha oscillations and sleep onset dynamics in humans.

Alpha oscillations play a vital role in managing the brain's resources, inhibiting neural activity as a function of their phase and amplitude, and are changed in many brain disorders. Developing minimally invasive tools to modulate alpha activity and identifying the parameters that determine its response to exogenous modulators is essential for the implementation of focussed interventions. We introduce Alpha Closed-Loop Auditory Stimulation (αCLAS) as an EEG-based method to modulate and investigate these brain rhythms in humans with specificity and selectivity, using targeted auditory stimulation. Across a series of independent experiments, we demonstrate that αCLAS alters alpha power, frequency, and connectivity in a phase, amplitude, and topography-dependent manner. Using single-pulse-αCLAS, we show that the effects of auditory stimuli on alpha oscillations can be explained within the theoretical framework of oscillator theory and a phase-reset mechanism. Finally, we demonstrate the functional relevance of our approach by showing that αCLAS can interfere with sleep onset dynamics in a phase-dependent manner.

The independence and predictivity of resting pain-free slow alpha frequency as a biomarker of pain: A reply to Mazaheri et al.

In response to Mazaheri et al.'s critique, we revisited our study (Valentini et al., 2022) on the relationship between peak alpha frequency (PAF) and pain. Their commentary prompted us to reassess our data to address the independence between slow and slowing alpha brain oscillations, as well as the predictivity of slow alpha oscillations in pain perception. Bayesian correlation analyses revealed mixed support for independence. Investigating predictivity, we found inconsistent associations between pre-PAF and unpleasantness ratings. We critically reflected on methodological and theoretical issues on the path to PAF validation as a pain biomarker. We emphasized the need for diversified methodology and analytical approaches as well as robust findings across research groups.

Blindly separated spontaneous network-level oscillations predict corticospinal excitability.

Objective.The corticospinal responses of the motor network to transcranial magnetic stimulation (TMS) are highly variable. While often regarded as noise, this variability provides a way of probing dynamic brain states related to excitability. We aimed to uncover spontaneously occurring cortical states that alter corticospinal excitability.Approach.Electroencephalography (EEG) recorded during TMS registers fast neural dynamics-unfortunately, at the cost of anatomical precision. We employed analytic Common Spatial Patterns technique to derive excitability-related cortical activity from pre-TMS EEG signals while overcoming spatial specificity issues.Main results.High corticospinal excitability was predicted by alpha-band activity, localized adjacent to the stimulated left motor cortex, and suggesting a travelling wave-like phenomenon towards frontal regions. Low excitability was predicted by alpha-band activity localized in the medial parietal-occipital and frontal cortical regions.Significance.We established a data-driven approach for uncovering network-level neural activity that modulates TMS effects. It requires no prior anatomical assumptions, while being physiologically interpretable, and can be employed in both exploratory investigation and brain state-dependent stimulation.

A randomized controlled trial of alpha phase-locked auditory stimulation to treat symptoms of sleep onset insomnia.

Sleep onset insomnia is a pervasive problem that contributes significantly to the poor health outcomes associated with insufficient sleep. Auditory stimuli phase-locked to slow-wave sleep oscillations have been shown to augment deep sleep, but it is unknown whether a similar approach can be used to accelerate sleep onset. The present randomized controlled crossover trial enrolled adults with objectively verified sleep onset latencies (SOLs) greater than 30 min to test the effect of auditory stimuli delivered at specific phases of participants' alpha oscillations prior to sleep onset. During the intervention week, participants wore an electroencephalogram (EEG)-enabled headband that delivered acoustic pulses timed to arrive anti-phase with alpha for 30 min (Stimulation). During the Sham week, the headband silently recorded EEG. The primary outcome was SOL determined by blinded scoring of EEG records. For the 21 subjects included in the analyses, stimulation had a significant effect on SOL according to a linear mixed effects model (p = 0.0019), and weekly average SOL decreased by 10.5 ± 15.9 min (29.3 ± 44.4%). These data suggest that phase-locked acoustic stimulation can be a viable alternative to pharmaceuticals to accelerate sleep onset in individuals with prolonged sleep onset latencies. Trial Registration: This trial was first registered on clinicaltrials.gov on 24/02/2023 under the name Sounds Locked to ElectroEncephalogram Phase For the Acceleration of Sleep Onset Time (SLEEPFAST), and assigned registry number NCT05743114.

Globus pallidus externus drives increase in network-wide alpha power with propofol-induced loss-of-consciousness in humans.

States of consciousness are likely mediated by multiple parallel yet interacting cortico-subcortical recurrent networks. Although the mesocircuit model has implicated the pallidocortical circuit as one such network, this circuit has not been extensively evaluated to identify network-level electrophysiological changes related to loss of consciousness (LOC). We characterize changes in the mesocircuit in awake versus propofol-induced LOC in humans by directly simultaneously recording from sensorimotor cortices (S1/M1) and globus pallidus interna and externa (GPi/GPe) in 12 patients with Parkinson disease undergoing deep brain stimulator implantation. Propofol-induced LOC is associated with increases in local power up to 20 Hz in GPi, 35 Hz in GPe, and 100 Hz in S1/M1. LOC is likewise marked by increased pallidocortical alpha synchrony across all nodes, with increased alpha/low beta Granger causal (GC) flow from GPe to all other nodes. In contrast, LOC is associated with decreased network-wide beta coupling and beta GC from M1 to the rest of the network. Results implicate an important and possibly central role of GPe in mediating LOC-related increases in alpha power, supporting a significant role of the GPe in modulating cortico-subcortical circuits for consciousness. Simultaneous LOC-related suppression of beta synchrony highlights that distinct oscillatory frequencies act independently, conveying unique network activity.

Abnormal resting-state EEG phase dynamics distinguishes major depressive disorder and bipolar disorder.

Changes in EEG have been reported in both major depressive disorder (MDD) and bipolar disorder (BD). Specifically, power changes in EEG alpha and theta frequency bands during rest and task are known in both disorders. This leaves open whether there are changes in yet another component of the electrophysiological EEG signal, namely phase-related processes that may allow for distinguishing MDD and BD. For that purpose, we investigate EEG-based spontaneous phase in the resting state of MDD, BD and healthy controls. Our main findings show: (i) decreased spontaneous phase variability in frontal theta of both MDD and BD compared to HC; (ii) decreased spontaneous phase variability in central-parietal alpha in MDD compared to both BD and HC; (iii) increased delays or lags of alpha phase cycles in MDD (but not in BD), which (iv) correlate with the decreased phase variability in MDD. Together, we show similar (decreased frontal theta variability) and distinct (decreased central-parietal alpha variability with increased lags or delays) findings in the spontaneous phase dynamics of MDD and BD. This suggests potential relevance of theta and alpha phase dynamics in distinguishing MDD and BD in clinical differential-diagnosis.