The dynamics of statistical learning in visual search and its interaction with salience processing: An EEG study.

Visual attention can be guided by statistical regularities in the environment, that people implicitly learn from past experiences (statistical learning, SL). Moreover, a perceptually salient element can automatically capture attention, gaining processing priority through a bottom-up attentional control mechanism. The aim of our study was to investigate the dynamics of SL and if it shapes attentional target selection additively with salience processing, or whether these mechanisms interact, e.g. one gates the other. In a visual search task, we therefore manipulated target frequency (high vs. low) across locations while, in some trials, the target was salient in terms of colour. Additionally, halfway through the experiment, the high-frequency location changed to the opposite hemifield. EEG activity was simultaneously recorded, with a specific interest in two markers related to target selection and post-selection processing, respectively: N2pc and SPCN. Our results revealed that both SL and saliency significantly enhanced behavioural performance, but also interacted with each other, with an attenuated saliency effect at the high-frequency target location, and a smaller SL effect for salient targets. Concerning processing dynamics, the benefit of salience processing was more evident during the early stage of target selection and processing, as indexed by a larger N2pc and early-SPCN, whereas SL modulated the underlying neural activity particularly later on, as revealed by larger late-SPCN. Furthermore, we showed that SL was rapidly acquired and adjusted when the spatial imbalance changed. Overall, our findings suggest that SL is flexible to changes and, combined with salience processing, jointly contributes to establishing attentional priority.

Separate and overlapping mechanisms of statistical regularities and salience processing in the occipital cortex and dorsal attention network.

Attention selects behaviorally relevant inputs for in-depth processing. Beside the role of traditional signals related to goal-directed and stimulus-driven control, a debate exists regarding the mechanisms governing the effect of statistical regularities on attentional selection, and how these are integrated with other control signals. Using a visuo-spatial search task under fMRI, we tested the joint effects of statistical regularities and stimulus-driven salience. We found that both types of signals modulated occipital activity in a spatially specific manner. Salience acted primarily by reducing the attention bias towards the target location when associated with irrelevant distractors, while statistical regularities reduced this attention bias when the target was presented at a low probability location, particularly at the lower levels of the visual hierarchy. In addition, we found that both statistical regularities and salience activated the dorsal frontoparietal network. Additional exploratory analyses of functional connectivity revealed that only statistical regularities modulated the inter-regional coupling between the posterior parietal cortex and the occipital cortex. These results show that statistical regularities and salience signals are both spatially represented at the occipital level, but that their integration into attentional processing priorities relies on dissociable brain mechanisms.

Statistical learning of target and distractor spatial probability shape a common attentional priority computation.

Converging evidence recently put forward the notion that dedicated neurocognitive mechanisms do exist for the suppression of salient, but irrelevant distractors. Along this line, it is plausible to hypothesize that, in appropriate contexts, experience-dependent forms of attentional learning might selectively induce plastic changes within this dedicated circuitry, thus allowing an independent shaping of priorities at the service of attentional filtering. Conversely, previous work suggested that statistical learning (SL) of both target and distractor spatial probability distributions converge in adjusting only the overall attentional priority of locations: in fact, in the presence of an independent manipulation, either related to the target or to the distractor only, SL induces indirect effects (e.g., changes in filtering efficiency due to an uneven distribution of targets), suggesting that SL-induced plastic changes affect a shared neural substrate. Here we tested whether, when (conflicting) target- and distractor-related manipulations are concurrently applied to the very same locations, dedicated mechanisms might support the selective encoding of spatial priority in relation to the specific attentional operation involved. In three related experiments, human healthy participants discriminated the direction of a target arrow, while ignoring a salient distractor, if present; both target and distractor spatial probability distributions were concurrently manipulated in relation to each single location. Critically, the selection bias produced by the target-related SL was marginally reduced by an adverse distractor contingency, and the suppression bias generated by the distractor-related SL was erased, or even reversed, by an adverse target contingency. Our results suggest that even conflicting target- and distractor-related SL manipulations result in the adjustment of a unique spatial priority computation, likely because the process directly relies on direct plastic alterations of shared spatial priority map(s).

Integrated effects of top-down attention and statistical learning during visual search: An EEG study.

The present study aims to investigate how the competition between visual elements is solved by top-down and/or statistical learning (SL) attentional control (AC) mechanisms when active together. We hypothesized that the "winner" element that will undergo further processing is selected either by one AC mechanism that prevails over the other, or by the joint activity of both mechanisms. To test these hypotheses, we conducted a visual search experiment that combined an endogenous cueing protocol (valid vs. neutral cue) and an imbalance of target frequency distribution across locations (high- vs. low-frequency location). The unique and combined effects of top-down control and SL mechanisms were measured on behaviour and amplitudes of three evoked-response potential (ERP) components (i.e., N2pc, P1, CNV) related to attentional processing. Our behavioural results showed better performance for validly cued targets and for targets in the high-frequency location. The two factors were found to interact, so that SL effects emerged only in the absence of top-down guidance. Whereas the CNV and P1 only displayed a main effect of cueing, for the N2pc we observed an interaction between cueing and SL, revealing a cueing effect for targets in the low-frequency condition, but not in the high-frequency condition. Thus, our data support the view that top-down control and SL work in a conjoint, integrated manner during target selection. In particular, SL mechanisms are reduced or even absent when a fully reliable top-down guidance of attention is at play.

Effects of top-down and bottom-up attention on post-selection posterior contralateral negativity.

We examined the effect of combined top-down and bottom-up attentional control sources in easy and difficult visual search tasks. Applying a new analysis on previously acquired data, we focused on the sustained posterior contralateral negativity (SPCN) and the response-locked posterior contralateral negativity (RLpcN), to better understand processes following target selection. We used the signed-area approach to measure the negative area, where the signal was either locked to the target or the response onsets. We further split the RLpcN into an early and a late segment to capture the dynamics of selection and post-selection processes. In Experiment 1, participants reported the orientation of a uniquely tilted target. In Experiment 2, participants reported the position of a small gap within the uniquely tilted target. In both experiments, endogenous cues manipulated top-down attention (valid vs. neutral), and salient color singletons (either the target or a distractor) manipulated bottom-up attention. We hypothesized that the SPCN and the later segment of the RLpcN would be modulated by task difficulty and target salience, as they are associated with post-selection processes, such as response selection and working memory. The early segment of the RLpcN was hypothesized to be modulated by the cueing manipulation and presence of a salient distractor, as they affect target selection. An effect of distractor presence was observed on the early segment of the RLpcN, and our results further supported the hypotheses regarding the SPCN and the later segment of the RLpcN, providing novel insights into post-selection processes in visual search.

Dynamic causal interactions between occipital and parietal cortex explain how endogenous spatial attention and stimulus-driven salience jointly shape the distribution of processing priorities in 2D visual space.

Visuo-spatial attention prioritizes the processing of relevant inputs via different types of signals, including current goals and stimulus salience. Complex mixtures of these signals engage in everyday life situations, but little is known about how these signals jointly modulate distributed patterns of activity across the occipital regions that represent visual space. Here, we measured spatio-topic, quadrant-specific occipital activity during the processing of visual displays containing both task-relevant targets and salient color-singletons. We computed spatial bias vectors indexing the effect of attention in 2D space, as coded by distributed activity in the occipital cortex. We found that goal-directed spatial attention biased activity towards the target and that salience further modulated this endogenous effect: salient distractors decreased the spatial bias, while salient targets increased it. Analyses of effective connectivity revealed that the processing of salient distractors relied on the modulation of the bidirectional connectivity between the occipital and the posterior parietal cortex, as well as the modulation of the lateral interactions within the occipital cortex. These findings demonstrate that goal-directed attention and salience jointly contribute to shaping processing priorities in the occipital cortex and highlight that multiple functional paths determine how spatial information about these signals is distributed across occipital regions.

An EEG study of the combined effects of top-down and bottom-up attentional selection under varying task difficulty.

We examined the effect of combined top-down and bottom-up attentional control sources, using known attention-related EEG components that are thought to reflect target selection (N2pc) and distractor suppression (PD ). We used endogenous cues (valid vs. neutral) for top-down attentional control, and salience in the form of color singletons (either the target or a distractor) for bottom-up attentional control in visual search. Crucially, in two experiments, the task was of increasing difficulty, reporting the orientation of a tilted target (Experiment 1), or the position of a small gap within the target among tilted non-targets (Experiment 2). Our results showed strong cueing effects on RT and accuracy in both experiments, demonstrating a general facilitation of responses to validly cued targets. Whereas the processing of salient targets was not improved compared with non-salient targets, the presence of a salient distractor consistently worsened performance. The N2pc and PD were only observed in trials where targets were preceded by neutral cues in Experiment 1, and for validly cued targets and salient neutrally cued targets in Experiment 2. A cueing effect was found on the PD in Experiment 1, showing an amplitude reduction in trials where the target was validly cued. These results support the idea that bottom-up attentional allocation occurs only when top-down allocation of attention is absent or inefficient. Therefore, these results indicate that attentional selection and suppression during visual search are both influenced by top-down cueing and give support to theories that focus on the interaction between the two types of attention.

How feature context alters attentional template switching.

In real-world tasks visual attention is rarely aimed at a single object. Humans rather "forage" the visual scene for information, dynamically switching attentional templates. Several visual search studies have found that observers often use suboptimal attentional control strategies, possibly to avoid effort. Here, we investigated with a foraging paradigm if observers' reluctance to switch between attentional templates increases with template specificity. To that end, we manipulated the feature context of displays in which participants "foraged" moving stimuli on a tablet-PC. Experiment 1 (N = 35) revealed a decline in switching tendency and foraging efficiency with increasing feature-space distance between target alternatives. Experiment 2 (N = 36) found even lower flexibility with distractor color close to target colors and strongest impairments with distractor color in between target colors. Our results demonstrate that visual information sampling is most flexible when broad (instead of very specific) templates and relational search strategies are possible (e.g., attending to "redder" objects), with implications for both attention research and applications, especially in visual-foraging-like tasks, such as baggage screening or medical image assessment. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Predictive brain: Addressing the level of representation by reviewing perceptual hysteresis.

In recent years, the idea that the prediction of sensory input is one of the major computational goals of the nervous system led to the development of several large-scale theories of brain functioning, such as different versions of the Bayesian approach to brain functions, predictive coding theories of cognition and the Free-energy principle. During the years, various empirical phenomena have been re-interpreted within such frames, and have been considered as consequences of predictive processing. Here we focus on perceptual hysteresis, or serial dependence, as an exemplary case. We unravel a potential gap in the predictive frameworks and raise the idea that alternative explanations of this effect can solve this issue, as they address the type of cognitive and neural representations involved.

The role of the vestibular system in value attribution to positive and negative reinforcers.

Somatic inputs originating from bioregulatory processes can guide cognition and behavior. One such bodily signal, mostly overlooked so far, is represented by visuo-vestibular coupling and its alteration, which in extreme cases may result in motion sickness. We argued that the inherently perturbed interoceptive state that follows can be a powerful determinant of human motivated behavior, resulting in a blunted response to appetitive stimuli and an exaggerated response to noxious ones. We sought to assess such differential impact of visuo-vestibular mismatches on value through a task involving conflict monitoring. We therefore administered to 42 healthy participants a modified version of the Flankers task, in which distractors (arrows, pointing in either a congruent or incongruent direction) signaled the availability of monetary incentives (gains, losses, or neutral trials). While performing the task, participants received either galvanic vestibular stimulation (GVS), or sham stimulation. We have found impaired behavioral performances when value, which was attached to task-irrelevant information, was at stake. Gains and losses, interestingly, dissociated, and only the latter caused enhanced interference costs in the task, suggesting that negative incentives may be more effective in capturing human attention than positive ones. Finally, we have found some weak evidence for GVS to further increase the processing of losses, as suggested by even larger interference costs in this condition. Results were, however, overall ambiguous, and suggest that much more research is needed to better understand the link between the vestibular system and motivation.

Modulating the influence of recent trial history on attentional capture via transcranial magnetic stimulation (TMS) of right TPJ.

In visual search, salient yet task-irrelevant distractors in the stimulus array interfere with target selection. This is due to the unwanted shift of attention towards the salient stimulus-the so-called attentional capture effect, which delays deployment of attention onto the target. Although powerful and automatic, attentional capture by a salient distractor is nonetheless antagonized by distractor-filtering mechanisms and is further modulated by cross-trial contingencies: The distractor cost is typically more robust when no distraction has been experienced in the immediate past, compared to when a distractor was present on the immediately preceding trial. Here, we used transcranial magnetic stimulation (TMS) to shed light on the causal role of two crucial nodes of the ventral attention network, namely the Temporo-Parietal Junction (TPJ) and the Middle Frontal Gyrus (MFG), in the exogenous control of attention (i.e., attentional capture) and its history-dependent modulation. Participants were asked to discriminate the direction of a target arrow while ignoring a task-irrelevant salient distractor, when present. Immediately after display onset, 10 Hz triple-pulse TMS was delivered either to TPJ or MFG on the right hemisphere. Results demonstrated that stimulation of right TPJ-but not of right MFG, strongly modulated attentional capture as a function of the type of previous trial, by somewhat enhancing the distractor-related cost when the preceding trial was a distractor-absent trial and significantly decreasing the cost when the preceding trial was a distractor-present trial. These findings indicate that TMS of right TPJ exacerbates the effect of the recent history, likely reflecting enhanced updating of the predictive model that dynamically governs proactive distractor-filtering mechanisms. More generally, the results attest to a role of TPJ in mediating the history-dependent modulation of attentional capture.

The Topography of Visually Guided Grasping in the Premotor Cortex: A Dense-Transcranial Magnetic Stimulation (TMS) Mapping Study.

Visuomotor transformations at the cortical level occur along a network where posterior parietal regions are connected to homologous premotor regions. Grasping-related activity is represented in a diffuse, ventral and dorsal system in the posterior parietal regions, but no systematic causal description of a premotor counterpart of a similar diffuse grasping representation is available. To fill this gap, we measured the kinematics of right finger movements in 17 male and female human participants during grasping of three objects of different sizes. Single-pulse transcranial magnetic stimulation was applied 100 ms after visual presentation of the object over a regular grid of 8 spots covering the left premotor cortex (PMC) and 2 Sham stimulations. Maximum finger aperture during reach was used as the feature to classify object size in different types of classifiers. Classification accuracy was taken as a measure of the efficiency of visuomotor transformations for grasping. Results showed that transcranial magnetic stimulation reduced classification accuracy compared with Sham stimulation when it was applied to 2 spots in the ventral PMC and 1 spot in the medial PMC, corresponding approximately to the ventral PMC and the dorsal portion of the supplementary motor area. Our results indicate a multifocal representation of object geometry for grasping in the PMC that matches the known multifocal parietal maps of grasping representations. Additionally, we confirm that, by applying a uniform spatial sampling procedure, transcranial magnetic stimulation can produce cortical functional maps independent of a priori spatial assumptions.SIGNIFICANCE STATEMENT Visually guided actions activate a large frontoparietal network. Here, we used a dense grid of transcranial magnetic stimulation spots covering the whole premotor cortex (PMC), to identify with accurate spatial mapping the functional specialization of the human PMC during grasping movement. Results corroborate previous findings about the role of the ventral PMC in preshaping the fingers according to the size of the target. Crucially, we found that the medial part of PMC, putatively covering the supplementary motor area, plays a direct role in object grasping. In concert with findings in nonhuman primates, these results indicate a multifocal representation of object geometry for grasping in the PMC and expand our understanding of how our brain integrates visual and motor information to perform visually guided actions.

Rehabilitation and Biomarkers of Stroke Recovery: Study Protocol for a Randomized Controlled Trial.

Background: Stroke is a leading cause of disability. Nonetheless, the care pathway for stroke rehabilitation takes partially into account the needs of chronic patients. This is due in part to the lack of evidence about the mechanisms of recovery after stroke, together with the poor knowledge of related and influencing factors. Here we report on the study protocol "Rehabilitation and Biomarkers of Stroke Recovery," which consists of 7 work-packages and mainly aim to investigate the effects of long-term neurorehabilitation on stroke patients and to define a related profile of (clinical-biological, imaging, neurophysiological, and genetic-molecular) biomarkers of long-term recovery after stroke. The work-package 1 will represent the main part of this protocol and aims to compare the long-term effects of intensive self-rehabilitation vs. usual (rehabilitation) care for stroke. Methods: We planned to include a total of 134 adult subacute stroke patients (no more than 3 months since onset) suffering from multidomain disability as a consequence of first-ever unilateral ischemic stroke. Eligible participants will be randomly assigned to one of the following groups: intensive self-rehabilitation (based on the principles of "Guided Self-Rehabilitation Contract") vs. usual care (routine practice). Treatment will last 1 year, and patients will be evaluated every 3 months according to their clinical presentation. The following outcomes will be considered in the main work-package: Fugl-Meyer assessment, Cognitive Oxford Screen Barthel Index, structural and functional neuroimaging, cortical excitability, and motor and somatosensory evoked potentials. Discussion: This trial will deal with the effects of an intensive self-management rehabilitation protocol and a related set of biomarkers. It will also investigate the role of training intensity on long-term recovery after stroke. In addition, it will define a set of biomarkers related to post-stroke recovery and neurorehabilitation outcome in order to detect patients with greater potential and define long-term individualized rehabilitation programs. Clinical Trial Registration: www.ClinicalTrials.gov, identifier: NCT04323501.

Two Distinct Systems Represent Contralateral and Ipsilateral Sensorimotor Processes in the Human Premotor Cortex: A Dense TMS Mapping Study.

Animal brains contain behaviorally committed representations of the surrounding world, which integrate sensory and motor information. In primates, sensorimotor mechanisms reside in part in the premotor cortex (PM), where sensorimotor neurons are topographically clustered according to functional specialization. Detailed functional cartography of the human PM is still under investigation. We explored the topographic distribution of spatially dependent sensorimotor functions in healthy volunteers performing left or right, hand or foot, responses to visual cues presented in the left or right hemispace, thus combining independently stimulus side, effector side, and effector type. Event-related transcranial magnetic stimulation was applied to single spots of a dense grid of 10 points on the participants' left hemiscalp, covering the whole PM. Results showed: (1) spatially segregated hand and foot representations, (2) focal representations of contralateral cues and movements in the dorsal PM, and (3) distributed representations of ipsilateral cues and movements in the ventral and dorso-medial PM. The present novel causal information indicates that (1) the human PM is somatotopically organized and (2) the left PM contains sensory-motor representations of both hemispaces and of both hemibodies, but the hemispace and hemibody contralateral to the PM are mapped on a distinct, nonoverlapping cortical region compared to the ipsilateral ones.

Probing the Neural Mechanisms for Distractor Filtering and Their History-Contingent Modulation by Means of TMS.

In visual search, the presence of a salient, yet task-irrelevant, distractor in the stimulus array interferes with target selection and slows down performance. Neuroimaging data point to a key role of the frontoparietal dorsal attention network in dealing with visual distractors; however, the respective roles of different nodes within the network and their hemispheric specialization are still unresolved. Here, we used transcranial magnetic stimulation (TMS) to evaluate the causal role of two key regions of the dorsal attention network in resisting attentional capture by a salient singleton distractor: the frontal eye field (FEF) and the cortex within the intraparietal sulcus (IPS). The task of the participants (male/female human volunteers) was to discriminate the pointing direction of a target arrow while ignoring a task-irrelevant salient distractor. Immediately after stimulus onset, triple-pulse 10 Hz TMS was delivered either to IPS or FEF on either side of the brain. Results indicated that TMS over the right FEF significantly reduced the behavioral cost engendered by the salient distractor relative to left FEF stimulation. No such effect was obtained with stimulation of IPS on either side of brain. Interestingly, this FEF-dependent reduction in distractor interference interacted with the contingent trial history, being maximal when no distractor was present on the previous trial relative to when there was one. Our results provide direct causal evidence that the right FEF houses key mechanisms for distractor filtering, pointing to a pivotal role of the frontal cortex of the right hemisphere in limiting interference from an irrelevant but attention-grabbing stimulus.SIGNIFICANCE STATEMENT Visually conspicuous stimuli attract our attention automatically and interfere with performance by diverting resources away from the main task. Here, we applied transcranial magnetic stimulation over four frontoparietal cortex locations (frontal eye field and intraparietal sulcus in each hemisphere) to identify regions of the dorsal attention network that help limit interference from task-irrelevant, salient distractors. Results indicate that the right FEF participates in distractor-filtering mechanisms that are recruited when a distracting stimulus is encountered. Moreover, right FEF implements adjustments in distraction-filtering mechanisms following recent encounters with distractors. Together, these findings indicate a different hemispheric contribution of the left versus right dorsal frontal cortex to distraction filtering. This study expands our understanding of how our brains select relevant targets in the face of task-irrelevant, salient distractors.

Laws of concatenated perception: Vision goes for novelty, decisions for perseverance.

Every instant of perception depends on a cascade of brain processes calibrated to the history of sensory and decisional events. In the present work, we show that human visual perception is constantly shaped by two contrasting forces exerted by sensory adaptation and past decisions. In a series of experiments, we used multilevel modeling and cross-validation approaches to investigate the impact of previous stimuli and decisions on behavioral reports during adjustment and forced-choice tasks. Our results revealed that each perceptual report is permeated by opposite biases from a hierarchy of serially dependent processes: Low-level adaptation repels perception away from previous stimuli, whereas decisional traces attract perceptual reports toward the recent past. In this hierarchy of serial dependence, "continuity fields" arise from the inertia of decisional templates and not from low-level sensory processes. This finding is consistent with a Two-process model of serial dependence in which the persistence of readout weights in a decision unit compensates for sensory adaptation, leading to attractive biases in sequential perception. We propose a unified account of serial dependence in which functionally distinct mechanisms, operating at different stages, promote the differentiation and integration of visual information over time.

Getting rid of visual distractors: the why, when, how, and where.

Distractor suppression, or the ability to disregard salient distractors while dealing with task-relevant information, is a key component of selective attention. Recent research has shown that distractor suppression can take place in different circumstances and present itself in different guises, which is presumably paralleled by a multiplicity of underlying neural mechanisms. In this review article, we discuss a number of central themes concerning distractor suppression and the underlying neural mechanisms, and also highlight several unresolved issues that will have to be addressed in future investigations.

Revealing Dissociable Attention Biases in Chronic Smokers Through an Individual-Differences Approach.

Addiction is accompanied by attentional biases (AB), wherein drug-related cues grab attention independently of their perceptual salience. AB have emerged in different flavours depending on the experimental approach, and their clinical relevance is still debated. In chronic smokers we sought evidence for dissociable attention abnormalities that may play distinct roles in the clinical manifestations of the disorder. Fifty smokers performed a modified visual probe-task measuring two forms of AB and their temporal dynamics, and data on their personality traits and smoking history/status were collected. Two fully dissociable AB effects were found: A Global effect, reflecting the overall impact of smoke cues on attention, and a Location-specific effect, indexing the impact of smoke cues on visuospatial orienting. Importantly, the two effects could be neatly separated from one another as they: (i) unfolded with dissimilar temporal dynamics, (ii) were accounted for by different sets of predictors associated with personality traits and smoking history and (iii) were not correlated with one another. Importantly, the relevance of each of these two components in the single individual depends on a complex blend of personality traits and smoking habits, a result that future efforts addressing the clinical relevance of addiction-related AB should take into careful consideration.

Optic Nerve Degeneration and Reduced Contrast Sensitivity Due to Folic Acid Deficiency: A Behavioral and Electrophysiological Study in Rhesus Monkeys.

Purpose: The purpose of the research was to elucidate the role of folic acid (B9) deficiency in the development of nutritional optic neuritis and to characterize the neurophysiological consequences of optic nerve degeneration in the cortical visual system. Methods: A combined behavioral and electrophysiological approach was applied to study luminance contrast sensitivity in two macaque monkeys affected by nutritional optic neuritis and in two healthy monkeys for comparison. For one monkey, a follow-up approach was applied to compare visual performance before onset of optic neuropathy, during the disease, and after treatment. Results: Optic nerve degeneration developed as a consequence of insufficient dietary intake of folic acid in two exemplars of macaque monkeys. The degeneration resulted in markedly reduced luminance contrast sensitivity as assessed behaviorally. In one monkey, we also measured visual activity in response to varying contrast at the level of single neurons in the cortical visual system and found a striking reduction in contrast sensitivity, as well as a marked increase in the latency of neuronal responses. Prolonged daily folate supplementation resulted in a significant recovery of function. Conclusions: Folic acid deficiency per se can lead to the development of optic nerve degeneration in otherwise healthy adult animals. The optic nerve degeneration strongly affects contrast sensitivity and leads to a distinct reduction in the strength and velocity of the incoming signal to cortical visual areas of the macaque brain, without directly affecting excitability and functional properties of cortical neurons.