Left parietal tACS at alpha frequency induces a shift of visuospatial attention

Background Voluntary shifts of visuospatial attention are associated with a lateralization of parieto-occipital alpha power (7-13Hz), i.e. higher power in the hemisphere ipsilateral and lower power contralateral to the locus of attention. Recent noninvasive neuromodulation studies demonstrated that alpha power can be experimentally increased using transcranial alternating current stimulation (tACS). Objective/Hypothesis We hypothesized that tACS at alpha frequency over the left parietal cortex induces shifts of attention to the left hemifield. However, spatial attention shifts not only occur voluntarily (endogenous/ top-down), but also stimulus-driven (exogenous/ bottom-up). To study the task-specificity of the potential effects of tACS on attentional processes, we administered three conceptually different spatial attention tasks. Methods 36 healthy volunteers were recruited from an academic environment. In two separate sessions, we applied either high-density tACS at 10Hz, or sham tACS, for 35–40 minutes to their left parietal cortex. We systematically compared performance on endogenous attention, exogenous attention, and stimulus detection tasks. Results In the endogenous attention task, a greater leftward bias in reaction times was induced during left parietal 10Hz tACS as compared to sham. There were no stimulation effects in either the exogenous attention or the stimulus detection task. Conclusion The study demonstrates that high-density tACS at 10Hz can be used to modulate visuospatial attention performance. The tACS effect is task-specific, indicating that not all forms of attention are equally susceptible to the stimulation

Hearing through your eyes: neural basis of audiovisual cross-activation, revealed by transcranial alternating current stimulation

Some people experience auditory sensations when seeing visual flashes or movements. This prevalent synaesthesia-like ‘visual-evoked auditory response’ (vEAR) could result either from over-exuberant cross-activation between brain areas, and/or reduced inhibition of normally-occurring cross-activation. We have used transcranial alternating current stimulation (tACS) to test these theories. We applied tACS at 10Hz (alpha-band frequency) or 40Hz (gamma-band), bilaterally either to temporal or occipital sites, while measuring same/different discrimination of paired auditory (A) versus visual (V) 'Morse code' sequences. At debriefing, participants were classified as vEAR or non-vEAR depending on whether they reported 'hearing' the silent flashes. In non-vEAR participants, temporal 10Hz tACS caused impairment of A performance, which correlated with improved V; conversely under occipital tACS, poorer V performance correlated with improved A. This reciprocal pattern suggests that sensory cortices are normally mutually inhibitory, and that alpha-frequency tACS may bias the balance of competition between them. vEAR participants showed no tACS effects, consistent with reduced inhibition, or enhanced cooperation between modalities. In addition, temporal 40Hz tACS impaired V performance, specifically in individuals who showed a performance advantage for V (relative to A). Gamma-frequency tACS may therefore modulate the ability of these individuals to benefit from recoding flashes into the auditory modality, possibly by disrupting cross-activation of auditory areas by visual stimulation. Our results support both theories, suggesting that vEAR may depend on disinhibition of normally-occurring sensory cross-activation, which may be expressed more strongly in some individuals. Furthermore, endogenous alpha and gamma-frequency oscillations may function respectively to inhibit or promote this cross-activation

After-effects of 10 Hz tACS over the prefrontal cortex on phonological word decisions

Introduction Previous work in the language domain has shown that 10 Hz rTMS of the left or right posterior inferior frontal gyrus (pIFG) in the prefrontal cortex impaired phonological decision-making, arguing for a causal contribution of the bilateral pIFG to phonological processing. However, the neurophysiological correlates of these effects are unclear. The present study addressed the question whether neural activity in the prefrontal cortex could be modulated by 10 Hz tACS and how this would affect phonological decisions. Methods In three sessions, 24 healthy participants received tACS at 10 Hz or 16.18 Hz (control frequency) or sham stimulation over the bilateral prefrontal cortex before task processing. Resting state EEG was recorded before and after tACS. We also recorded EEG during task processing. Results Relative to sham stimulation, 10 Hz tACS significantly facilitated phonological response speed. This effect was task-specific as tACS did not affect a simple control task. Moreover, 10 Hz tACS significantly increased theta power during phonological decisions. The individual increase in theta power was positively correlated with the behavioral facilitation after 10 Hz tACS. Conclusion Our results show a facilitation of phonological decisions after 10 Hz tACS over the bilateral prefrontal cortex. This might indicate that 10 Hz tACS increased task-related activity in the stimulated area to a level that was optimal for phonological performance. The significant correlation with the individual increase in theta power suggests that the behavioral facilitation might be related to increased theta power during language processing

Phase of beta-frequency tACS over primary motor cortex modulates corticospinal excitability

The assessment of corticospinal excitability by means of transcranial magnetic stimulation-induced motor evoked potentials is an established diagnostic tool in neurophysiology and a widely used procedure in fundamental brain research. However, concern about low reliability of these measures has grown recently. One possible cause of high variability of MEPs under identical acquisition conditions could be the influence of oscillatory neuronal activity on corticospinal excitability. Based on research showing that transcranial alternating current stimulation can entrain neuronal oscillations we here test whether alpha or beta frequency tACS can influence corticospinal excitability in a phase-dependent manner. We applied tACS at individually calibrated alpha- and beta-band oscillation frequencies, or we applied sham tACS. Simultaneous single TMS pulses time locked to eight equidistant phases of the ongoing tACS signal evoked MEPs. To evaluate offline effects of stimulation frequency, MEP amplitudes were measured before and after tACS. To evaluate whether tACS influences MEP amplitude, we fitted one-cycle sinusoids to the average MEPs elicited at the different phase conditions of each tACS frequency. We found no frequency-specific offline effects of tACS. However, beta-frequency tACS modulation of MEPs was phase-dependent. Post hoc analyses suggested that this effect was specific to participants with low (<19 Hz) intrinsic beta frequency. In conclusion, by showing that beta tACS influences MEP amplitude in a phase-dependent manner, our results support a potential role attributed to neuronal oscillations in regulating corticospinal excitability. Moreover, our findings may be useful for the development of TMS protocols that improve the reliability of MEPs as a meaningful tool for research applications or for clinical monitoring and diagnosis. (C) 2018 Elsevier Ltd. All rights reserved

Midfrontal theta transcranial alternating current stimulation modulates behavioural adjustment after error execution

Cognitive control during conflict monitoring, error processing, and post-error adjustment appear to be associated with the occurrence of midfrontal theta (MFϴ). While this association is supported by correlational EEG studies, much less is known about the possible causal link between MFϴ and error and conflict processing. In the present study, we aimed to explore the role of band-specific effects in modulating the error system during a conflict resolution. In turn, we delivered transcranial alternating current stimulation (tACS) at different frequency bands (delta δ, theta θ, alpha α, beta β, gamma γ) and sham stimulation over the medial frontal cortex (MFC) in 36 healthy participants performing a modified version of the Flanker task. Task performance and reports about the sensations (e.g. visual flickering, cutaneous burning) induced by the different frequency bands, were also recorded. We found that online θ-tACS increased the response speed to congruent stimuli after error execution with respect to sham stimulation. Importantly, the accuracy following the errors did not decrease because of speed-accuracy trade off. Moreover, tACS evoked visual and somatosensory sensations were significantly stronger at α-tACS and β-tACS compared to other frequencies. Our findings suggest that theta activity plays a causative role in modulating behavioural adjustments during perceptual choices in a stimulus-response conflict task. © 2018 Federation of European Neuroscience Societies and John Wiley &amp; Sons Lt

Phase Dependency of the Human Primary Motor Cortex and Cholinergic Inhibition Cancelation during Beta tACS

The human motor cortex has a tendency to resonant activity at about 20 Hz so stimulation should more readily entrain neuronal populations at this frequency. We investigated whether and how different interneuronal circuits contribute to such resonance by using transcranial magnetic stimulation (TMS) during transcranial alternating current stimulation (tACS) at motor (20 Hz) and a nonmotor resonance frequency (7 Hz). We tested different TMS interneuronal protocols and triggered TMS pulses at different tACS phases. The effect of cholinergic short-latency afferent inhibition (SAI) was abolished by 20 Hz tACS, linking cortical beta activity to sensorimotor integration. However, this effect occurred regardless of the tACS phase. In contrast, 20 Hz tACS selectively modulated MEP size according to the phase of tACS during single pulse, GABAAergic short-interval intracortical inhibition (SICI) and glutamatergic intracortical facilitation (ICF). For SICI this phase effect was more marked during 20 Hz stimulation. Phase modulation of SICI also depended on whether or not spontaneous beta activity occurred at ~20 Hz, supporting an interaction effect between tACS and underlying circuit resonances. The present study provides in vivo evidence linking cortical beta activity to sensorimotor integration, and for beta oscillations in motor cortex being promoted by resonance in GABAAergic interneuronal circuits

Beta Power May Mediate the Effect of Gamma-TACS on Motor Performance

Transcranial alternating current stimulation (tACS) is becoming an important method in the field of motor rehabilitation because of its ability to non-invasively influence ongoing brain oscillations at arbitrary frequencies. However, substantial variations in its effect across individuals are reported, making tACS a currently unreliable treatment tool. One reason for this variability is the lack of knowledge about the exact way tACS entrains and interacts with ongoing brain oscillations. The present crossover stimulation study on 20 healthy subjects contributes to the understanding of cross-frequency effects of gamma (70 Hz) tACS over the contralateral motor cortex by providing empirical evidence which is consistent with a role of low- (12~-20 Hz) and high- (20-~30 Hz) beta power as a mediator of gamma-tACS on motor performance.Comment: 7 pages, 5 figures, in Proceedings of IEEE Engineering in Medicine and Biology Conference, July 2019 (IEEE license notice

Does alpha phase modulate visual target detection? Three experiments with tACS-phase-based stimulus presentation

In recent years, the influence of alpha (7–13 Hz) phase on visual processing has received a lot of attention. Magneto‐/encephalography (M/EEG) studies showed that alpha phase indexes visual excitability and task performance. Studies with transcranial alternating current stimulation (tACS) aim to modulate oscillations and causally impact task performance. Here, we applied right occipital tACS (O2 location) to assess the functional role of alpha phase in a series of experiments. We presented visual stimuli at different pre‐determined, experimentally controlled, phases of the entraining tACS signal, hypothesizing that this should result in an oscillatory pattern of visual performance in specifically left hemifield detection tasks. In experiment 1, we applied 10 Hz tACS and used separate psychophysical staircases for six equidistant tACS‐phase conditions, obtaining contrast thresholds for detection of visual gratings in left or right hemifield. In experiments 2 and 3, tACS was at EEG‐based individual peak alpha frequency. In experiment 2, we measured detection rates for gratings with (pseudo‐)fixed contrast. In experiment 3, participants detected brief luminance changes in a custom‐built LED device, at eight equidistant alpha phases. In none of the experiments did the primary outcome measure over phase conditions consistently reflect a one‐cycle sinusoid. However, post hoc analyses of reaction times (RT) suggested that tACS alpha phase did modulate RT for specifically left hemifield targets in both experiments 1 and 2 (not measured in experiment 3). This observation requires future confirmation, but is in line with the idea that alpha phase causally gates visual inputs through cortical excitability modulation

Brain-state determines learning improvements after transcranial alternating-current stimulation to frontal cortex

Published in final edited form as: Brain Stimul. 2018 ; 11(4): 723–726. doi:10.1016/j.brs.2018.02.008BACKGROUND Theories of executive control propose that communication between medial frontal cortex (MFC) and lateral prefrontal cortex (lPFC) is critical for learning. 6-Hz phase synchronization may be the mechanism by which neural activity between MFC and lPFC is coordinated into a functional network. Recent evidence suggests that switching from eyes closed to open may induce a change in brain-state reflected by enhanced executive control and related functional connectivity. OBJECTIVE/HYPOTHESIs To examine whether causal manipulation of MFC and lPFC can improve learning according to the brain-state induced by switching from eyes closed to open. METHODS Within-subjects, sham-controlled, double-blind study of 30 healthy subjects, each receiving 6-Hz in-phase high definition transcranial alternating-current stimulation (HD-tACS) applied to MFC and right lPFC prior to performing a time estimation task. RESULTS HD-tACS with eyes open improved learning ability relative to sham, whereas HD-tACS with eyes closed had no significant effect on behavior. CONCLUSION Results suggest a phase-sensitive mechanism in frontal cortex mediates components of learning performance in a state-dependent manner.Accepted manuscrip

Modulating brain oscillations to drive brain function

Do neuronal oscillations play a causal role in brain function? In a study in this issue of PLOS Biology, Helfrich and colleagues address this long-standing question by attempting to drive brain oscillations using transcranial electrical current stimulation. Remarkably, they were able to manipulate visual perception by forcing brain oscillations of the left and right visual hemispheres into synchrony using oscillatory currents over both hemispheres. Under this condition, human observers more often perceived an inherently ambiguous visual stimulus in one of its perceptual instantiations. These findings shed light on the mechanisms underlying neuronal computation. They show that it is the neuronal oscillations that drive the visual experience, not the experience driving the oscillations. And they indicate that synchronized oscillatory activity groups brain areas into functional networks. This points to new ways for controlled experimental and possibly also clinical interventions for the study and modulation of brain oscillations and associated functions