Effects of spatial location on distractor interference.

When target and distractor stimuli are close together, they activate the same neurons and there is ambiguity as to what the neural activity represents. It has been suggested that the ambiguity is resolved by spatial competition between target and nontarget stimuli. A competitive advantage is conveyed by bottom-up biases (e.g., stimulus saliency) and top-down biases (e.g., the match to a stored representation of the target stimulus). Here, we tested the hypothesis that regions with high perceptual performance may provide a bottom-up bias, resulting in increased distractor interference. Initially, we focused on two known anisotropies. At equal distance from central fixation, perceptual performance is better along the horizontal than the vertical meridian, and in the lower than in the upper visual hemifield. Consistently, interference from distractors on the horizontal meridian was greater than interference from distractors on the vertical meridian. However, distractors in the lower hemifield interfered less than distractors in the upper visual hemifield, which is contrary to the known anisotropy. These results were obtained with targets and distractors on opposite meridians. Further, we observed greater interference from distractors on the meridians compared with distractors on the diagonals, possibly reflecting anisotropies in attentional scanning. Overall, the results are only partially consistent with the hypothesis that distractor interference is larger for distractors on regions with high perceptual performance.

The PD Reflects Selection of Nontarget Locations, Not Distractor Suppression.

In visual search tasks, negative features provide information about stimuli that can be excluded from search. It has been shown that these negative features help participants to locate the target, possibly by attentional suppression of stimuli sharing the negative feature. Attentional suppression is assumed to be reflected in an event-related potential, the PD component. To provide a further test of these assumptions, we presented the color of the distractor at the start of a trial and asked participants to find the other colored stimulus in the subsequent search display. Consistent with attentional suppression, we observed a PD to a lateral distractor shown with a vertical target. However, the PD occurred in this condition only when target and distractor could also be on opposite sides of fixation. The effect of trial context on the PD suggests that the PD reflects a search strategy whereby participants select stimuli opposite to the distractor when trials with opposite placements occur during the experiment. Therefore, the PD to the distractor may in fact be an N2pc to the opposite stimulus, indicating that the distractor is not suppressed, but avoided by redirecting attentional selection to the opposite side.

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.

Does attentional suppression occur at the level of perception or decision-making? Evidence from Gaspelin et al.'s (2015) probe letter task.

Visual attention is often inadvertently captured by salient stimuli. It was suggested that it is possible to prevent attentional capture in some search tasks by suppressing salient stimuli below baseline. Evidence for attentional suppression comes from a probe task that was interleaved with the main search task. In the probe task of Gaspelin et al. (Psychol Sci 26(11):1740-1750, 2015. https://doi.org/10.1177/0956797615597913 ), letters were shown on the stimuli of the search display and participants had to identify as many letters as possible. Performance was found to be worse for letters shown on the distractor compared to non-salient non-target stimuli, suggesting that distractor processing was suppressed below baseline. However, it is unclear whether suppression occurred at the level of perception or decision-making because participants may have reported letters on the distractor less frequently than letters on nontargets. This decision-level bias may have degraded performance for letters on distractor compared to nontarget stimuli without changing perception. After replicating the original findings, we conducted two experiments where we avoided report bias by cueing only a single letter for report. We found that the difference between distractor and nontarget stimuli was strongly reduced, suggesting that decision-level processes contribute to attentional suppression. In contrast, the difference between target and non-target stimuli was unchanged, suggesting that it reflected perceptual-level enhancement of the target stimuli.

Biased Competition between Targets and Distractors Reduces Attentional Suppression: Evidence from the Positivity Posterior Contralateral and Distractor Positivity.

The biased competition account claims that competition between two stimuli increases when they are close together compared with when they are far apart. The reason is that nearby stimuli are more likely to be represented in the same receptive fields, requiring top-down or bottom-up biases to resolve the ambiguity. Consistent with biased competition, previous research showed that an index of attentional enhancement, the N2pc component, was attenuated when two targets were close together. In contrast, it is unclear whether distractor processing would also be attenuated when the distractor is close to the target. To answer this question, we used the additional singleton paradigm where a target is sometimes accompanied by a more salient, but entirely irrelevant, distractor. In the conditions of interest, the distance between the target and the distractor was systematically manipulated whereas the eccentricity to central fixation was always the same. The results showed that two indices of attentional suppression, the positivity posterior contralateral and distractor positivity components, were attenuated when the distractor was close to the target. Consistent with biased competition, attentional suppression of distractors was inhibited when the distance between target and distractor was short. The reduced attentional suppression of distractors with nearby targets may contribute to the increased behavioral interference with close distractors.

Attentional Templates Are Sharpened through Differential Signal Enhancement, Not Differential Allocation of Attention.

In visual search, the internal representation of the target feature is referred to as the attentional template. The attentional template can be broad or precise depending on the task requirements. In singleton search, the attentional template is broad because the target is the only colored element in the display. In feature search, a precise attentional template is required because the target is in a specific color in an array of varied colors. To measure the precision of the attentional template, we used a cue-target paradigm where cueing benefits decrease when the cue color differs from the target color. Consistent with broad and precise attentional templates, the decrease of cueing effects was stronger in feature than in singleton search. Measurements of ERPs showed that the N2pc elicited by the cue decreased with increasing color difference, suggesting that attention was more strongly captured by cues that were similar to the target. However, the cue-elicited N2pc did not differ between feature and singleton search, making it unlikely to reflect the mechanism underlying attentional template precision. Furthermore, there was no evidence for attentional suppression as there was no cue-elicited PD, even in conditions where the cueing benefit turned into a same-location cost. However, an index of signal enhancement, the contralateral positivity, reflected attention template precision. In general, there was sensory enhancement of the stimulus appearing at the cued location in the search display. With broad attentional templates, any stimulus at the cued location was enhanced, whereas enhancement was restricted to target-matching colors with precise attentional templates.

Visual selective attention and the control of tracking eye movements: a critical review.

People's eyes are directed at objects of interest with the aim of acquiring visual information. However, processing this information is constrained in capacity, requiring task-driven and salience-driven attentional mechanisms to select few among the many available objects. A wealth of behavioral and neurophysiological evidence has demonstrated that visual selection and the motor selection of saccade targets rely on shared mechanisms. This coupling supports the premotor theory of visual attention put forth more than 30 years ago, postulating visual selection as a necessary stage in motor selection. In this review, we examine to which extent the coupling of visual and motor selection observed with saccades is replicated during ocular tracking. Ocular tracking combines catch-up saccades and smooth pursuit to foveate a moving object. We find evidence that ocular tracking requires visual selection of the speed and direction of the moving target, but the position of the motion signal may not coincide with the position of the pursuit target. Further, visual and motor selection can be spatially decoupled when pursuit is initiated (open-loop pursuit). We propose that a main function of coupled visual and motor selection is to serve the coordination of catch-up saccades and pursuit eye movements. A simple race-to-threshold model is proposed to explain the variable coupling of visual selection during pursuit, catch-up and regular saccades, while generating testable predictions. We discuss pending issues, such as disentangling visual selection from preattentive visual processing and response selection, and the pinpointing of visual selection mechanisms, which have begun to be addressed in the neurophysiological literature.

Capture by Context Elements, Not Attentional Suppression of Distractors, Explains the PD with Small Search Displays.

Top-down control of attention allows us to resist attentional capture by salient stimuli that are irrelevant to our current goals. Recently, it was proposed that attentional suppression of salient distractors contributes to top-down control by biasing attention away from the distractor. With small search displays, attentional suppression of salient distractors may even result in reduced RTs on distractor-present trials. In support of attentional suppression, electrophysiological measures revealed a positivity between 200 and 300 msec contralateral to the distractor, which has been referred to as distractor positivity (PD). We reexamined distractor benefits with small search displays and found that the positivity to the distractor was followed by a negativity to the distractor. The negativity, referred to as N2pc, is considered an index of attentional selection of the contralateral element. Thus, attentional suppression of the distractor (PD) preceded attentional capture (N2pc) by the distractor, which is at odds with the idea that attentional suppression avoids attentional capture by the distractor. Instead, we suggest that the initial "PD" is not a positivity to the distractor but rather a negativity (N2pc) to the contralateral context element, suggesting that, initially, the context captured attention. Subsequently, the distractor was selected because, paradoxically, participants searched all lateral target positions (even when irrelevant) before they examined the vertical positions. Consistent with this idea, search times were shorter for lateral than vertical targets. In summary, the early voltage difference in small search displays is unrelated to distractor suppression but may reflect capture by the context.

Looking up improves performance in verbal tasks.

Earlier research suggested that gaze direction has an impact on cognitive processing. It is likely that horizontal gaze direction increases activation in specific areas of the contralateral cerebral hemisphere. Consistent with the lateralization of memory functions, we previously showed that shifting gaze to the left improves visuo-spatial short-term memory. In the current study, we investigated the effect of unilateral gaze on verbal processing. We expected better performance with gaze directed to the right because language is lateralized in the left hemisphere. Also, an advantage of gaze directed upward was expected because local processing and object recognition are facilitated in the upper visual field. Observers directed their gaze at one of the corners of the computer screen while they performed lexical decision, grammatical gender and semantic discrimination tasks. Contrary to expectations, we did not observe performance differences between gaze directed to the left or right, which is consistent with the inconsistent literature on horizontal asymmetries with verbal tasks. However, RTs were shorter when observers looked at words in the upper compared to the lower part of the screen, suggesting that looking upwards enhances verbal processing.

Motion integration is anisotropic during smooth pursuit eye movements.

Smooth pursuit eye movements (pursuit) are used to minimize the retinal motion of moving objects. During pursuit, the pattern of motion on the retina carries not only information about the object movement but also reafferent information about the eye movement itself. The latter arises from the retinal flow of the stationary world in the direction opposite to the eye movement. To extract the global direction of motion of the tracked object and stationary world, the visual system needs to integrate ambiguous local motion measurements (i.e., the aperture problem). Unlike the tracked object, the stationary world's global motion is entirely determined by the eye movement and thus can be approximately derived from motor commands sent to the eye (i.e., from an efference copy). Because retinal motion opposite to the eye movement is dominant during pursuit, different motion integration mechanisms might be used for retinal motion in the same direction and opposite to pursuit. To investigate motion integration during pursuit, we tested direction discrimination of a brief change in global object motion. The global motion stimulus was a circular array of small static apertures within which one-dimensional gratings moved. We found increased coherence thresholds and a qualitatively different reflexive ocular tracking for global motion opposite to pursuit. Both effects suggest reduced sampling of motion opposite to pursuit, which results in an impaired ability to extract coherence in motion signals in the reafferent direction. We suggest that anisotropic motion integration is an adaptation to asymmetric retinal motion patterns experienced during pursuit eye movements. NEW & NOTEWORTHY This study provides a new understanding of how the visual system achieves coherent perception of an object's motion while the eyes themselves are moving. The visual system integrates local motion measurements to create a coherent percept of object motion. An analysis of perceptual judgments and reflexive eye movements to a brief change in an object's global motion confirms that the visual and oculomotor systems pick fewer samples to extract global motion opposite to the eye movement.

A novel dissociation between representational momentum and representational gravity through response modality.

When people are required to indicate the vanishing location of a moving object, systematic biases forward, in the direction of motion, and downward, in the direction of gravity, are usually found. Both these displacements, called representational momentum and representational gravity, respectively, are thought to reflect anticipatory internal mechanisms aiming to overcome neural delays in the perception of motion. We challenge this view. There may not be such a single mechanism. Although both representational momentum and representational gravity follow a specific time-course, compatible with an anticipation of the object's dynamics, they do not seem to be commensurable with each other, as they are differentially modulated by relevant variables, such as eye movements and strength of motion signals. We found separate response components, one related to overt motor localization behaviour and one limited to purely perceptual judgement. Representational momentum emerged only for the motor localization task, revealing a motor overshoot. In contrast, representational gravity was mostly evident for spatial perceptual judgements. We interpret the results in support of a partial dissociation in the mechanisms that give rise to representational momentum and representational gravity, with the former but not the latter strongly modulated by the enrolment of the motor system.

Reference-frames in vision: Contributions of attentional tracking to nonretinotopic perception in the Ternus-Pikler display.

Perception depends on reference frames. For example, the "true" cycloidal motion trajectory of a reflector on a bike's wheel is invisible because we perceive the reflector motion relative to the bike's motion trajectory, which serves as a reference frame. To understand such an object-based motion perception, we suggested a "two-stage" model in which first reference frames are computed based on perceptual grouping (bike) and then features are attributed (reflector motion) based on group membership. The overarching goal of this study was to investigate how multiple features (i.e., motion, shape, and color) interact with attention to determine retinotopic or nonretinotopic reference frames. We found that, whereas tracking by focal attention can generate nonretinotopic reference-frames, the effect is rather small compared with motion-based grouping. Combined, our results support the two-stage model and clarify how various features and cues can work in conjunction or in competition to determine prevailing groups. These groups in turn establish reference frames according to which features are processed and bound together.

Placeholder objects shape spatial attention effects before eye movements.

In the time leading up to a saccade, the saccade target is perceptually enhanced compared to other objects in the visual field. This enhancement is attributed to a shift of spatial attention toward the target. We examined whether the presence of visual objects is critical for the perceptual enhancement at the saccade target to occur. We hypothesized that attention may need an object to focus on in order to be effective. We conducted four experiments using a dual-task design, where participants performed eye movements either to a location demarked by a placeholder or to an empty screen location where no object was displayed. At the same time, they discriminated a probe flashed at the location targeted by the eye movement or at one of two control locations. A strong perceptual advantage at the saccade target location was observed only when placeholders were displayed at the time of probe presentation. The complete absence of placeholders (Experiment 1), the presence of placeholders before but not during probe presentation (Experiment 3), and the presence of objects only around the saccade target (Experiments 3 and 4) led to a strong reduction in the saccade-target benefit. We conclude that placeholders may indeed be necessary to observe presaccadic enhancement at the saccade target. However, this is not because placeholders provide an object to focus attention on, but rather because they produce a masking (or crowding) effect. This detrimental effect is overcome by the presaccadic shift of attention, resulting in heightened perception only at the saccade target object.

Stronger interference from distractors in the right hemifield during visual search.

The orientation-bias hypothesis states that there is a bias to attend to the right visual hemifield (RVF) when there is spatial competition between stimuli in the left and right hemifield [Pollmann, S. (1996). A pop-out induced extinction-like phenomenon in neurologically intact subjects. Neuropsychologia, 34(5), 413-425. doi: 10.1016/0028-3932(95)00125-5 ]. In support of this hypothesis, stronger interference was reported for RVF distractors with contralateral targets. In contrast, previous studies using rapid serial visual presentation (RSVP) found stronger interference from distractors in the left visual hemifield (LVF). We used the additional singleton paradigm to test whether this discrepancy was due to the different distractor features that were employed (colour vs. orientation). Interference from the colour distractor with contralateral targets was larger in the RVF than in the LVF. However, the asymmetrical interference disappeared when observers had to search for an inconspicuous colour target instead of the inconspicuous shape target. We suggest that the LVF orienting-bias is limited to situations where search is driven by bottom-up saliency (singleton search) instead of top-down search goals (feature search). In contrast, analysis of the literature suggests the opposite for the LVF bias in RSVP tasks. Thus, the attentional asymmetry may depend on whether the task involves temporal or spatial competition, and whether search is based on bottom-up or top-down signals.

Gaze-cueing requires intact face processing - Insights from acquired prosopagnosia.

Gaze-cueing is the automatic spatial orienting of attention in the direction of perceived gaze. Participants respond faster to targets located at positions congruent with the direction of gaze, compared to incongruent ones (gaze cueing effect, GCE). However, it still remains unclear whether its occurrence depends on intact integration of information from the entire eye region or face, rather than simply the presence of the eyes per se. To address this question, we investigated the GCE in PS, an extensively studied case of pure acquired prosopagnosia. In our gaze-cueing paradigm, we manipulated the duration at which cues were presented (70ms vs. 400ms) and the availability of facial information (full-face vs. eyes-only). For 70ms cue duration, we found a context-dependent dissociation between PS and controls: PS showed a GCE for eyes-only stimuli, whereas controls showed a GCE only for full-face stimuli. For 400ms cue duration, PS showed gaze-cueing independently of stimulus context, whereas in healthy controls a GCE again emerged only for full-face stimuli. Our findings suggest that attentional deployment based on the gaze direction of briefly presented faces requires intact processing of facial information, which affords salience to the eye region.

Perceptual enhancement prior to intended and involuntary saccades.

Prior to an eye movement, attention is gradually shifted toward the point where the saccade will land. Our goal was to better understand the allocation of attention in an oculomotor capture paradigm for saccades that go straight to the eye movement target and for saccades that go to a distractor and are followed by corrective saccades to the target (i.e., involuntary saccades). We also sought to test facilitation at the future retinotopic location of target and nontarget objects, with the principal aim of verifying whether the remapping process accounts for the retinal displacement caused by involuntary saccades. Two experiments were run employing a dual-task design, primarily requiring participants to perform saccades toward a target while discriminating an asymmetric cross presented briefly before saccade onset. The results clearly show perceptual facilitation at the target location for goal-directed saccades and at the distractor location when oculomotor capture occurred. Facilitation was observed at a location relating to the remapping of a future saccade landing point, in sequences of oculomotor capture. In contrast, performance remained unaffected at the remapped location of a salient distracting object, which was not looked at. The findings are taken as evidence that presaccadic enhancement occurs prior to involuntary and voluntary saccades alike and that the remapping process also indiscriminatingly accounts for the retinal displacement caused by either.

The effect of gaze direction on the different components of visuo-spatial short-term memory.

Cerebral asymmetries and cortical regions associated with the upper and lower visual field were investigated using shifts of gaze. Earlier research suggests that gaze shifts to the left or right increase activation of specific areas of the contralateral hemisphere. We asked whether looking at one quadrant of the visual field facilitates the recall in various visuo-spatial tasks. The different components of visuo-spatial memory were investigated by probing memory for a stimulus matrix in each quadrant of the screen. First, memory for visual images or patterns was probed with a matrix of squares that was simultaneously presented and had to be reconstructed by mouse click. Better memory performance was found in the upper left quadrant compared to the three other quadrants indicating that both laterality and elevation are important. Second, positional memory was probed by subsequently presenting squares which prevented the formation of a visual image. Again, we found that gaze to the upper left facilitated performance. Third, memory for object-location binding was probed by asking observers to associate objects to particular locations. Higher performance was found with gaze directed to the lower quadrants irrespective of lateralization, confirming that only some components of visual short-term memory have shared neural substrates.

Amygdala activation for eye contact despite complete cortical blindness.

Cortical blindness refers to the loss of vision that occurs after destruction of the primary visual cortex. Although there is no sensory cortex and hence no conscious vision, some cortically blind patients show amygdala activation in response to facial or bodily expressions of emotion. Here we investigated whether direction of gaze could also be processed in the absence of any functional visual cortex. A well-known patient with bilateral destruction of his visual cortex and subsequent cortical blindness was investigated in an fMRI paradigm during which blocks of faces were presented either with their gaze directed toward or away from the viewer. Increased right amygdala activation was found in response to directed compared with averted gaze. Activity in this region was further found to be functionally connected to a larger network associated with face and gaze processing. The present study demonstrates that, in human subjects, the amygdala response to eye contact does not require an intact primary visual cortex.

Ocular tracking responses to background motion gated by feature-based attention.

Involuntary ocular tracking responses to background motion offer a window on the dynamics of motion computations. In contrast to spatial attention, we know little about the role of feature-based attention in determining this ocular response. To probe feature-based effects of background motion on involuntary eye movements, we presented human observers with a balanced background perturbation. Two clouds of dots moved in opposite vertical directions while observers tracked a target moving in horizontal direction. Additionally, they had to discriminate a change in the direction of motion (±10° from vertical) of one of the clouds. A vertical ocular following response occurred in response to the motion of the attended cloud. When motion selection was based on motion direction and color of the dots, the peak velocity of the tracking response was 30% of the tracking response elicited in a single task with only one direction of background motion. In two other experiments, we tested the effect of the perturbation when motion selection was based on color, by having motion direction vary unpredictably, or on motion direction alone. Although the gain of pursuit in the horizontal direction was significantly reduced in all experiments, indicating a trade-off between perceptual and oculomotor tasks, ocular responses to perturbations were only observed when selection was based on both motion direction and color. It appears that selection by motion direction can only be effective for driving ocular tracking when the relevant elements can be segregated before motion onset.

Gaze direction affects visuo-spatial short-term memory.

Hemispheric asymmetries were investigated by changing the horizontal position of stimuli that had to be remembered in a visuo-spatial short-term memory task. Observers looked at matrices containing a variable number of filled squares on the left or right side of the screen center. At stimulus offset, participants reproduced the positions of the filled squares in an empty response matrix. Stimulus and response matrices were presented in the same quadrant. We observed that memory performance was better when the matrices were shown on the left side of the screen. We distinguished between recall strategies that relied on visual or non-visual (verbal) cues and found that the effect of gaze position occurred more reliably in participants using visual recall strategies. Overall, the results show that there is a solid enhancement of visuo-spatial short-term memory when observers look to the left. In contrast, vertical position had no influence on performance. We suggest that unilateral gaze to the left activates centers in the right hemisphere contributing to visuo-spatial memory.