Current foveal inspection and previous peripheral preview influence subsequent eye movement decisions.

Humans visually inspect the world with their fovea and select new parts of the scene using saccadic eye movements. Foveal inspection and the decision of where and when to look next proceed simultaneously, but there is mixed evidence concerning their independence. Here, we tested their interdependence using drift-diffusion modeling. Participants first made a saccade to a predetermined inspection target and subsequently decided between two selection targets. We found that the inspected target's meaningfulness and the opportunity to preview it peripherally affects fixation durations and the upcoming saccadic selection. Drift-diffusion modeling showed that meaningfulness and the absence of peripheral preview can both delay the subsequent saccadic decision process and affect the rate at which peripheral information is accumulated. Our results thus show that foveal inspection and peripheral selection are dependent on each other and that peripheral information can be maintained across the saccade to influence subsequent eye movement decisions.

A nearby distractor does not influence hand movements.

When interacting with the environment, our manual actions are often preceded by an eye movement. This suggests that the processes underlying target selection in hand and eye movements may be coupled. It is known that when a distractor is presented close to a target, the endpoint of an eye movement will be biased towards the distractor. The size of this so-called global effect decreases when more viewing time is available. Here we investigate whether a similar effect is also present in hand movements. If the processes underlying target selection for hand and eye movements are indeed coupled, a similar bias should be present in hand movements as well. To test this, we adopted a classic global effect paradigm but applied it to goal-directed hand movements. We show that the endpoints of hand movements are unbiased for all but one participant, irrespective of the viewing time. These results suggest that the processes underlying target selection for hand movements operate independently from those for eye movements.

Rapid spatial oculomotor updating across saccades is malleable.

The oculomotor system uses a sophisticated updating mechanism to adjust for large retinal displacements which occur with every saccade. Previous studies have shown that updating operates rapidly and starts before saccade is initiated. Here we used saccade adaptation to alter life-long expectations about how a saccade changes the location of an object on the retina. Participants made a sequence of one horizontal and one vertical saccade and ignored an irrelevant distractor. The time-course of oculomotor updating was estimated using saccade curvature of the vertical saccade, relative to the distractor. During the first saccade both saccade targets were shifted on 80% of trials, which induced saccade adaptation (Experiment 1). Critically, since the distractor was left stationary, successful saccade adaptation (e.g., saccade becoming shorter) meant that after the first saccade the distractor appeared in a different hemifield than without adaptation. After adaptation, second saccades curved away only from the newly learned distractor location starting at 80 ms after the first saccade. When on the minority of trials (20%) the targets were not shifted, saccades again first curved away from the newly learned (now empty) location, but then quickly switched to curving away from the life-long learned, visible location. When on some trials the distractor was removed during the first saccade, saccades curved away only from the newly learned (but empty) location (Experiment 2). The results show that updating of locations across saccades is not only fast, but is highly malleable, relying on recently learned sensorimotor contingencies.

Visual information is required to reduce the global effect.

When a distractor appears in close proximity to a saccade target, the saccadic end point is biased towards the distractor. This so-called global effect reduces with the latency of the saccade if the saccade is visually guided. We recently reported that the global effect does not reduce with the latency of a double-step memory-guided saccade. The aim of this study was to investigate why the global effect in memory-guided saccades does not show the typically observed reduction with saccadic latency. One possibility is that reduction of the global effect requires continuous access to visual information about target and distractor locations, which is lacking in the case of a memory-guided saccade. Alternatively, participants may be inclined to routinely preprogram a memory-guided saccade at the moment the visual information disappears, with the result that a memory-guided saccade is typically programmed on the basis of an earlier representation than necessary. To distinguish between these alternatives, two potential targets were presented, and participants were asked to make a saccade to one of them after a delay. In one condition, the target identity was precued, allowing preprogramming of the saccade, while in another condition, it was revealed by a retro cue after the delay. The global effect remained present in both conditions. Increasing visual exposure of target and distractor led to a reduction of the global effect, irrespective of whether participants could preprogram a saccade or not. The results suggest that continuous access to visual information is required in order to eliminate the global effect.

Voluntary and Involuntary Control of Attention in Adolescents Born Very Preterm: A Study of Eye Movements.

Very preterm birth is associated with attention deficits that interfere with academic performance. A better understanding of attention processes is necessary to support very preterm born children. This study examined voluntary and involuntary attentional control in very preterm born adolescents by measuring saccadic eye movements. Additionally, these control processes were related to symptoms of inattention, intelligence, and academic performance. Participants included 47 very preterm and 61 full-term born 13-years-old adolescents. Oculomotor control was assessed using the antisaccade and oculomotor capture paradigm. Very preterm born adolescents showed deficits in antisaccade but not in oculomotor capture performance, indicating impairments in voluntary but not involuntary attentional control. These impairments mediated the relation between very preterm birth and inattention, intelligence, and academic performance.

Spatiotopic and saccade-specific transsaccadic memory for object detail.

The content and nature of transsaccadic memory are still a matter of debate. Brief postsaccadic target blanking was demonstrated to recover transsaccadic memory and defeat saccadic suppression of displacement. We examined whether blanking would also support transsaccadic transfer of detailed form information. Observers saccaded to a peripheral, checkerboard-like stimulus and reported whether an intrasaccadic change had occurred in its upper or lower half. On half of the trials, the stimulus was blanked for 200 ms with saccade onset. In a fixation condition, observers kept fixation but the stimulus was displaced from periphery to fixation, mimicking the retinal events of the saccade condition. Results show that stimulus blanking improves transsaccadic change detection, with performance being far superior to the retinally equivalent fixation condition. Our findings argue in favor of a remapped memory trace that can be accessed only in the blanking condition, when not being overwritten by the salient postsaccadic stimulus.

Detection of object displacement during a saccade is prioritized by the oculomotor system.

The human eye-movement system is equipped with a sophisticated updating mechanism that can adjust for large retinal displacements produced by saccadic eye movements. The nature of this updating mechanism is still highly debated. Previous studies have demonstrated that updating can occur very rapidly and is initiated before the start of a saccade. In the present study, we used saccade curvature to demonstrate that the oculomotor system is tuned for detecting object displacements during saccades. Participants made a sequence of saccades while ignoring an irrelevant distractor. Curvature of the second saccade relative to the distractor was used to estimate the time course of updating. Saccade curvature away from the presaccadic location of the distractor emerged as early as 80 ms after the first saccade when the distractor was displaced during a saccade. This is about 50 ms earlier than when a distractor was only present before a saccade, only present after a saccade, or remained stationary across a saccade. This shows that the oculomotor system prioritizes detection of object displacements during saccades, which may be useful for guiding corrective saccades. The results also challenge previous views by demonstrating the additional role of postsaccadic information in updating target-distractor competition across saccades.

Updating spatial working memory in a dynamic visual environment.

The present review describes recent developments regarding the role of the eye movement system in representing spatial information and keeping track of locations of relevant objects. First, we discuss the active vision perspective and why eye movements are considered crucial for perception and attention. The second part focuses on the question of how the oculomotor system is used to represent spatial attentional priority, and the role of the oculomotor system in maintenance of this spatial information. Lastly, we discuss recent findings demonstrating rapid updating of information across saccadic eye movements. We argue that the eye movement system plays a key role in maintaining and rapidly updating spatial information. Furthermore, we suggest that rapid updating emerges primarily to make sure actions are minimally affected by intervening eye movements, allowing us to efficiently interact with the world around us.

The Limits of Predictive Remapping of Attention Across Eye Movements.

With every eye movement, visual input projected onto our retina changes drastically. The fundamental question of how we keep track of relevant objects and movement targets has puzzled scientists for more than a century. Recent advances suggested that this can be accomplished through the process of predictive remapping of visual attention to the future post-saccadic locations of relevant objects. Evidence for the existence of predictive remapping of attention was first provided by Rolfs et al. (2011) (Nature Neuroscience, 14, 252-256). However, they used a single distant control location away from the task-relevant locations, which could have biased the allocation of visual attention. In this study we used a similar experimental paradigm as Rolfs et al. (2011), but probed attention equally likely at all possible locations. Our results showed that discrimination performance was higher at the remapped location than at a distant control location, but not compared to the other two control locations. A re-analysis of the results obtained by Rolfs et al. (2011) revealed a similar pattern. Together, these findings suggest that it is likely that previous reports of the predictive remapping of attention were due to a diffuse spread of attention to the task-relevant locations rather than to a specific shift toward the target's future retinotopic location.

Forget binning and get SMART: Getting more out of the time-course of response data.

Many experiments aim to investigate the time-course of cognitive processes while measuring a single response per trial. A common first step in the analysis of such data is to divide them into a limited number of bins. As we demonstrate here, the way one chooses these bins can considerably influence the resulting time-course. As a solution to this problem, we here present the smoothing method for analysis of response time-course (SMART)-a complete package for reconstructing the time-course from one-sample-per-trial data and performing statistical analysis. After smoothing the data, the SMART weights the data based on the effective number of data points per participant. A cluster-based permutation test then determines at which moments the responses differ from a baseline or between two conditions. We show here that, in contrast to contemporary binning methods, the chosen temporal resolution has a negligible effect on the SMART reconstructed time-course. To facilitate its use, the SMART method, accompanied by a tutorial, is available as an open-source package.

Target-distractor competition cannot be resolved across a saccade.

When a distractor is presented in close spatial proximity to a target, a saccade tends to land in between the two objects rather than on the target. This robust phenomenon (also referred to as the global effect) is thought to reflect unresolved competition between target and distractor. It is unclear whether this landing bias persists across saccades since a saccade displaces the retinotopic representations of target and distractor. In the present study participants made successive saccades towards two saccadic targets which were presented simultaneously with an irrelevant distractor in close proximity to the second saccade target. The second saccade was either visually-guided or memory-guided. For the memory-guided trials, the second saccade showed a landing bias towards the location of the distractor, despite the disappearance of the distractor after the first saccade. In contrast, for the visually-guided trials, the bias was corrected and the landing bias was eliminated, even for saccades with the shortest intersaccadic intervals. This suggests that the biased saccade plan was remapped across the first saccade. Therefore, we conclude that the target-distractor competition was not resolved across a saccade, but can be resolved based on visual information that is available after a saccade.

Distractor displacements during saccades are reflected in the time-course of saccade curvature.

Every time we make a saccade we form a prediction about where objects are going to be when the eye lands. This is crucial since the oculomotor system is retinotopically organized and every saccade drastically changes the projection of objects on the retina. We investigated how quickly the oculomotor system accommodates new spatial information when a distractor is displaced during a saccade. Participants performed sequences of horizontal and vertical saccades and oculomotor competition was induced by presenting a task-irrelevant distractor before the first saccade. On half of the trials the distractor remained in the same location after the first saccade and on the other half the distractor moved during the first saccade. Curvature of the second saccade was used to track target-distractor competition. At short intersaccadic intervals, saccades curved away from the original distractor location, confirming that in the oculomotor system spatiotopic representations emerge rapidly and automatically. Approximately 190 ms after the first saccade, second saccades curved away from the new distractor location. These results show that after a saccade the oculomotor system is initially driven by the spatial prediction made before the saccade, but it is able to quickly update these spatial predictions based on new visual information.

Rapid updating of spatial working memory across saccades.

Each time we make an eye movement, positions of objects on the retina change. In order to keep track of relevant objects their positions have to be updated. The situation becomes even more complex if the object is no longer present in the world and has to be held in memory. In the present study, we used saccadic curvature to investigate the time-course of updating a memorized location across saccades. Previous studies have shown that a memorized location competes with a saccade target for selection on the oculomotor map, which leads to saccades curving away from it. In our study participants performed a sequence of two saccades while keeping a location in memory. The trajectory of the second saccade was used to measure when the memorized location was updated after the first saccade. The results showed that the memorized location was rapidly updated with the eyes curving away from its spatial coordinates within 130 ms after the first eye movement. The time-course of updating was comparable to the updating of an exogenously attended location, and depended on how well the location was memorized.

Eye abduction reduces but does not eliminate competition in the oculomotor system.

Although it is well established that there is a tight coupling between covert attention and the eye movement system there is an ongoing controversy whether this relationship is functional. Previous studies demonstrated that disrupting the ability to execute an eye movement interferes with the allocation of covert attention. One technique that prevents the execution of an eye movement involves the abduction of the eye in the orbit while presenting the stimuli outside of the effective oculomotor range (Craighero, Nascimben, & Fadiga, 2004). Although eye abduction is supposed to disrupt activation of the oculomotor program responsible for the shift of covert attention, this crucial assumption has never been tested experimentally. In the present study we used saccadic curvature to examine whether eye abduction eliminates the target-distractor competition in the oculomotor system. We experimentally reduced the ability to execute saccades by abducting the eye by 30° (monocular vision). This way the peripheral part of the temporal hemifield was located outside the oculomotor range. Participants made a vertical eye movement while on some trials a distractor was shown either inside or outside of the oculomotor range. The curvature away from distractors located outside the oculomotor range was reduced, but not completely eliminated. This confirms that eye abduction influences the activation of the oculomotor program, but points to the fact that other forms of motor planning, such as head movements are also represented in the oculomotor system. The results are in line with the idea that covert attention is an emerging property of movement planning, but is not restricted to saccade planning.

The time course of attentional bias to cues of threat and safety.

It is well known that relative to neutral stimuli, attention is biased towards processing stimuli that convey threat. In a previous study in which a particular stimulus (e.g. a blue diamond) was associated with the delivery of an electrical shock, the presence of the fear-conditioned stimulus interfered with the execution of voluntary eye movements to other locations. Here, we show that this effect not only occurs early in time, but remains present long after the fear-conditioned stimulus was removed from the screen. In a subsequent experiment, we associated the presence of a particular stimulus with safety, that is, when this stimulus was present it was certain that no electrical shock would be delivered. The presence of the safety signalling stimulus also interfered with the execution of voluntary saccades, but only when the time between stimulus and cue presentation was relatively long. The results indicate that both signals of threat and signals of safety interfere with execution of a saccade long after the source of threat or safety has been removed. However, only threatening stimuli affect saccade execution early in time, suggesting that threatening stimuli drive selection exogenously.

The Role of the Oculomotor System in Updating Visual-Spatial Working Memory across Saccades.

Visual-spatial working memory (VSWM) helps us to maintain and manipulate visual information in the absence of sensory input. It has been proposed that VSWM is an emergent property of the oculomotor system. In the present study we investigated the role of the oculomotor system in updating of spatial working memory representations across saccades. Participants had to maintain a location in memory while making a saccade to a different location. During the saccade the target was displaced, which went unnoticed by the participants. After executing the saccade, participants had to indicate the memorized location. If memory updating fully relies on cancellation driven by extraretinal oculomotor signals, the displacement should have no effect on the perceived location of the memorized stimulus. However, if postsaccadic retinal information about the location of the saccade target is used, the perceived location will be shifted according to the target displacement. As it has been suggested that maintenance of accurate spatial representations across saccades is especially important for action control, we used different ways of reporting the location held in memory; a match-to-sample task, a mouse click or by making another saccade. The results showed a small systematic target displacement bias in all response modalities. Parametric manipulation of the distance between the to-be-memorized stimulus and saccade target revealed that target displacement bias increased over time and changed its spatial profile from being initially centered on locations around the saccade target to becoming spatially global. Taken together results suggest that we neither rely exclusively on extraretinal nor on retinal information in updating working memory representations across saccades. The relative contribution of retinal signals is not fixed but depends on both the time available to integrate these signals as well as the distance between the saccade target and the remembered location.

Distractors associated with reward break through the focus of attention.

In the present study, we investigated the conditions in which rewarded distractors have the ability to capture attention, even when attention is directed toward the target location. Experiment 1 showed that when the probability of obtaining reward was high, all salient distractors captured attention, even when they were not associated with reward. This effect may have been caused by participants suboptimally using the 100%-valid endogenous location cue. Experiment 2 confirmed this result by showing that salient distractors did not capture attention in a block in which no reward was expected. In Experiment 3, the probability of the presence of a distractor was high, but it only signaled reward availability on a low number of trials. The results showed that those very infrequent distractors that signaled reward captured attention, whereas the distractors (both frequent and infrequent ones) not associated with reward were simply ignored. The latter experiment indicates that even when attention is directed to a location in space, stimuli associated with reward break through the focus of attention, but equally salient stimuli not associated with reward do not.

Visual input signaling threat gains preferential access to awareness in a breaking continuous flash suppression paradigm.

Visual input that signals threat is inherently relevant for survival. Accordingly, it has been demonstrated that threatening visual input elicits faster behavioral responses than non-threatening visual input. Considering that awareness is a prerequisite for performing demanding tasks and guiding novel behavior, we hypothesized that threatening visual input would gain faster access to awareness than non-threatening visual input. In the present study, we associated one of two basic visual stimuli, that were devoid of intrinsic relevance (colored annuli), with aversive stimulation (i.e., electric shocks) following a classical fear conditioning procedure. In the subsequent test phase no more electric shocks were delivered, and a breaking continuous flash suppression task was used to measure how fast these stimuli would access awareness. The results reveal that stimuli that were previously paired with an electric shock break through suppression faster than comparable stimuli that were not paired with an electric shock.

The role of context in volitional control of feature-based attention.

Visual selection can be biased toward nonspatial feature values such as color, but there is continued debate about whether this bias is subject to volitional control or whether it is an automatic bias toward recently seen target features (selection history). Although some studies have tried to separate these 2 sources of selection bias, mixed findings have not offered a clear resolution. The present work offers a possible explanation of conflicting findings by showing that the context in which a trial is presented can determine whether volitional control is observed. We used a cueing task that enabled independent assessments of the effects of color repetitions and current selection goals. When the target was presented among distractors with multiple colors (heterogeneous blocks), Experiment 1 revealed clear goal-driven selection effects, but these effects were eliminated when the target was a color singleton (pop-out blocks). When heterogeneous and pop-out displays were mixed within a block (Experiment 2), however, goal-driven selection was observed with both types of displays. In Experiment 3, this pattern was replicated using an encoding-limited task that included brief displays and an A' measure of performance. Thus, goal-driven selection of nonspatial features is potentiated in contexts where there is strong competition with distractors. Selection history has powerful effects, but we find clear evidence that observers can exert volitional control over feature-based attention. (PsycINFO Database Record

The Effects of Feature-Based Priming and Visual Working Memory on Oculomotor Capture.

Recently, it has been demonstrated that objects held in working memory can influence rapid oculomotor selection. This has been taken as evidence that perceptual salience can be modified by active working memory representations. The goal of the present study was to examine whether these results could also be caused by feature-based priming. In two experiments, participants were asked to saccade to a target line segment of a certain orientation that was presented together with a to-be-ignored distractor. Both objects were given a task-irrelevant color that varied per trial. In a secondary task, a color had to be memorized, and that color could either match the color of the target, match the color of the distractor, or it did not match the color of any of the objects in the search task. The memory task was completed either after the search task (Experiment 1), or before it (Experiment 2). The results showed that in both experiments the memorized color biased oculomotor selection. Eye movements were more frequently drawn towards objects that matched the memorized color, irrespective of whether the memory task was completed after (Experiment 1) or before (Experiment 2) the search task. This bias was particularly prevalent in short-latency saccades. The results show that early oculomotor selection performance is not only affected by properties that are actively maintained in working memory but also by those previously memorized. Both working memory and feature priming can cause early biases in oculomotor selection.