Relating alpha power modulations to competing visuospatial attention theories

Visuospatial attention theories often propose hemispheric asymmetries underlying the control of attention. In general support of these theories, previous EEG/MEG studies have shown that spatial attention is associated with hemispheric modulation of posterior alpha power (gating by inhibition). However, since measures of alpha power are typically expressed as lateralization scores, or collapsed across left and right attention shifts, the individual hemispheric contribution to the attentional control mechanism remains unclear. This is, however, the most crucial and decisive aspect in which the currently competing attention theories continue to disagree. To resolve this long-standing conflict, we derived predictions regarding alpha power modulations from Heilman's hemispatial theory and Kinsbourne's interhemispheric competition theory and tested them empirically in an EEG experiment. We used an attention paradigm capable of isolating alpha power modulation in two attentional states, namely attentional bias in a neutral cue condition and spatial orienting following directional cues. Differential alpha modulations were found for both hemispheres across conditions. When anticipating peripheral visual targets without preceding directional cues (neutral condition), posterior alpha power in the left hemisphere was generally lower and more strongly modulated than in the right hemisphere, in line with the interhemispheric competition theory. Intriguingly, however, while alpha power in the right hemisphere was modulated by both, cue-directed leftward and rightward attention shifts, the left hemisphere only showed modulations by rightward shifts of spatial attention, in line with the hemispatial theory. This suggests that the two theories may not be mutually exclusive, but rather apply to different attentional states

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

Calibrating rhythmic stimulation parameters to individual electroencephalography markers: The consistency of individual alpha frequency in practical lab settings

Rhythmic stimulation can be applied to modulate neuronal oscillations. Such ‘entrainment’ is optimized when stimulation frequency is individually calibrated based on magneto/encephalography markers. It remains unknown how consistent such individual markers are across days/sessions, within a session, or across cognitive states, hemispheres and estimation methods, especially in a realistic, practical, lab setting. We here estimated individual alpha frequency (IAF) repeatedly from short electroencephalography (EEG) measurements at rest or during an attention task (cognitive state), using single parieto-occipital electrodes in 24 participants on 4 days (between-sessions), with multiple measurements over an hour on 1 day (within-session). First, we introduce an algorithm to automatically reject power spectra without a sufficiently clear peak to ensure unbiased IAF estimations. Then we estimated IAF via the traditional ‘maximum’ method and a ‘Gaussian fit’ method. IAF was reliable within- and between-sessions for both cognitive states and hemispheres, though task-IAF estimates tended to be more variable. Overall, the ‘Gaussian fit’ method was more reliable than the ‘maximum’ method. Furthermore, we evaluated how far from an approximated ‘true’ task-related IAF the selected ‘stimulation frequency’ was, when calibrating this frequency based on a short rest-EEG, a short task-EEG, or simply selecting 10 Hz for all participants. For the ‘maximum’ method, rest-EEG calibration was best, followed by task-EEG, and then 10 Hz. For the ‘Gaussian fit’ method, rest-EEG and task-EEG-based calibration were similarly accurate, and better than 10 Hz. These results lead to concrete recommendations about valid, and automated, estimation of individual oscillation markers in experimental and clinical settings

Neural tracking of phrases in spoken language comprehension is automatic and task-dependent

Linguistic phrases are tracked in sentences even though there is no one-to-one acoustic phrase marker in the physical signal. This phenomenon suggests an automatic tracking of abstract linguistic structure that is endogenously generated by the brain. However, all studies investigating linguistic tracking compare conditions where either relevant information at linguistic timescales is available, or where this information is absent altogether (e.g., sentences versus word lists during passive listening). It is therefore unclear whether tracking at phrasal timescales is related to the content of language, or rather, results as a consequence of attending to the timescales that happen to match behaviourally relevant information. To investigate this question, we presented participants with sentences and word lists while recording their brain activity with magnetoencephalography (MEG). Participants performed passive, syllable, word, and word-combination tasks corresponding to attending to four different rates: one they would naturally attend to, syllable-rates, word-rates, and phrasal-rates, respectively. We replicated overall findings of stronger phrasal-rate tracking measured with mutual information for sentences compared to word lists across the classical language network. However, in the inferior frontal gyrus (IFG) we found a task effect suggesting stronger phrasal-rate tracking during the word-combination task independent of the presence of linguistic structure, as well as stronger delta-band connectivity during this task. These results suggest that extracting linguistic information at phrasal rates occurs automatically with or without the presence of an additional task, but also that IFG might be important for temporal integration across various perceptual domains

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

“Broadband Alpha Transcranial Alternating Current Stimulation”: Exploring a new biologically calibrated brain stimulation protocol

Transcranial alternating current stimulation (tACS) can be used to study causal contributions of oscillatory brain mechanisms to cognition and behavior. For instance, individual alpha frequency (IAF) tACS was reported to enhance alpha power and impact visuospatial attention performance. Unfortunately, such results have been inconsistent and difficult to replicate. In tACS, stimulation generally involves one frequency, sometimes individually calibrated to a peak value observed in an M/EEG power spectrum. Yet, the ‘peak’ actually observed in such power spectra often contains a broader range of frequencies, raising the question whether a biologically calibrated tACS protocol containing this fuller range of alpha-band frequencies might be more effective. Here, we introduce ‘Broadband-alpha-tACS’, a complex individually calibrated electrical stimulation protocol. We band-pass filtered left posterior resting-state EEG data around the IAF (+/- 2 Hz), and converted that time series into an electrical waveform for tACS stimulation of that same left posterior parietal cortex location. In other words, we stimulated a brain region with a ‘replay’ of its own alpha-band frequency content, based on spontaneous activity. Within-subjects (N=24), we compared to a sham tACS session the effects of broadband-alpha tACS, power-matched spectral inverse (‘alpha-removed’) control tACS, and individual alpha frequency tACS, on EEG alpha power and performance in an endogenous attention task previously reported to be affected by alpha tACS. Broadband-alpha-tACS significantly modulated attention task performance (i.e., reduced the rightward visuospatial attention bias in trials without distractors, and reduced attention benefits). Alpha-removed tACS also reduced the rightward visuospatial attention bias. IAF-tACS did not significantly modulate attention task performance compared to sham tACS, but also did not statistically significantly differ from broadband-alpha-tACS. This new broadband-alpha tACS approach seems promising, but should be further explored and validated in future studies