Covert Shifts Of Attention Research Articles

Oscillatory brain activity in the canonical alpha-band conceals distinct mechanisms in attention.

Brain oscillations in the alpha-band (8-14Hz) over posterior areas have been linked to specific processes in attention and perception. In particular, decreases in alpha-amplitude are thought to reflect activation of perceptually relevant brain areas for target engagement, while increases in alpha-amplitude have been associated with inhibition for distractor suppression. Traditionally, these alpha-changes have been viewed as two facets of the same process. However, more recent evidence calls for revisiting this interpretation. Here, we concurrently recorded MEG/EEG in 32 participants (19 females) during covert visuo-spatial attention shifts (spatial cues), and two control conditions (neutral cue, no-attention cue), while tracking fixational eye-movements. In disagreement with a single, perceptually relevant alpha-process, we found the typical alpha-modulations contra- and ipsilateral to the attention focus to be triple dissociated in their timing, topography, and spectral features: Ipsilateral alpha-increases occurred early, over occipital sensors, at a high alpha-frequency (10-14Hz) and were expressed during spatial attention (alpha spatial cue > neutral cue). By contrast, contralateral alpha-decreases occurred later, over parietal sensors, at a lower alpha-frequency (7-10Hz) and were associated with attention deployment in general (alpha spatial&neutral cue < no-attention cue). Additionally, the lateralized early alpha-increases but not -decreases during spatial attention coincided in time with directionally biased microsaccades. Overall, these findings suggest that the attention-related early alpha-increases and late -decreases reflect distinct, likely reflexive versus endogenously controlled attention mechanisms. We conclude that there is more than one perceptually relevant posterior alpha-oscillation, which need to be dissociated for a detailed account of their roles in perception and attention.Significance statement This study provides novel insights into perceptually relevant brain oscillations in the "canonical" alpha-band, and the neural correlates of covert and overt attention processes. This by simultaneously recording MEG and EEG, and isolating by design visuospatial and temporal expectations respectively, while tracking fixational eye-movements concurrently to the nominally covert attention shifts. The data reveal the presence of two, spatio-temporally and spectrally dissociated patterns of posterior alpha-changes that are distinctively associated with deployment of spatial and temporal anticipation, and eye-movement activity. This refines our understanding of the role of brain oscillations in perception and attention, the neural underpinnings of attention deployment in space and time, and provides methodological pointers for the study of perceptually relevant oscillatory activity through MEG/EEG.

Differences between overt, covert and natural attention shifts to emotional faces

In daily life, individuals pay attention to emotional facial expressions and dynamically choose how to shift their attention, i.e. either overtly (with eye-movements) or covertly (without eye-movements). However, research on attention to emotional faces has mostly been conducted in controlled laboratory settings, in which people were instructed where to look. The current preregistered study co-registered EEG and eye-tracking to investigate differences in emotion-driven attention between instructed and uninstructed natural attention shifts in 48 adults. While a central stimulus was presented to the participant, a face appeared in the periphery, showing either a happy, neutral or an angry expression. In three counterbalanced blocks participants were instructed to either move their eyes overtly to the peripheral face, keep fixating the center and therefore covertly shift their attention, or freely look wherever they would like to look. We found that emotional content had stronger effects on the amplitude of the Early Posterior Negativity when participants shifted attention naturally, and that natural shifts of attention differed from instructed shifts in both saccade behavior and neural mechanisms. In summary, our results emphasize the importance of investigating modulation of attention using paradigms that allow participants to allocate their attention naturally.

Orienting of attention and spatial cognition.

Humans orient to their sensory world through foveation of target location or through covert shifts of attention. Orienting provides primacy to the selected location and in humans improves the precision of discrimination. Covert orienting appears to arise separately from the mechanisms involved in saccadic eye movements. Covert orienting can serve to prioritize processing the target even increasing its subjective intensity and its acuity. However, this network does not appear to be involved in the operations related to binding and segmentation. Cells exist in the early visual cortex that are activated by both color and form features without attention, however, color and form appear to remain independent even when oriented to the target that is required to be reported. An understanding of the pathways that connect attention networks to memory networks may allow us to understand more complex aspects of spatial cognition and enhance orienting and thus improve spatial cognition.

A computational account of transsaccadic attentional allocation based on visual gain fields

Coordination of goal-directed behavior depends on the brain's ability to recover the locations of relevant objects in the world. In humans, the visual system encodes the spatial organization of sensory inputs, but neurons in early visual areas map objects according to their retinal positions, rather than where they are in the world. How the brain computes world-referenced spatial information across eye movements has been widely researched and debated. Here, we tested whether shifts of covert attention are sufficiently precise in space and time to track an object's real-world location across eye movements. We found that observers' attentional selectivity is remarkably precise and is barely perturbed by the execution of saccades. Inspired by recent neurophysiological discoveries, we developed an observer model that rapidly estimates the real-world locations of objects and allocates attention within this reference frame. The model recapitulates the human data and provides a parsimonious explanation for previously reported phenomena in which observers allocate attention to task-irrelevant locations across eye movements. Our findings reveal that visual attention operates in real-world coordinates, which can be computed rapidly at the earliest stages of cortical processing.

EEG decoding reveals task-dependent recoding of sensory information in working memory

Working memory (WM) supports future behavior by retaining perceptual information obtained in the recent past. The present study tested the hypothesis that WM recodes sensory information in a format that better supports behavioral goals. We recorded EEG while participants performed color delayed-estimation tasks where the colorwheel for the response was either randomly rotated or held fixed across trials. Accordingly, observers had to remember the exact colors in the Rotation condition, whereas they could prepare for a response based on the fixed mapping between the colors and their corresponding locations on the colorwheel in the No-Rotation condition. Results showed that the color reports were faster and more precise in the No-Rotation condition even when exactly the same set of colors were tested in both conditions. To investigate how the color information was maintained in the brain, we decoded the color using a multivariate EEG classification method. The decoding was limited to the stimulus encoding period in the Rotation condition, whereas it continued to be significant during the maintenance period in the No-Rotation condition, indicating that the color information was actively maintained in the condition. Follow-up analyses suggested that the prolonged decoding was not merely driven by the covert shift of attention but rather by the recoding of sensory information into an action-oriented response format. Together, these results provide converging evidence that WM flexibly recodes sensory information depending on the specific task context to optimize subsequent behavioral performance.

Spatial Decoding for Gaze Independent Brain-Computer Interface Based on Covert Visual Attention Shift Using Electroencephalography.

The gaze-independent brain-computer interface (BCI) device is used to re-establish interaction for individuals who have abnormal eye movement. It may be possible to control the BCI by shifting your attention spatially. However, spatial attention is rarely employed to increase the effectiveness of target detection and is typically used to provide a simple "yes" or "no" response to the target recognition inquiry. To improve the effectiveness of detecting target, it is crucial to take advantage of the possible advantages of spatial attention. N2-posterior-contralateral (N2pc) component reflects correlates of visual spatial attention and is used to determine target position. In this study, a long-short-term memory (LSTM) network is used to answer "yes/no" questions by decoding covert spatial attention based on N2pc characteristics using EEG signals. The proposed LSTM-based model's average decoding accuracy is 92.79%. The target detection efficiency was successfully increased by about 4% when compared to conventional machine learning algorithms. The proposed model is tested on the independent dataset to validate its performance. The results of this work show that N2pc characteristics can be employed in gaze-independent BCIs for tracking covert attention shifts, which may help persons with poor eye mobility to connect with their environment.

Is covert attention necessary for programming accurate saccades? Evidence from saccade-locked event-related potentials.

For decades, researchers have assumed that shifts of covert attention mandatorily occur prior to eye movements to improve perceptual processing of objects before they are fixated. However, recent research suggests that the N2pc component-a neural measure of covert attentional allocation-does not always precede eye movements. The current study investigated whether the N2pc component mandatorily precedes eye movements and assessed its role in the accuracy of gaze control. In three experiments, participants searched for a letter of a specific color (e.g., red) and directed gaze to it as a response. Electroencephalograms and eye movements were coregistered to determine whether neural markers of covert attention preceded the initial shift of gaze. The results showed that the presaccadic N2pc only occurred under limited conditions: when there were many potential target locations and distractors. Crucially, there was no evidence that the presence or magnitude of the presaccadic N2pc was associated with improved eye movement accuracy in any of the experiments. Interestingly, ERP decoding analyses were able to classify the target location well before the eyes started to move, which likely reflects a presaccadic cognitive process that is distinct from the attentional process measured by the N2pc. Ultimately, we conclude that the covert attentional mechanism indexed by the N2pc is not necessary for precise gaze control.

Are microsaccades biased similarly during external and internal shifts of covert attention?

Are microsaccades biased similarly during external and internal shifts of covert attention?

Head and body cues guide eye movements and facilitate target search in real-world videos.

Static gaze cues presented in central vision result in observer shifts of covert attention and eye movements, and benefits in perceptual performance in the detection of simple targets. Less is known about how dynamic gazer behaviors with head and body motion influence search eye movements and performance in perceptual tasks in real-world scenes. Participants searched for a target person (yes/no task, 50% presence), whereas watching videos of one to three gazers looking at a designated person (50% valid gaze cue, looking at the target). To assess the contributions of different body parts, we digitally erase parts of the gazers in the videos to create three different body parts/whole conditions for gazers: floating heads (only head movements), headless bodies (only lower body movements), and the baseline condition with intact head and body. We show that valid dynamic gaze cues guided participants' eye movements (up to 3 fixations) closer to the target, speeded the time to foveate the target, reduced fixations to the gazers, and improved target detection. The effect of gaze cues in guiding eye movements to the search target was the smallest when the gazer's head was removed from the videos. To assess the inherent information about gaze goal location for each body parts/whole condition, we collected perceptual judgments estimating gaze goals by a separate group of observers with unlimited time. Observers' perceptual judgments showed larger estimate errors when the gazer's head was removed. This suggests that the reduced eye movement guidance from lower body cueing is related to observers' difficulty extracting gaze information without the presence of the head. Together, the study extends previous work by evaluating the impact of dynamic gazer behaviors on search with videos of real-world cluttered scenes.

Exploring the relationship between oculomotor preparation and gaze-cued covert shifts in attention.

Eye gaze plays dual perceptual and social roles in everyday life. Gaze allows us to select information, while also indicating to others where we are attending. There are situations, however, where revealing the locus of our attention is not adaptive, such as when playing competitive sports or confronting an aggressor. It is in these circumstances that covert shifts in attention are assumed to play an essential role. Despite this assumption, few studies have explored the relationship between covert shifts in attention and eye movements within social contexts. In the present study, we explore this relationship using the saccadic dual-task in combination with the gaze-cueing paradigm. Across two experiments, participants prepared an eye movement or fixated centrally. At the same time, spatial attention was cued with a social (gaze) or non-social (arrow) cue. We used an evidence accumulation model to quantify the contributions of both spatial attention and eye movement preparation to performance on a Landolt gap detection task. Importantly, this computational approach allowed us to extract a measure of performance that could unambiguously compare covert and overt orienting in social and non-social cueing tasks for the first time. Our results revealed that covert and overt orienting make separable contributions to perception during gaze-cueing, and that the relationship between these two types of orienting was similar for both social and non-social cueing. Therefore, our results suggest that covert and overt shifts in attention may be mediated by independent underlying mechanisms that are invariant to social context.

Pre-saccadic attention relies more on suppression than does covert attention.

During covert and pre-saccadic attentional shifts, it is unclear how facilitation and suppression processes interact for target selection. A recent countermanding task pointed to greater suppression at unattended locations during trials with saccades compared to trials without saccades (i.e., fixation and successful stop trials), whereas target facilitation did not differ. It is unknown whether this finding is restricted to countermanding paradigms that involve inhibitory processes. To test this, we adapted Gaspelin and colleagues (2015)'s attention capture task where, within the same block, one location was primed with frequent line discrimination trials, and all locations were occasionally probed using letters report trials. Participants also performed a baseline condition without priming. We tested 15 participants and examined how performance at non-primed locations was affected by covert versus pre-saccadic attention in blocks of four or six items, as well as by position from the primed location and timing from saccade onset. For both attention conditions, letter report at non-primed locations was worse compared to baseline, demonstrating suppression, and letter report at primed location was better, demonstrating facilitation. In saccades trials, letter report was better at primed locations and worse at non-primed locations compared to fixation trials. The timing of this additional pre-saccadic suppression differed from saccadic suppression. In both attention conditions, suppression was greater when primed and non-primed locations were within the same hemifield or in diagonal opposite quadrants. These results confirmed that attention preceding saccade execution suppressed non-primed locations to a larger extent than covert attention, with the same spatial quadrant effect.

Virtual occlusion effects on the perception of self-initiated visual stimuli

Virtual reality (VR) has established itself as a useful tool in the study of human perception in the laboratory. A recent study introduced a new approach to examine visual sensory attenuation (SA) effects in VR. Hand movements triggered the appearance of Gabor stimuli, which were either presented behind the participant’s hand – not rendered in VR (“virtual occlusion”) - or elsewhere on the display. Virtual occlusion led to a rightward shift of the psychometric curve, suggesting that self-generated hand movements reduced the perceived contrast of the stimulus. Since such attenuation effects might provide a window into the predictive processing of the sensory and cognitive apparatus, we sought to better understand the nature of the virtual occlusion effects. In our study, the presentation of test stimuli was either self-initiated, self-initiated with a variable delay, or triggered externally; the test stimuli were occluded or not. In conflict with our hypothesis, we found moderate to strong evidence for an absence of any horizontal shifts between the psychometric curves. However, virtual occlusion was associated with a decrease in the slope of the psychometric function. Our results suggest that virtual occlusion attenuated the relative perceptual sensitivity, so that participants had more difficulty discriminating contrast differences when the test stimulus was presented behind the hand. We tentatively conclude that, in the visual domain, the discriminability of stimulus intensity is modified by internal predictive cues (i.e., proprioception), possibly linked to shifts in covert spatial attention.

A functional role for oculomotor preparation in mental arithmetic evidenced by the abducted eye paradigm.

Solving subtraction and addition problems is accompanied by spontaneous leftward and rightward gaze shifts, respectively. These shifts have been related to attentional processes involved in mental arithmetic, but whether these processes induce overt attentional shifts mediated by the activation of the motor programs underlying lateral eye movements or covert shifts only is still unknown. Here, we used the abducted eye paradigm to selectively disrupt activation of the oculomotor system and prevent oculomotor preparation, which affects overt but not covert attentional shifts. Participants had to mentally solve addition and subtraction problems while fixating a screen positioned either in front of them or laterally to their left or right such that they were physically unable to programme and execute saccades further into their temporal field while they still could do so in their nasal field. In comparison to the frontal condition, rightward eye abduction impaired additions (with carrying), and leftward eye abduction impaired subtractions (with borrowing) showing that at least some arithmetic problems rely on processes dedicated to overt attentional shifts. We propose that when solving arithmetic problems requires procedures such as carrying and borrowing, oculomotor mechanisms operating on a mental space transiently built in working memory are recruited to represent one numerical magnitude in relation to another (e.g. the first operand and the result).

Pupil size variations reveal covert shifts of attention induced by numbers

The pupil light response is more than a pure reflexive mechanism that reacts to the amount of light entering the eye. The pupil size may also react to the luminance of objects lying in the visual periphery, revealing the locus of covert attention. In the present study, we took advantage of this response to study the spatial coding of abstract concepts with no physical counterpart: numbers. The participants' gaze was maintained fixed in the middle of a screen whose left and right parts were dark or bright, and variations in pupil size were recorded during an auditory number comparison task. The results showed that small numbers accentuated pupil dilation when the darker part of the screen was on the left, while large numbers accentuated pupil dilation when the darker part of the screen was on the right. This finding provides direct evidence for covert attention shifts on a left-to-right oriented mental spatial representation of numbers. From a more general perspective, it shows that the pupillary response to light is subject to modulation from spatial attention mechanisms operating on mental contents.

The dynamics of microsaccade amplitude reflect shifting of covert attention

Attention flexibly shifts between spatial locations to accommodate task demands. The present study examined if the dynamics of attentional shifting are seen in microsaccades whose direction has been shown to accompany the shifts of covert attention. In a spatial cueing task, the cue predicted the target location on 100%, 75%, or 50% of the trials. The results revealed that microsaccade rate and amplitude were both reduced following cue onset and then rebounded. Both microsaccade rate and amplitude were biased towards the opposite direction of the cue and then returned to the cued direction. Importantly, the cue validity modulated the temporal profile of microsaccade amplitude but had little impact on the temporal profile of microsaccade rate. In line with this, the cueing effect measured with target response accuracy was correlated with the microsaccade amplitude only. These results indicate that the temporal dynamics of microsaccade amplitude reflect shifting of covert attention.

Involvement of the dorsal and ventral attention networks invisual attention span.

Visual attention span (VAS), which refers to the window size of multielement parallel processing in a short time, plays an important role in higher‐level cognition (e.g., reading) as required by encoding large amounts of information input. However, it is still a matter of debate about the underlying neural mechanism of VAS. In the present study, a modified visual 1‐back task was designed by using nonverbal stimuli and nonverbal responses, in which possible influences of target presence and position were considered to identify more pure VAS processing. A task‐driven functional magnetic resonance imaging (fMRI) experiment was then performed, and 30 healthy adults participated in this study. Results of confirmatory and exploratory analyses consistently revealed that both dorsal attention network (DAN) and ventral attention network (VAN) were significantly activated during this visual simultaneous processing. In particular, more significant activation in the left superior parietal lobule (LSPL), as compared to that in the bilateral inferior frontal gyrus (IFGs), suggested a greater involvement of DAN in VAS‐related processing in contrast to VAN. In addition, it was also found that the activation in temporoparietal junctions (TPJs) were suppressed during multielement processing only in the target‐absent condition. The current results suggested the recruitment of LSPL in covert attentional shifts and top‐down control of VAS resources distribution during the rapid visual simultaneous processing, as well as the involvement of bilateral IFGs (especially RIFG) in both VAS processing and inhibitory control. The present findings might bring some enlightenments for diagnosis of the atypicality of attentional disorders and reading difficulties.

Reading Specific Small Saccades Predict Individual Phonemic Awareness and Reading Speed.

Small fixational eye-movements are a fundamental aspect of vision and thought to reflect fine shifts in covert attention during active viewing. While the perceptual benefits of these small eye movements have been demonstrated during a wide range of experimental tasks including during free viewing, their function during reading remains surprisingly unclear. Previous research demonstrated that readers with increased microsaccade rates displayed longer reading speeds. To what extent increased fixational eye movements are, however, specific to reading and might be indicative of reading skill deficits remains, however, unknown. To address this topic, we compared the eye movement scan paths of 13 neurotypical individuals and 13 subjects diagnosed with developmental dyslexia during short story reading and free viewing of natural scenes. We found that during reading only, dyslexics tended to display small eye movements more frequently compared to neurotypicals, though this effect was not significant at the population level, as it could also occur in slow readers not diagnosed as dyslexics. In line with previous research, neurotypical readers had twice as many regressive compared to progressive microsaccades, which did not occur during free viewing. In contrast, dyslexics showed similar amounts of regressive and progressive small fixational eye movements during both reading and free viewing. We also found that participants with smaller fixational saccades from both neurotypical and dyslexic samples displayed reduced reading speeds and lower scores during independent tests of reading skill. Slower readers also displayed greater variability in the landing points and temporal occurrence of their fixational saccades. Both the rate and spatio-temporal variability of fixational saccades were associated with lower phonemic awareness scores. As none of the observed differences between dyslexics and neurotypical readers occurred during control experiments with free viewing, the reported effects appear to be directly related to reading. In summary, our results highlight the predictive value of small saccades for reading skill, but not necessarily for developmental dyslexia.

Visual attentional orienting by eye gaze: A meta-analytic review of the gaze-cueing effect.

Given limitations in the amount of visual information that a person can simultaneously process through to conscious perception, selective visual attention is necessary. Visual signals in the environment aid this selection process by triggering reflexive shifts of covert attention to locations of potential importance. One such signal appears to be others' eye gaze. Indeed, a gaze-cueing effect, whereby healthy adults respond faster to targets that are presented at locations cued rather than miscued by eye gaze has been consistently observed in the empirical literature. Critically though, the influences of task and cue features on this effect are not well understood. To address this gap, we report a meta-analytic integration of 423 gaze-cueing effects using a multilevel approach. A gaze-cueing effect emerged across all levels of the assessed task and cue features, indicating that others' eye gaze reliably directs observers' attention. We found that the magnitude of the gaze-cueing effect was moderated by whether direct gaze cues preceded directional gaze cues or not; the cue-target stimulus onset asynchrony (SOA), whether participants had to detect, localize, or categorize targets; and the cue's facial expression. Whether or not the gaze cue remained on screen after the target appeared, and whether schematic faces, computer-generated faces, or images of real faces were used as cues, did not appear to reliably function as moderators. The theoretical implications of these findings are discussed, particularly in relation to the social attention system. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

The Study of Security Priming on Avoidant Attentional Biases: Combining Microsaccadic Eye-Movement Measurement With a Dot-Probe Task.

Microsaccades are small fixational eye movements that have shown to index covert attentional shifts. The present experiment combined microsaccades with performance measures from a dot-probe task to study influences of attachment security priming on the attentional biases of individuals high in attachment avoidance. Security priming is an experimental manipulation aimed at boosting felt security. Using a randomized, mixed design, we measured differences in attentional vigilance toward angry and neutral faces as a function of priming (neutral vs. secure) and attachment avoidance. Individuals high in avoidance habitually tend to withdraw from, or otherwise dismiss, emotionally salient stimuli. Here, we operationalized attentional withdrawal based on both task performance in the dot-probe task and microsaccadic movements. In addition, unlike previous studies where priming salience for the individual participant has been unclear, we used a standardized narrative method for attachment script assessment, securing an indication of how strongly each participant was primed. Dot-probe data significantly captured the link between avoidance and attentional disengagement, though from all facial stimuli (angry and neutral). Although microsaccadic movements did not capture avoidant attentional disengagement, they positively correlated to dot-probe data suggesting measurement convergence. Avoidance was associated with weaker security priming and no overall effect of priming on attention was found, indicating a need for further exploration of suitable priming methods to bypass avoidant deactivation. Our results provide a first indication that, as an implicit looking measure, microsaccadic movements can potentially reveal where early attention is directed at the exact moment of stimulus presentation.

Have a look - the role of covert attention shifts for object integration: Evidence from pupillometry.

Have a look - the role of covert attention shifts for object integration: Evidence from pupillometry.