Distinct patterns of spatial attentional modulation of steady-state visual evoked magnetic fields (SSVEFs) in subdivisions of the human early visual cortex.

In recent years, steady-state visual evoked potentials (SSVEPs) became an increasingly valuable tool to investigate neural dynamics of competitive attentional interactions and brain-computer interfaces. This is due to their good signal-to-noise ratio, allowing for single-trial analysis, and their ongoing oscillating nature that enables to analyze temporal dynamics of facilitation and suppression. Given the popularity of SSVEPs, it is surprising that only a few studies looked at the cortical sources of these responses. This is in particular the case when searching for studies that assessed the cortical sources of attentional SSVEP amplitude modulations. To address this issue, we used a typical spatial attention task and recorded neuromagnetic fields (MEG) while presenting frequency-tagged stimuli in the left and right visual fields, respectively. Importantly, we controlled for attentional deployment in a baseline period before the shifting cue. Subjects either attended to a central fixation cross or to two peripheral stimuli simultaneously. Results clearly showed that signal sources and attention effects were restricted to the early visual cortex: V1, V2, hMT+, precuneus, occipital-parietal, and inferior-temporal cortex. When subjects attended to central fixation first, shifting attention to one of the peripheral stimuli resulted in a significant activation increase for the to-be-attended stimulus with no activation decrease for the to-be-ignored stimulus in hMT+ and inferio-temporal cortex, but significant SSVEF decreases from V1 to occipito-parietal cortex. When attention was first deployed to both rings, shifting attention away from one ring basically resulted in a significant activation decrease in all areas for the then-to-be-ignored stimulus.

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.

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.

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.

Higher amplitudes in steady-state visual evoked potentials driven by square-wave versus sine-wave contrast modulation - A dual-laboratory study.

Steady-state visual evoked potentials (ssVEPs) are an established tool for assessing visuocortical responses in visual perception and attention. They have the same temporal frequency characteristics as a periodically modulated stimulus (e.g., in contrast or luminance) that drives them. It has been hypothesized that the amplitude of a given ssVEP may depend on the shape of the stimulus modulation function, but the size and robustness of these effects is not well established. The current study systematically compared the effect of the two most common functions in the ssVEP literature, square-wave and sine-wave functions. Across two laboratories, we presented mid-complex color patterns to 30 participants with square-wave or sine-wave contrast modulation and at different driving frequencies (6 Hz, 8.57 Hz, 15 Hz). When ssVEPs were analyzed independently for the samples, with each laboratory's standard processing pipeline, ssVEP amplitudes in both samples decreased at higher driving frequencies and square-wave modulation evoked higher amplitudes at lower frequencies (i.e., 6 Hz, 8.57 Hz) compared to sine-wave modulation. These effects were replicated when samples were aggregated and analyzed with the same processing pipeline. In addition, when using signal-to-noise ratios as outcome measures, this joint analysis indicated a somewhat weaker effect of increased ssVEP amplitudes to square-wave modulation at 15 Hz. The present study suggests that square-wave modulation should be used in ssVEP research when the goal is to maximize signal amplitude or signal-to-noise ratio. Given effects of modulation function across laboratories, and data processing pipelines, the findings appear robust to differences in data collection and analysis.

Global attentional selection of visual features is not associated with selective modulation of posterior alpha-band activity.

Attending to a single feature, such as color or motion, leads to global modulation of neural processing associated with the representation of the attended features. Alpha-band modulations are hypothesized to be a marker (and even a mechanism) of the modulation of neural processing. By adopting a previously used attentional shifting paradigm, we examined whether alpha-band dynamics are linked to sustained Feature-Based-Attentional (FBA) selection. For this purpose, we presented task-irrelevant flickering random dot kinematograms (RDKs) in the periphery that either did or did not share the to-be-attended color of centrally presented task-relevant RDKs and should thus be subject to global FBA selection. Steady-state visual evoked potentials (SSVEPs) and alpha-band activity associated with these task-irrelevant RDKs were analyzed to quantify FBA modulation. Overall, the SSVEP results replicated previous findings: relative to a pre-cue baseline, SSVEP amplitudes for peripheral RDKs were significantly enhanced when these RDKs shared the to-be-attended color of the central RDKs and were not modulated when they shared the centrally to-be-ignored color. Nevertheless, there were no differences in alpha-band amplitude modulations between signals recorded contralateral to the RDKs sharing the centrally attended color and RDKs sharing the centrally ignored color. Hence, alpha-band modulations seem not to index the sustained global selection of attended over unattended feature values within the same feature dimension.

Attentional capture is modulated by stimulus saliency in visual search as evidenced by event-related potentials and alpha oscillations.

This study used a typical four-item search display to investigate top-down control over attentional capture in an additional singleton paradigm. By manipulating target and distractor color and shape, stimulus saliency relative to the remaining items was systematically varied. One group of participants discriminated the side of a dot within a salient orange target (ST group) presented with green circles (fillers) and a green diamond distractor. A second group discriminated the side of the dot within a green diamond target presented with green circle fillers and a salient orange square distractor (SD group). Results showed faster reaction times and a shorter latency of the N2pc component in the event-related potential (ERP) to the more salient targets in the ST group. Both salient and less salient distractors elicited Pd components of equal amplitude. Behaviorally, no task interference was observed with the less salient distractor, indicating the prevention of attentional capture. However, reaction times were slower in the presence of the salient distractor, which conflicts with the hypothesis that the Pd reflects proactive distractor suppression. Contrary to recent proposals that elicitation of the Pd requires competitive interactions with a target, we found a greater Pd amplitude when the distractor was presented alone. Alpha-band amplitudes decreased during target processing (event-related desynchronization), but no significant amplitude enhancement was observed at electrodes contralateral to distractors regardless of their saliency. The results demonstrate independent neural mechanisms for target and distractor processing and support the view that top-down guidance of attention can be offset (counteracted) by relative stimulus saliency.

Information theoretical limits for quantum optimal control solutions: error scaling of noisy control channels.

Accurate manipulations of an open quantum system require a deep knowledge of its controllability properties and the information content of the implemented control fields. By using tools of information and quantum optimal control theory, we provide analytical bounds (information-time bounds) to characterize our capability to control the system when subject to arbitrary sources of noise. Moreover, since the presence of an external noise field induces open quantum system dynamics, we also show that the results provided by the information-time bounds are in very good agreement with the Kofman-Kurizki universal formula describing decoherence processes. Finally, we numerically test the scaling of the control accuracy as a function of the noise parameters, by means of the dressed chopped random basis (dCRAB) algorithm for quantum optimal control.

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.

An integrative model for neural dynamics of feature selection in visual object processing: An explorative study.

Objects and features represent fundamental selection units of visual attention. The associated mechanisms, however, had been mostly explored in isolation. Not least for this reason, apparently contradictory theories persist about feature selection in visual object processing. Our previous study demonstrated the time-dependent nature of this process. In the present explorative study, we used electroencephalographic recordings to examine feature-specific selection during the time course of object processing. For this, we analyzed the feature-specific selection negativities (SNs), i.e., the SNs when human subjects attended to or ignored one (color or shape) relative to attending to all (color and shape) constituent features of an object (a square). We found that the feature-specific SN amplitude of the task-relevant feature was significantly decreased in a time bin from 500 to 850 ms after trial onset in comparison to an earlier (100-450 ms) and a later time bin (900-1250 ms). Whereas, the feature-specific SN amplitude of the task-irrelevant feature was firstly positive, indicating no specific selection of this feature, and secondly, it marginally increased from the first to the second time bin. This leads to the proposed integrative model, which indicates that sustained feature-specific prioritization only emerges over time, this process consisting of three phases: after initial selection of the relevant feature and inhibition of the irrelevant feature (first time bin), further inhibition of the irrelevant feature is accompanied by co-inhibition of the relevant feature, causing prioritization to decline (second time bin), before the relevant feature eventually regains priority by being facilitated again (third time bin).

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.

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.

Emotional content overrides spatial attention.

Spatial attention is our capacity to attend to or ignore particular regions of our spatial environment. However, some classes of stimuli may be able to override our efforts to ignore them. Here we assessed the relationship between involuntary attentional capture with emotional images and spatial attention at early stages of perceptual processing. Multiple scenes of unpleasant and neutral content were displayed in rapid serial visual presentation (RSVP) streams that elicited the steady-state visual evoked potential (SSVEP), a neural marker of selective attention at early visual areas. In a spatial cueing task, participants were cued to covertly attend to RSVP streams presented at 4 and 6 Hz presentation rates in the left and right visual hemifields. The task was to detect square targets occasionally displayed within the image streams, responding only to those appearing on the cued side. The RSVP streams were always neutral pictures in one visual hemifield but would unpredictably switch from neutral to aversive content in the other visual hemifield. We found that SSVEP amplitude was consistently modulated by a change in emotional valence of image streams, regardless of whether the change in content occurred in the attended or unattended spatial location, reflecting an automatic sensory amplification for affective stimuli. The present data provide further evidence in support that emotional images can attract visual processing resources independently of spatial attention allocation, and are consistent with sustained sensory facilitation of early visual areas through re-entrant feedback projections from higher-order cortical areas involved in the extraction of affective information.

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.

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.

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.

Alignment of Continuous Auditory and Visual Distractor Stimuli Is Leading to an Increased Performance.

Information across different senses can affect our behavior in both positive and negative ways. Stimuli aligned with a target stimulus can lead to improved behavioral performances, while competing, transient stimuli often negatively affect our task performance. But what about subtle changes in task-irrelevant multisensory stimuli? Within this experiment we tested the effect of the alignment of subtle auditory and visual distractor stimuli on the performance of detection and discrimination tasks respectively. Participants performed either a detection or a discrimination task on a centrally presented Gabor patch, while being simultaneously subjected to a random dot kinematogram, which alternated its color from green to red with a frequency of 7.5 Hz and a continuous tone, which was either a frequency modulated pure tone for the audiovisual congruent and incongruent conditions or white noise for the visual control condition. While the modulation frequency of the pure tone initially differed from the modulation frequency of the random dot kinematogram, the modulation frequencies of both stimuli could align after a variable delay, and we measured accuracy and reaction times around the possible alignment time. We found increases in accuracy for the audiovisual congruent condition suggesting subtle alignments of multisensory background stimuli can increase performance on the current task.