Dynamic alpha power modulations and slow negative potentials track natural shifts of spatio-temporal attention.

Alpha power modulations and slow negative potentials have previously been associated with anticipatory processes in spatial and temporal top-down attention. In typical experimental designs, however, neural responses triggered by transient stimulus onsets can interfere with attention-driven activity patterns and our interpretation of such. Here, we investigated these signatures of spatio-temporal attention in a dynamic paradigm free from potentially confounding stimulus-driven activity using electroencephalography. Participants attended the cued side of a bilateral stimulus rotation and mentally counted how often one of two remembered sample orientations (i.e., the target) was displayed while ignoring the uncued side and non-target orientation. Afterwards, participants performed a delayed match-to-sample task, in which they indicated if the orientation of a probe stimulus matched the corresponding sample orientation (previously target or non-target). We observed dynamic alpha power reductions and slow negative waves around task-relevant points in space and time (i.e., onset of the target orientation in the cued hemifield) over posterior electrodes contralateral to the locus of attention. In contrast to static alpha power lateralization, these dynamic signatures correlated with subsequent memory performance (primarily detriments for matching probes of the non-target orientation), suggesting a preferential allocation of attention to task-relevant locations and time points at the expense of reduced resources and impaired performance for information outside the current focus of attention. Our findings suggest that humans can naturally and dynamically focus their attention at relevant points in space and time and that such spatio-temporal attention shifts can be reflected by dynamic alpha power modulations and slow negative potentials.

Event-related Potentials Indicate Target Processing in the Absence of Distractor Suppression during Rapid Serial Visual Presentation.

BACKGROUND: In our modern world we are exposed to a steady stream of information containing important as well as irrelevant information. Therefore, our brains have to constantly select relevant over distracting items and further process the selected information. Whereas there is good evidence that even in rapid serial streams of presented information relevant targets can be actively selected, it is less clear whether and how distracting information is de-selected and suppressed in such scenarios. METHODS: To address this issue we recorded electroencephalographic activity during a rapid serial visual presentation paradigm in which healthy, young human volunteers had to encode visual targets into short-term memory while salient visual distractors and neutral filler items needed to be ignored. Event-related potentials were analyzed in 3D source space and compared between stimulus types. RESULTS: A negative wave between around 170 and 230 ms after stimulus onset resembling the N2pc component was identified that dissociated between target stimuli and distractors as well as filler items. This wave appears to reflect target selection processes. However, there was no electrophysiological signature identified that would indicate an active distractor suppression mechanism. CONCLUSIONS: The obtained results suggest that unlike in situations where target stimuli and distractors are presented simultaneously, targets can be selected without the need for active suppression of distracting information in serial presentations with a clear and regular temporal structure. It is assumed that temporal expectation supports efficient target selection by the brain.

Alpha-tACS alters attentional control but not cognitive functions as video games do: A psychophysical investigation based on the theory of visual attention.

Video game players' faster speed of information processing has been shown to coincide with altered posterior alpha power modulation, that is, brain oscillatory activity around 10 Hz. Thus, it was proposed that improved cognitive processing in video game players may be related to differential alpha activity. However, a causal relationship thereof has not yet been established. We addressed this by conducting a non-invasive brain stimulation study to demonstrate that modulating alpha power using transcranial alternating current stimulation (tACS) may impact on speed of information processing. Furthermore, we aimed to show that this effect correlated with altered attentional control, for example, visuospatial attention and/or top-down control processing, given that this has been suggested to contribute to video gaming effects. Therefore, we recruited 19 non-video game players to undergo one of five brain stimulation conditions while performing a visual short-term memory task at five different days, respectively. Thus, we applied tACS either at 10 Hz (alpha frequency) or at 16.18 Hz (control frequency) either over their left or right posterior parietal cortex (PPC) or a sham stimulation. Individuals' speed of information processing, visuospatial attention and top-down control processing were operationalised using a computational modelling approach based on the theory of visual attention. We found that alpha-tACS applied over individuals' left PPC altered their visuospatial attention orientation but not their speed of information processing. Thus, we were not able to establish a causal relationship between speed of information processing and altered visuospatial attention processing through alpha power modulation using non-invasive brain stimulation.

Spatio-temporal dynamics of oscillatory brain activity during the observation of actions and interactions between point-light agents.

Predicting actions from non-verbal cues and using them to optimise one's response behaviour (i.e. interpersonal predictive coding) is essential in everyday social interactions. We aimed to investigate the neural correlates of different cognitive processes evolving over time during interpersonal predictive coding. Thirty-nine participants watched two agents depicted by moving point-light stimuli while an electroencephalogram (EEG) was recorded. One well-recognizable agent performed either a 'communicative' or an 'individual' action. The second agent either was blended into a cluster of noise dots (i.e. present) or was entirely replaced by noise dots (i.e. absent), which participants had to differentiate. EEG amplitude and coherence analyses for theta, alpha and beta frequency bands revealed a dynamic pattern unfolding over time: Watching communicative actions was associated with enhanced coupling within medial anterior regions involved in social and mentalising processes and with dorsolateral prefrontal activation indicating a higher deployment of cognitive resources. Trying to detect the agent in the cluster of noise dots without having seen communicative cues was related to enhanced coupling in posterior regions for social perception and visual processing. Observing an expected outcome was modulated by motor system activation. Finally, when the agent was detected correctly, activation in posterior areas for visual processing of socially relevant features was increased. Taken together, our results demonstrate that it is crucial to consider the temporal dynamics of social interactions and of their neural correlates to better understand interpersonal predictive coding. This could lead to optimised treatment approaches for individuals with problems in social interactions.

Theta:gamma phase coupling and evoked gamma activity reflect the fidelity of mental templates during memory matching in visual perception.

Top-down predictions of future events shaped by prior experience are an important control mechanism to allocate limited attentional resources more efficiently and are thought to be implemented as mental templates stored in memory. Increased evoked gamma activity and theta:gamma phase-phase coupling over parieto-occipital areas have previously been observed when mental templates meet matching visual stimuli. Here, we investigated how these signatures evolve during the formation of new mental templates and how they relate to the fidelity of such. Based on single-trial feedback, participants learned to classify target shapes as matching or mismatching with preceding cue sequences. In the end of the experiment, they were asked to freely reproduce targets as means of template fidelity. We observed fidelity-dependent increments of matching-related gamma phase locking and theta:gamma phase coupling in early visual areas around 100-200-ms poststimulus over time. Theta:gamma phase synchronization and evoked gamma activity might serve as complementary signatures of memory matching in visual perception; theta:gamma phase synchronization seemingly most important in early phases of learning and evoked gamma activity being essential for transition of mental templates into long-term memory.

Modulating verbal working memory with fronto-parietal transcranial electric stimulation at theta frequency: Does it work?

Oscillatory theta activity in a fronto-parietal network has been associated with working memory (WM) processes and may be directly related to WM performance. In their seminal study, Polanía et al. (2012) (de-)coupled a fronto-parietal theta-network by applying transcranial alternating current stimulation (tACS), and showed that anti-phase tACS led to slower and in-phase tACS to faster response times in a verbal WM task compared to placebo stimulation. In the literature, this 'synchronization-desynchronization' effect has only been partly replicated, and electric field modelling suggests that it might not be the fronto-parietal network that is primarily stimulated during in-phase tACS with a shared return electrode. This provides one possible reason for inconsistency in the literature. In this study, we aimed to reproduce the findings reported by Polanía et al. (2012). We also aimed to investigate whether in-phase theta tACS with multiple close-by return electrodes for focal stimulation of the frontal and the parietal cortex will have at least as much of a facilitatory effect as the in-phase stimulation as indicated by Polania et al. (2012). In a single-trial distributional analysis, we explored whether mean, variation and right-skewness of the response time distribution are affected. Against our hypothesis, we found no 'synchronization-desynchronization' effect by fronto-parietal theta tACS on response times using the same delayed letter discrimination task and stimulation parameters in two experiments, both between-subjects and within-subjects. However, we could show that in a more demanding 3-back task, fronto-parietal in-phase and in-phase focal theta tACS substantially improved task performance compared to placebo stimulation.

EEG cross-frequency phase synchronization as an index of memory matching in visual search.

Visual perception is influenced by our expectancies about incoming sensory information. It is assumed that mental templates of expected sensory input are created and compared to actual input, which can be matching or not. When such mental templates are held in working memory, cross-frequency phase synchronization (CFS) between theta and gamma band activity has been proposed to serve matching processes between prediction and sensation. We investigated how this is affected by the number of activated templates that could be matched by comparing conditions where participants had to keep either one or multiple templates in mind for successful visual search. We found a transient CFS between EEG theta and gamma activity in an early time window around 150 ms after search display presentation, in right hemispheric parietal cortex. Our results suggest that for single template conditions, stronger transient theta-gamma CFS at posterior sites contralateral to target presentation can be observed than for multiple templates. This can be interpreted as evidence to the idea of sequential attentional templates. But mainly, it is understood in line with previous theoretical accounts strongly arguing for transient synchronization between posterior theta and gamma phase as a neural correlate of matching incoming sensory information with contents from working memory and as evidence for limitations in memory matching during multiple template search.

Enhancing visual motion discrimination by desynchronizing bifocal oscillatory activity.

Visual motion discrimination involves reciprocal interactions in the alpha band between the primary visual cortex (V1) and mediotemporal areas (V5/MT). We investigated whether modulating alpha phase synchronization using individualized multisite transcranial alternating current stimulation (tACS) over V5 and V1 regions would improve motion discrimination. We tested 3 groups of healthy subjects with the following conditions: (1) individualized In-Phase V1alpha-V5alpha tACS (0° lag), (2) individualized Anti-Phase V1alpha-V5alpha tACS (180° lag) and (3) sham tACS. Motion discrimination and EEG activity were recorded before, during and after tACS. Performance significantly improved in the Anti-Phase group compared to the In-Phase group 10 and 30 min after stimulation. This result was explained by decreases in bottom-up alpha-V1 gamma-V5 phase-amplitude coupling. One possible explanation of these results is that Anti-Phase V1alpha-V5alpha tACS might impose an optimal phase lag between stimulation sites due to the inherent speed of wave propagation, hereby supporting optimized neuronal communication.

The speed of parietal theta frequency drives visuospatial working memory capacity.

The speed of theta brain oscillatory activity is thought to play a key role in determining working memory (WM) capacity. Individual differences in the length of a theta cycle (ranging between 4 and 7 Hz) might determine how many gamma cycles (>30 Hz) can be nested into a theta wave. Gamma cycles are thought to represent single memory items; therefore, this interplay could determine individual memory capacity. We directly tested this hypothesis by means of parietal transcranial alternating current stimulation (tACS) set at slower (4 Hz) and faster (7 Hz) theta frequencies during a visuospatial WM paradigm. Accordingly, we found that 4-Hz tACS enhanced WM capacity, while 7-Hz tACS reduced WM capacity. Notably, these effects were found only for items presented to the hemifield contralateral to the stimulation site. This provides causal evidence for a frequency-dependent and spatially specific organization of WM storage, supporting the theta-gamma phase coupling theory of WM capacity.

Slow Theta tACS of the Right Parietal Cortex Enhances Contralateral Visual Working Memory Capacity.

Recent research suggests alteration of visual working memory capacity by modulation of parietal theta frequency via transcranial alternating current stimulation (tACS). However, it remains to be clarified whether this effect is partly driven by co-stimulation of prefrontal cortex and subcortical structures. It was hypothesized that focal tACS over the parietal lobe without additional prefrontal or subcortical stimulation should lead to similar effects as reported in the literature. Healthy, young participants were tested on a visual working memory paradigm while receiving either focal parietal tACS at 4 Hz, at 7 Hz or sham stimulation. Focal right posterior 4 Hz tACS led to increased working memory capacity strictly for the visual hemifield contralateral to stimulation. Exclusive stimulation of posterior cortex by 4 Hz tACS replicates effects recently reported in literature, confirming that stimulation of the prefrontal cortex or subcortical structures are not a primary driver of these observations.

The involvement of alpha oscillations in voluntary attention directed towards encoding episodic memories.

Forming episodic memories is often driven by top-down processes of allocating attention towards voluntarily remembering the details of an episode. This attention orientation is needed to make sure that information is encoded for later remembering. Here we designed an episodic long-term memory (LTM) EEG experiment where we examined brain oscillatory activity associated with attention allocation towards the temporal link between an item and its context. The remembering of this temporal conjunction is crucial for item-context binding and hence for the formation of episodic memories. Participants saw a background picture and a word in a central position on a computer screen and were instructed to memorise (a) the picture only, (b) the word, (c) both individually (i.e. ignoring their co-occurrence) and (d) both with them being presented together. Attention allocation towards item-context binding was associated with oscillatory alpha desynchronization in the upper alpha band (10-13 Hz) over dominantly left posterior brain areas. The results highlight the role of alpha desynchronization in voluntary attention allocation towards the temporal conjunction of item and its context in episodic binding and the involvement of posterior brain areas. The pattern of results suggest that they most likely reflect additional visual processes recruited by attentional mechanisms and do not tap into neural processes of item-context binding per se. Moreover, it indicates that the involvement of alpha oscillations in cognitive processes may be more complex.

Distributed Cortical Phase Synchronization in the EEG Reveals Parallel Attention and Working Memory Processes Involved in the Attentional Blink.

Attentional blink (AB) describes a visuo-perceptual phenomenon in which the second of 2 targets within a rapid serial visual presentation stream is not detected. There are several cognitive models attempting to explain the fundamentals of this information processing bottleneck. Here, we used electroencephalographic recordings and the analysis of interregional phase synchronization of rhythmical brain activity to investigate the neural bases of the AB. By investigating the time course of interregional phase synchronization separately for trials in which participants failed to report the second target correctly (AB trials) and trials in which no AB occurred, and by clustering interregional connections based on their functional similarity, it was possible to define several distinct cortical networks. Analyzing these networks comprising phase synchronization--over a large spectrum of brain frequencies from theta to gamma activity--it was possible to identify neural correlates for cognitive subfunctions involved in the AB, such as the encoding of targets into working memory, tuning of attentional filters, and the recruitment of general cognitive resources. This parallel activation of functionally distinct neural processes substantiates the eligibility of several cognitive models on the AB.

EEG theta phase coupling during executive control of visual working memory investigated in individuals with schizophrenia and in healthy controls.

In healthy humans, it has been shown that executive functions are associated with increased frontal-midline EEG theta activity and theta phase coupling between frontal and posterior brain regions. In individuals with schizophrenia, central executive functions are supposed to be heavily impaired. Given that theta phase coupling is causally involved in central executive functions, one would expect that patients with an executive function deficit should display abnormal EEG theta synchronization. We therefore investigated executive functioning in 21 healthy controls and 21 individuals with schizophrenia while they performed a visuospatial delayed match to sample task. The task required either high executive demands (manipulation of content in working memory [WM]) or low executive demands (retention of WM content). In addition, WM load (one vs. three items) was varied. Results indicated higher frontal theta activity for manipulation processes than for retention processes in patients with schizophrenia, as compared with healthy controls, independently of WM load. Furthermore, individuals with schizophrenia revealed a reduction in theta phase coupling during early stages of the delay period for retention, as well as for manipulation processes at high-WM loads. Deviations in theta phase coupling in individuals with schizophrenia were mainly characterized by aberrant fronto-posterior connections, but also by attenuated posterior connections during manipulation of high-WM load. To conclude, fronto-parietal theta coupling seems to be substantially involved in executive control, whereas frontal theta activity seems to reflect general task demands, such as deployment of attentional resources during WM.

Two brakes are better than one: the neural bases of inhibitory control of motor memory traces.

Inhibitory control of actions is one important aspect in daily life to warrant adequate context related behavior. Alpha activity (oscillatory brain activity around 10Hz) has been suggested to play a major role for the implementation of inhibitory control. In the present study electrophysiological correlates of voluntary suppression of acquired, memorized motor actions have been compared to the suppression of novel motor actions. Multichannel EEG analyses of alpha power and alpha phase coherence were used. Healthy subjects were asked to inhibit the execution of either well-trained, memorized or untrained, novel sequential finger movements depending on the respective context. An increase of focal upper alpha activity at bilateral sensorimotor cortices was found during suppression of movements independent of whether these were memorized or novel. This represents a memory unspecific mechanism of motor cortical inhibition. In contrast, interregional phase synchronization between frontal and (left) central recording sites showed a differential effect with decoupling during suppression of memorized movements which was not the case with novel ones. Increase of fronto-central coupling at upper alpha frequency during retrieval of the memory trace and decrease during suppression of retrieval were obtained. This further supports the view of the functional relevance of upper alpha oscillations as a mechanism of context-dependent sustained inhibition of memory contents.

Subthalamic nucleus stimulation restores the efferent cortical drive to muscle in parallel to functional motor improvement.

Pathological synchronization in large-scale motor networks constitutes a pathophysiological hallmark of Parkinson's disease (PD). Corticomuscular synchronization in PD is pronounced in lower frequency bands (< 10 Hz), whereas efficient cortical motor integration in healthy persons is driven in the beta frequency range. Electroencephalogram and electromyogram recordings at rest and during an isometric precision grip task were performed in four perioperative sessions in 10 patients with PD undergoing subthalamic nucleus deep-brain stimulation: (i) 1 day before (D0); (ii) 1 day after (D1); (iii) 8 days after implantation of macroelectrodes with stimulation off (D8StimOff); and (iv) on (D8StimOn). Analyses of coherence and phase delays were performed in order to challenge the effects of microlesion and stimulation on corticomuscular coherence (CMC). Additionally, local field potentials recorded from the subthalamic nucleus on D1 allowed comprehensive mapping of motor-related synchronization in subthalamocortical and cerebromuscular networks. Motor performance improved at D8StimOn compared with D0 and D8StimOff paralleled by a reduction of muscular activity and CMC in the theta band (3.9-7.8 Hz) and by an increase of CMC in the low-beta band (13.7-19.5 Hz). Efferent motor cortical drives to muscle presented mainly below 10 Hz on D8StimOff that were suppressed on D8StimOn and occurred on higher frequencies from 13 to 45 Hz. On D1, coherence of the high-beta band (20.5-30.2 Hz) increased during movement compared with rest in subthalamomuscular and corticomuscular projections, whereas it was attenuated in subcorticocortical projections. The present findings lend further support to the concept of pathological network synchronization in PD that is beneficially modulated by stimulation.

Brain rhythms: How control gets into working memory.

New research suggests that frontal midline theta EEG activity in humans controls activity in parietal cortex associated with memory maintenance. In turn, the speed of this frontal theta is modulated by the number of items to be handled, potentially indicating strong bidirectional communication within a fronto-parietal network.

Spatio-Temporal Brain Dynamic Differences in Fluid Intelligence.

Human fluid intelligence is closely linked to the sequential solving of complex problems. It has been associated with a distributed cognitive control or multiple-demand (MD) network, comprising regions of lateral frontal, insular, dorsomedial frontal, and parietal cortex. Previous neuroimaging research suggests that the MD network may orchestrate the allocation of attentional resources to individual parts of a complex task: in a complex target detection task with multiple independent rules, applied one at a time, reduced response to rule-critical events across the MD network in lower fluid intelligence was observed. This was in particular the case with increasing task complexity (i.e., larger sets of rules), and was accompanied by impairment in performance. Here, we examined the early spatiotemporal neural dynamics of this process in electroencephalography (EEG) source analyses using a similar task paradigm. Levels of fluid intelligence specifically predicted early neural responses in a left inferiorparietal MD region around 200-300 ms post stimulus onset. Evoked source amplitudes in left parietal cortex within this early time window also correlated with behavioural performance measures. Like in previous research, we observed impaired performance in lower fluid intelligence with increasing number of task rules. This links fluid intelligence to a process of attentional focus on those parts of a task that are most critical for the current behaviour. Within the MD system, our time re-resolved measures suggest that the left parietal cortex specifically impacts on early processes of attentional focus on task critical features. This is novel evidence on the neurocognitive correlates of fluid intelligence suggesting that individual differences are critically linked to an early process of attentional focus on task-relevant information, which is supported by left parietal MD regions.

Seeing a Bayesian ghost: Sensorimotor activation leads to an illusory social perception.

Based on our prior experiences we form social expectations and anticipate another person's response. Under certain conditions, these expectations can be so strong that they lead to illusory perception of another person who is actually not there (i.e., seeing a Bayesian ghost). We used EEG to investigate the neural correlates of such illusory social perception. Our results showed that activation of the premotor cortex predicted the occurrence of the Bayesian ghost, whereas its actual appearance was later accompanied by activation in sensorimotor and adjacent parietal regions. These findings confirm that our perception of others is so strongly affected by prior expectations, in such a way they can prompt illusory social perceptions associated with activity change in brain regions relevant for action perception. They also contribute to a better understanding of social interaction in healthy individuals as well as persons with mental illnesses, which can be characterized by illusory perception and social interaction difficulties.

The truth about lying: inhibition of the anterior prefrontal cortex improves deceptive behavior.

Recent neuroimaging studies have indicated a predominant role of the anterior prefrontal cortex (aPFC) in deception and moral cognition, yet the functional contribution of the aPFC to deceptive behavior remains unknown. We hypothesized that modulating the excitability of the aPFC by transcranial direct current stimulation (tDCS) could reveal its functional contribution in generating deceitful responses. Forty-four healthy volunteers participated in a thief role-play in which they were supposed to steal money and then to attend an interrogation with the Guilty Knowledge Test. During the interrogation, participants received cathodal, anodal, or sham tDCS. Remarkably, inhibition of the aPFC by cathodal tDCS did not lead to an impairment of deceptive behavior but rather to a significant improvement. This effect manifested in faster reaction times in telling lies, but not in telling the truth, a decrease in sympathetic skin-conductance response and feelings of guilt while deceiving the interrogator and a significantly higher lying quotient reflecting skillful lying. Increasing the excitability of the aPFC by anodal tDCS did not affect deceptive behavior, confirming the specificity of the stimulation polarity. These findings give causal support to recent correlative data obtained by functional magnetic resonance imaging studies indicating a pivotal role of the aPFC in deception.

Cause or consequence? Alpha oscillations in visuospatial attention.

A well-established finding in the literature of human studies is that alpha activity (rhythmical brain activity around 10 Hz) shows retinotopic amplitude modulation during shifts in visual attention. Thus, it has long been argued that alpha amplitude modulation might play a crucial role in attention-driven alterations in visual information processing. Recently, there has been a revival of the topic, driven in part by new studies directly investigating the possible causal relationship between alpha activity and responses to visual input, both neuronally and perceptually. Here, we discuss evidence for and against a causal role of alpha activity in visual attentional processing. We conclude with hypotheses regarding the mechanisms by which top-down-modulated alpha activity in the parietal cortex might select visual information for attentive processing.