Mechanisms of sustained perceptual entrainment after stimulus offset.

Temporal alignment of neural activity to rhythmic stimulation has been suggested to result from a resonating internal neural oscillator mechanism, but can also be explained by interval-based temporal prediction. Here, we investigate behavioural and brain responses in the post-stimulation period to compare an oscillatory versus an interval-based account. Hickok et al.'s (2015) behavioural paradigm yielded results that relate to a neural oscillatory entrainment mechanism. We adapted the paradigm to an event-related potential (ERP) suitable design: a periodic sequence was followed, in half of the trials, by near-threshold targets embedded in noise. The targets were played in various phases in relation to the preceding sequences' period. Participants had to detect whether targets were played or not, and their EEG was recorded. Both behavioural results and the P300 component of the ERP were not only partially consistent with an oscillatory mechanism but also partially consistent with an interval-based attentional gain mechanism. Instead, data obtained in the post-entrainment period can best be explained with a combination of both mechanisms.

Early sensory gain control is dominated by obligatory and global feature-based attention in top-down shifts of combined spatial and feature-based attention.

What are the dynamics of global feature-based and spatial attention, when deployed together? In an attentional shifting experiment, flanked by three control experiments, we investigated neural temporal dynamics of combined attentional shifts. For this purpose, orange- and blue-frequency-tagged spatially overlapping Random Dot Kinematograms were presented in the left and right visual hemifield to elicit continuous steady-state-visual-evoked-potentials. After being initially engaged in a fixation cross task, participants were at some point in time cued to shift attention to one of the Random Dot Kinematograms, to detect and respond to brief coherent motion events, while ignoring all such events in other Random Dot Kinematograms. The analysis of steady-state visual-evoked potentials allowed us to map time courses and dynamics of early sensory-gain modulations by attention. This revealed a time-invariant amplification of the to-be attended color both at the attended and the unattended side, followed by suppression for the to-be-ignored color at attended and unattended sides. Across all experiments, global and obligatory feature-based selection dominated early sensory gain modulations, whereas spatial attention played a minor modulatory role. However, analyses of behavior and neural markers such as alpha-band activity and event-related potentials to target- and distractor-event processing, revealed clear modulations by spatial attention.

Feature-based Attentional Amplitude Modulations of the Steady-state Visual Evoked Potentials Reflect Blood Oxygen Level Dependent Changes in Feature-sensitive Visual Areas.

Recent EEG studies have investigated basic principles of feature-based attention by means of frequency-tagged random dot kinematograms in which different colors are simultaneously presented at different temporal frequencies to elicit steady-state visual evoked potentials (SSVEPs). These experiments consistently showed global facilitation of the to-be-attended random dot kinematogram-a basic principle of feature-based attention. SSVEP source estimation suggested that posterior visual cortex from V1 to area hMT+/V5 is broadly activated by frequency-tagged stimuli. What is presently unknown is whether the feature-based attentional facilitation of SSVEPs is a rather unspecific neural response including all visual areas that follow the "on/off," or whether SSVEP feature-based amplitude enhancements are driven by activity in visual areas most sensitive to a specific feature, such as V4v in the case of color. Here, we leverage multimodal SSVEP-fMRI recordings in human participants and a multidimensional feature-based attention paradigm to investigate this question. Attending to shape produced significantly greater SSVEP-BOLD covariation in primary visual cortex compared with color. SSVEP-BOLD covariation during color selection increased along the visual hierarchy, with greatest values in areas V3 and V4. Importantly, in area hMT+/V5, we found no differences between shape and color selection. Results suggest that SSVEP amplitude enhancements in feature-based attention is not an unspecific enhancement of neural activity in all visual areas following the "on/off." These findings open new avenues to investigating neural dynamics of competitive interactions in specific visual areas sensitive to a certain feature in a more economical way and better temporal resolution compared with fMRI.

The Distractor Positivity Component and the Inhibition of Distracting Stimuli.

There has been a long-lasting debate about whether salient stimuli, such as uniquely colored objects, have the ability to automatically distract us. To resolve this debate, it has been suggested that salient stimuli do attract attention but that they can be suppressed to prevent distraction. Some research supporting this viewpoint has focused on a newly discovered ERP component called the distractor positivity (PD), which is thought to measure an inhibitory attentional process. This collaborative review summarizes previous research relying on this component with a specific emphasis on how the PD has been used to understand the ability to ignore distracting stimuli. In particular, we outline how the PD component has been used to gain theoretical insights about how search strategy and learning can influence distraction. We also review alternative accounts of the cognitive processes indexed by the PD component. Ultimately, we conclude that the PD component is a useful tool for understanding inhibitory processes related to distraction and may prove to be useful in other areas of study related to cognitive control.

Sustained selective attention to chromatic information enhances visuocortical gain at the population level.

Prior work in selective attention research has shown that colour-selective attention enhances neural activity in visuocortical areas sensitive to the attended colour while suppressing activity in areas sensitive to ignored colours. However, it is currently unclear whether this effect is limited to attending to specific colour hues or extends to chromatic information more broadly. To investigate this question, we used steady-state visual evoked potentials (ssVEPs) frequency tagging to quantify participants' visuocortical responses to specific elements embedded in arrays of flickering, randomly moving mid-complex patterns. Participants were instructed to attend to either coloured or greyscale patterns while ignoring the others. We found that attending to either coloured or greyscale patterns produced robust increases in ssVEP amplitudes both compared to ignored stimuli and to baseline. There was however no evidence of suppressed responses to ignored patterns. These findings demonstrate that attentional selection based on the presence or absence of chromatic information prompts selectively enhanced visuocortical processing but this selective amplification is not accompanied by suppression of unattended stimuli. Findings are consistent with theoretical notions that predict strong competition between specific exemplars within a given feature dimension, such as red or green, but weak competition between broadly defined stimulus categories, such as chromatic versus non-chromatic.

Suppression of Unattended Features Is Independent of Task Relevance.

Feature-based attention serves the separation of relevant from irrelevant features. While global amplification of attended features is coherently described as a key mechanism for feature-based attention, nature and constituting factors of neural suppressive interactions are far less clear. One aspect of global amplification is its flexible modulation by the task relevance of the to-be-attended stimulus. We examined whether suppression is similarly modulated by their respective task relevance or is mandatory for all unattended features. For this purpose, participants saw a display of randomly moving dots with 3 distinct colors and were asked to report brief events of coherent motion for a cued color. Of the 2 unattended colored clouds, one contained distracting motion events while the other was irrelevant and without such motion events throughout the experiment. We used electroencephalography-derived steady-state visual-evoked potentials to investigate early visual processing of the attended, unattended, and irrelevant color under sustained feature-based attention. The analysis revealed a biphasic process with an early amplification of the to-be-attended color followed by suppression of the to-be-ignored color relative to a pre-cue baseline. Importantly, the neural dynamics for the unattended and always irrelevant color were comparable. Suppression is thus a mandatory mechanism affecting all unattended stimuli irrespective of their task relevance.

Electrophysiological evidence for target facilitation without distractor suppression in two-stimulus search displays.

This study used electrophysiological measures to investigate how attention is deployed to target and distractor stimuli during visual search using search displays with a small set-size. Participants viewed randomized sequences of two-item displays that consisted of either a target and a distractor (differing in color) or a pair of task-irrelevant filler stimuli having a third color, all presented in an ongoing stream of flickering gray circles. The allocation of attention was assessed by concurrent recordings of steady-state visual evoked potentials (SSVEPs) elicited by the flickering displays and perturbations of the endogenous alpha rhythm following each type of display. The aim was to test a central prediction of the signal suppression hypothesis, namely that the processing of distractors will be proactively suppressed below the level of filler stimuli. Amplitude modulations of both the SSVEP and the lateralized alpha rhythm provided converging evidence against early proactive suppression of highly salient distractors. Instead, these electrophysiological measures were consistent with the view that in this type of two-stimulus search task there is an initial capture of attention by all color-change stimuli (targets, distractors, and fillers) followed by a further focusing of attention upon the target, with no evidence for suppression of the distractor.

Dynamics of attentional allocation to targets and distractors during visual search.

There is much debate about the neural mechanisms that achieve suppression of salient distracting stimuli during visual search. The proactive suppression hypothesis asserts that if exposed to the same distractors repeatedly, these stimuli are actively inhibited before attention can be shifted to them. A contrasting proposal holds that attention is initially captured by salient distractors but is subsequently withdrawn. By concurrently measuring stimulus-driven and intrinsic brain potentials in 36 healthy human participants, we obtained converging evidence against early proactive suppression of distracting input. Salient distractors triggered negative event-related potentials (N1pc/N2pc), enhanced the steady-state visual evoked potential (SSVEP) relative to non-salient (filler) stimuli, and suppressed contralateral relative to ipsilateral alpha-band amplitudes-three electrophysiological measure associated with the allocation of attention-even though these distractors did not interfere with behavioral responses to the search targets. Furthermore, these measures indicated that both stimulus-driven and goal-driven allocations of attention occurred in conjunction with one another, with the goal-driven effect enhancing and prolonging the stimulus-driven effect. These results provide a new perspective on the traditional dichotomy between bottom-up and top-down attentional allocation. Control experiments revealed that continuous marking of the locations at which the search display items were presented resulted in a dramatic and unexpected conversion of the target-elicited N2pc into a shorter-latency N1pc in association with faster reaction times to the targets.

One decade of quantum optimal control in the chopped random basis.

The chopped random basis (CRAB) ansatz for quantum optimal control has been proven to be a versatile tool to enable quantum technology applications such as quantum computing, quantum simulation, quantum sensing, and quantum communication. Its capability to encompass experimental constraints-while maintaining an access to the usually trap-free control landscape-and to switch from open-loop to closed-loop optimization (including with remote access-or RedCRAB) is contributing to the development of quantum technology on many different physical platforms. In this review article we present the development, the theoretical basis and the toolbox for this optimization algorithm, as well as an overview of the broad range of different theoretical and experimental applications that exploit this powerful technique.

Shifting Attention in Feature Space: Fast Facilitation of the To-Be-Attended Feature Is Followed by Slow Inhibition of the To-Be-Ignored Feature.

In an explorative study, we investigated the time course of attentional selection shifts in feature-based attention in early visual cortex by means of steady-state visual evoked potentials (SSVEPs). To this end, we presented four flickering random dot kinematograms with red/blue, horizontal/vertical bars, respectively. Given the oscillatory nature of SSVEPs, we were able to investigate neural temporal dynamics of facilitation and inhibition/suppression when participants shifted attention either within (i.e., color to color) or between feature dimensions (i.e., color to orientation). Extending a previous study of our laboratory [Müller, M. M., Trautmann, M., & Keitel, C. Early visual cortex dynamics during top-down modulated shifts of feature-selective attention. Journal of Cognitive Neuroscience, 28, 643-655, 2016] to a full factorial design, we replicated a critical finding of our previous study: Facilitation of color was quickest, regardless of the origin of the shift (from color or orientation). Furthermore, facilitation of the newly to-be-attended and inhibition/suppression of the then to-be-ignored feature is not a time-invariant process that occurs instantaneously, but a biphasic one with longer time delays between the two processes. Interestingly, inhibition/suppression of the to-be-ignored feature after the shifting cue had a much longer latency with between- compared to within-dimensional shifts (by about 130-150 msec). The exploratory nature of our study is reasoned by two limiting factors: (a) Identical to our precursor study, we found no attentional SSVEP amplitude time course modulation for orientation, and (b) the signal-to-noise ratio for single trials was too poor to allow for reliable statistical testing of the latencies that were obtained with running t tests of averaged data.

Affective Bias without Hemispheric Competition: Evidence for Independent Processing Resources in Each Cortical Hemisphere.

We assessed the extent of neural competition for attentional processing resources in early visual cortex between foveally presented task stimuli and peripheral emotional distracter images. Task-relevant and distracting stimuli were shown in rapid serial visual presentation (RSVP) streams to elicit the steady-state visual evoked potential, which serves as an electrophysiological marker of attentional resource allocation in early visual cortex. A task-related RSVP stream of symbolic letters was presented centrally at 15 Hz while distracting RSVP streams were displayed at 4 or 6 Hz in the left and right visual hemifields. These image streams always had neutral content in one visual field and would unpredictably switch from neutral to unpleasant content in the opposite visual field. We found that the steady-state visual evoked potential amplitude was consistently modulated as a function of change in emotional valence in peripheral RSVPs, indicating sensory gain in response to distracting affective content. Importantly, the facilitated processing for emotional content shown in one visual hemifield was not paralleled by any perceptual costs in response to the task-related processing in the center or the neutral image stream in the other visual hemifield. Together, our data provide further evidence for sustained sensory facilitation in favor of emotional distracters. Furthermore, these results are in line with previous reports of a "different hemifield advantage" with low-level visual stimuli and are suggestive of independent processing resources in each cortical hemisphere that operate beyond low-level visual cues, that is, with complex images that impact early stages of visual processing via reentrant feedback loops from higher order processing areas.

Dissociable neural correlates of stimulation intensity and detection in somatosensation.

Somatosensory stimulation intensity and behavioral detection are positively related, and both correlate with neural responses. However, it is still controversial as to what extent stimulus intensity and early somatosensory evoked potentials (SEP) predict detection and how these parameters interact with pre-stimulus brain oscillatory states, which also influence sensory processing. Here we investigated how early SEP components encode stimulation intensity, how pre-stimulus alpha- and beta-band amplitudes interact with SEPs, and which neural markers predict stimulus detection. To this end, we randomly presented electrical finger nerve stimulation with various intensities distributed along the individual psychometric response function (including catch trials) while recording the EEG. Participants reported stimulus presence on a trial-by-trial basis (one-alternative-forced-choice). For the lowest (imperceptible) intensities, participants showed zero (behavioral) sensitivity despite measurable early cortical processing reflected by the P50 component. The P50 amplitude scaled with increasing stimulation intensities but was not predictive of stimulus detection. Instead, detection was associated with the later negative N150 component, as well as with pre-stimulus lowered somatosensory alpha- and increased frontal beta-band amplitudes. Our results give evidence for a serial representation of stimulus intensity and detection, as reflected by the P50 and N150 amplitude, respectively. Furthermore, stimulus detection seems to depend on the current brain state, rendering upcoming stimulation being reportable or not.

Reduction of somatosensory functional connectivity by transcranial alternating current stimulation at endogenous mu-frequency.

Alpha, the most prominent human brain rhythm, might reflect a mechanism of functional inhibition for gating neural processing. This concept has been derived predominantly from local measures of inhibition, while large-scale network mechanisms to guide information flow are largely unknown. Here, we investigated functional connectivity changes on a whole-brain level by concurrent transcranial alternating current stimulation (tACS) and resting-state functional MRI in humans. We specifically focused on somatosensory alpha-band oscillations by adjusting the tACS frequency to each individual´s somatosensory (mu-) alpha peak frequency (mu-tACS). Potential differences of Eigenvector Centrality of primary somatosensory cortex (S1) as well as on a whole brain level between mu-tACS and sham were analyzed. Our results demonstrate that mu-tACS induces a locally-specific decrease in whole-brain functional connectivity of left S1. An additional exploratory analysis revealed that this effect primarily depends on a decrease in functional connectivity between S1 and a network of regions that are crucially involved in somatosensory processing. Furthermore, the decrease in functional centrality was specific to mu-tACS and was not observed when tACS was applied in the gamma-range in an independent study. Our findings provide evidence that modulated somatosensory (mu-) alpha-activity may affect whole-brain network level activity by decoupling primary sensory areas from other hubs involved in sensory processing.

Attentional Facilitation of Constituent Features of an Object Does Not Spread Automatically along Object-defining Cortical Boundaries.

The integrated object account predicts that attention is spread across all features that constitute one object, regardless of their task relevance. We challenge that prediction with a novel stimulation technique that allows for simultaneous electrophysiological measurements of the allocation of attention to two distinct features within one object. A rotating square that flickers in different colors evoked two distinct steady-state visual evoked potentials (SSVEPs) for rotation and color, respectively. If the integrated object account were true, we would expect identical SSVEP amplitudes regardless of what feature participants attended. We found greater SSVEP amplitudes for the to-be-attended feature compared with the to-be-ignored feature. SSVEP amplitudes averaged across both features were significantly reduced when participants attended to both features, which was mirrored in behavioral costs, implying competitive interactions or a division of attentional resources. Surprisingly, this reduction in amplitude was mainly driven by the SSVEP amplitude elicited by color changes. In conclusion, our results challenge the integrated object account and highlight the extent to which color is "special" within feature space.

Time matters: Feature-specific prioritization follows feature integration in visual object processing.

Objects represent a fundamental selection unit of visual attention. However, at odds with the integrated competition account, our recent study demonstrated that attentional facilitation of constituent features does not spread automatically within an object, but instead depends on the specific task relevance of each feature. Here, we employed a novel experimental design, allowing simultaneous electrophysiological measurements of the allocation of attention to two distinct features (rotation and color) within one object (a square) during both trial-wise and block-wise cued shifts of attention. This was possible through the presentation of a square that evokes two distinct steady-state visual evoked potentials (SSVEPs) for its rotation and its color changes, respectively. Given the continuous oscillatory nature of SSVEPs, we were able to investigate the time course of neural activity in the early visual cortex of the human brain when subjects attended to one of the two features, compared to when the whole object was attended. This approach enabled us to uncover feature-based mechanisms of attention within one object, as well as their interaction with object-based mechanisms. Both behavioral and electrophysiological results indicate a biphasic process composed of an early transient integration of the constituent object features, followed by sustained mechanisms of feature selection with amplification of the to-be-attended feature, followed temporally by suppression of the to-be-ignored feature.

Rapid sensory gain with emotional distracters precedes attentional deployment from a foreground task.

The steady-state visual evoked potential (SSVEP), an electrophysiological marker of attentional resource allocation, has recently been demonstrated to serve as a neural signature of emotional content extraction from a rapid serial visual presentation (RSVP). SSVEP amplitude was reduced for streams of emotional relative to neutral scenes passively viewed at 6 Hz (~167 ms per image), but it was enhanced for emotional relative to neutral scenes when viewed as 4 Hz RSVP (250 ms per image). Here, we investigated whether these seemingly contradictory observations may be related to different dynamics in the allocation of attentional resources as a consequence of stimulation frequency. To this end, we advanced our distraction paradigm by presenting a visual foreground task consisting of randomly moving squares flickering at 15 Hz superimposed on task-irrelevant RSVP streams shown at 6 or 4 Hz, which could unpredictably switch from neutral to unpleasant content during the trial or remained neutral. Critically, our findings demonstrate that affective distractors captured attentional resources more strongly than their neutral counterparts, irrespective of whether they were presented at 6 or 4 Hz rate. Moreover, the emotion-dependent attentional deployment from the foreground task was temporally preceded by sustained sensory facilitation in response to emotional background images. Together, present findings provide evidence for rapid sustained visual facilitation but a rather slow attentional bias in favor of emotional distractors in early visual areas.

Alpha-Band Brain Oscillations Shape the Processing of Perceptible as well as Imperceptible Somatosensory Stimuli during Selective Attention.

Attention filters and weights sensory information according to behavioral demands. Stimulus-related neural responses are increased for the attended stimulus. Does alpha-band activity mediate this effect and is it restricted to conscious sensory events (suprathreshold), or does it also extend to unconscious stimuli (subthreshold)? To address these questions, we recorded EEG in healthy male and female volunteers undergoing subthreshold and suprathreshold somatosensory electrical stimulation to the left or right index finger. The task was to detect stimulation at the randomly alternated cued index finger. Under attention, amplitudes of somatosensory evoked potentials increased 50-60 ms after stimulation (P1) for both suprathreshold and subthreshold events. Prestimulus amplitude of peri-Rolandic alpha, that is mu, showed an inverse relationship to P1 amplitude during attention compared to when the finger was unattended. Interestingly, intermediate and high amplitudes of mu rhythm were associated with the highest P1 amplitudes during attention and smallest P1 during lack of attention, that is, these levels of alpha rhythm seemed to optimally support the behavioral goal ("detect" stimuli at the cued finger while ignoring the other finger). Our results show that attention enhances neural processing for both suprathreshold and subthreshold stimuli and they highlight a rather complex interaction between attention, Rolandic alpha activity, and their effects on stimulus processing.SIGNIFICANCE STATEMENT Attention is crucial in prioritizing processing of relevant perceptible (suprathreshold) stimuli: it filters and weights sensory input. The present study investigates the controversially discussed question whether this attention effect extends to imperceptible (subthreshold) stimuli as well. We found noninvasive EEG signatures for attentional modulation of neural events following perceptible and imperceptible somatosensory stimulation in human participants. Specifically, stimulus processing for both kinds of stimulation, subthreshold and suprathreshold, is enhanced by attention. Interestingly, Rolandic alpha rhythm strength and its influence on stimulus processing are strikingly altered by attention most likely to optimally achieve the behavioral goal.

It takes two to tango: Suppression of task-irrelevant features requires (spatial) competition.

In a recent electrophysiological study, we reported on global facilitation but local suppression of color stimuli in feature-based attention in human early visual cortex. Subjects attended to one of two centrally located superimposed red/blue random dot kinematograms (RDKs). Task-irrelevant single RDKs in the same colors were presented in the left and right periphery, respectively. Suppression of the to-be-ignored color was only present in the centrally located RDK but not in the one with the same color in the periphery. This result was at odds with the idea of active suppression of task-irrelevant features across the entire visual field. In the present study, we introduced competition in the periphery by superimposing the RDKs at the task-irrelevant location as well. With such competition, we found suppression of the task-irrelevant color in the centrally and peripherally located RDKs. Results clearly demonstrate that suppression of task-irrelevant features at task-irrelevant locations requires (spatial) competitive interactions and is not an inherent neural mechanism in feature-based attention as was found for global facilitation.

Neural mechanisms of divided feature-selective attention to colour.

Feature-based attentional selection of colour is challenging to investigate due to the multidimensional nature of colour-space. When attending concurrently to features from different feature dimensions (e.g. red and horizontal), the attentional selections of the separate dimensions are largely independent. Therefore, if colour constitutes multiple independent feature dimensions for attentional purposes, concurrently attending to two colours should be effective and independent of the specific configuration of target and distractor colours. Here, observers attended concurrently to two out of four fully overlapping random dot kinematograms of different colours, and the allocation of attention to each colour was assessed separately by recordings of steady-state visual evoked potentials. The magnitude of attention effects depended on colour proximity and was well described by a simple model which suggested that colour space is rescaled in an adaptive manner to achieve attentional selection. In conclusion, different spatially overlaid colours can be attended concurrently with an efficiency that is determined by their configuration in colour space, supporting the idea that (at least in terms of hue) colour acts as a single dimension for attentional purposes.

Global Enhancement but Local Suppression in Feature-based Attention.

A key property of feature-based attention is global facilitation of the attended feature throughout the visual field. Previously, we presented superimposed red and blue randomly moving dot kinematograms (RDKs) flickering at a different frequency each to elicit frequency-specific steady-state visual evoked potentials (SSVEPs) that allowed us to analyze neural dynamics in early visual cortex when participants shifted attention to one of the two colors. Results showed amplification of the attended and suppression of the unattended color as measured by SSVEP amplitudes. Here, we tested whether the suppression of the unattended color also operates globally. To this end, we presented superimposed flickering red and blue RDKs in the center of a screen and a red and blue RDK in the left and right periphery, respectively, also flickering at different frequencies. Participants shifted attention to one color of the superimposed RDKs in the center to discriminate coherent motion events in the attended from the unattended color RDK, whereas the peripheral RDKs were task irrelevant. SSVEP amplitudes elicited by the centrally presented RDKs confirmed the previous findings of amplification and suppression. For peripherally located RDKs, we found the expected SSVEP amplitude increase, relative to precue baseline when color matched the one of the centrally attended RDK. We found no reduction in SSVEP amplitude relative to precue baseline, when the peripheral color matched the unattended one of the central RDK, indicating that, while facilitation in feature-based attention operates globally, suppression seems to be linked to the location of focused attention.