Perisaccadic Compression Research Articles

Fixation related shifts of perceptual localization counter to saccade direction.

Perisaccadic compression of the perceived location of flashed visual stimuli toward a saccade target occurs from about 50 ms before a saccade. Here we show that between 150 and 80 ms before a saccade, perceived locations are shifted toward the fixation point. To establish the cause of the "reverse" presaccadic perceptual distortion, participants completed several versions of a saccade task. After a cue to saccade, a probe bar stimulus was briefly presented within the saccade trajectory. In Experiment 1 participants made (a) overlap saccades with immediate return saccades, (b) overlap saccades, and (c) step saccades. In Experiment 2 participants made gap saccades in complete darkness. In Experiment 3 participants maintained fixation with the probe stimuli masked at various interstimulus intervals. Participants indicated the bar's location using a mouse cursor. In all conditions in Experiment 1 presaccadic compression was preceded by compression toward the initial fixation. In Experiment 2, saccadic compression was maintained but the preceding countercompression was not observed. Stimuli masked at fixation were not compressed. This suggests the two opposing compression effects are related to the act of executing an eye movement. They are also not caused by the requirement to make two sequential saccades ending at the initial fixation location and are not caused by continuous presence of the fixation markers. We propose that countercompression is related to fixation activity and is part of the sequence of motor preparations to execute a cued saccade.

Evidence that perisaccadic compression is related to uncertainty about the target's position

Evidence that perisaccadic compression is related to uncertainty about the target's position

Perisaccadic visual perception

Primates use frequent, rapid eye movements to sample their visual environment. This is a fruitful strategy to make the best use of the highly sensitive foveal part of the retina, but it requires neural mechanisms to bind the rapidly changing visual input into a single, stable percept. Studies investigating these neural mechanisms have typically assumed that perisaccadic perception in nonhuman primates matches that of humans. We tested this assumption by performing identical experiments in human and nonhuman primates. Our data confirm that perisaccadic visual perception of macaques and humans is qualitatively similar. Specifically, we found a reduction in detectability and mislocalization of targets presented at the time of saccades. We also found substantial differences between human and nonhuman primates. Notably, in nonhuman primates, localization that requires knowledge of eye position was less precise, nonhuman primates detected fewer perisaccadic stimuli, and perisaccadic compression was not towards the saccade target. The qualitative similarities between species support the view that the nonhuman primate is ideally suited to study aspects of brain function—such as those relying on foveal vision—that are uniquely developed in primates. The quantitative differences, however, demonstrate the need for a reassessment of the models purportedly linking neural response changes at the time of saccades with the behavioral phenomena of perisaccadic reduction of detectability and mislocalization.

A predictive coding model of gaze shifts and the underlying neurophysiology

ABSTRACTA comprehensive model of gaze control must account for a number of empirical observations at both the behavioural and neurophysiological levels. The computational model presented in this article can simulate the coordinated movements of the eye, head, and body required to perform horizontal gaze shifts. In doing so it reproduces the predictable relationships between the movements performed by these different degrees of freedom (DOFs) in the primate. The model also accounts for the saccadic undershoot that accompanies large gaze shifts in the biological visual system. It can also account for our perception of a stable external world despite frequent gaze shifts and the ability to perform accurate memory-guided and double-step saccades. The proposed model also simulates peri-saccadic compression: the mis-localization of a briefly presented visual stimulus towards the location that is the target for a saccade. At the neurophysiological level, the proposed model is consistent with the existence of cortical neurons tuned to the retinal, head-centred, body-centred, and world-centred locations of visual stimuli and cortical neurons that have gain-modulated responses to visual stimuli. Finally, the model also successfully accounts for peri-saccadic receptive field (RF) remapping which results in reduced responses to stimuli in the current RF location and an increased sensitivity to stimuli appearing at the location that will be occupied by the RF after the saccade. The proposed model thus offers a unified explanation for this seemingly diverse range of phenomena. Furthermore, as the proposed model is an implementation of the predictive coding theory, it offers a single computational explanation for these phenomena and relates gaze shifts to a wider framework for understanding cortical function.

Does memory affect perisaccadic compression?

Does memory affect perisaccadic compression?

Compression of Space for Low Visibility Probes.

Stimuli briefly flashed just before a saccade are perceived closer to the saccade target, a phenomenon known as perisaccadic compression of space (Ross et al., 1997). More recently, we have demonstrated that brief probes are attracted towards a visual reference when followed by a mask, even in the absence of saccades (Zimmermann et al., 2014a). Here, we ask whether spatial compression depends on the transient disruptions of the visual input stream caused by either a mask or a saccade. Both of these degrade the probe visibility but we show that low probe visibility alone causes compression in the absence of any disruption. In a first experiment, we varied the regions of the screen covered by a transient mask, including areas where no stimulus was presented and a condition without masking. In all conditions, we adjusted probe contrast to make the probe equally hard to detect. Compression effects were found in all conditions. To obtain compression without a mask, the probe had to be presented at much lower contrasts than with masking. Comparing mislocalizations at different probe detection rates across masking, saccades and low contrast conditions without mask or saccade, Experiment 2 confirmed this observation and showed a strong influence of probe contrast on compression. Finally, in Experiment 3, we found that compression decreased as probe duration increased both for masks and saccades although here we did find some evidence that factors other than simply visibility as we measured it contribute to compression. Our experiments suggest that compression reflects how the visual system localizes weak targets in the context of highly visible stimuli.

Interference during eye movement preparation shifts the timing of perisaccadic compression.

Our perception of the surrounding environment remains stable despite the fact that we frequently change the retinal position of input by rapid gaze shifts (saccades). There is a long-standing debate whether visual stability depends on an active mechanism using an efference copy of the impending saccadic motor command. Behavioral studies showing changes in perception around the time of saccades are consistent with a predictive mechanism, but previous studies of perceptual effects in humans confounded saccade programming with the resulting physical eye movement. In three experiments, we used a saccadic inhibition (SI) paradigm to delay saccadic onset while participants were performing a perisaccadic localization task. As expected, the perceived position of the probe stimulus was systematically biased (compressed) toward the saccadic goal, already during the presaccadic interval. In the SI condition, the localization error was shifted in time, in line with it following saccade intention rather than execution. The pattern was not the consequence of the probe being captured by the timing of the flashed distractor, but depended instead on the delay in saccadic onset time caused by SI. Importantly, the same configurations of perceptual probes presented with a flashed backward mask when participants maintained fixation did not lead to similar localization errors as saccade trials. This pattern of results is consistent with an active, sensorimotor explanation for perisaccadic mislocalization and, more generally, theories emphasizing the role of motor prediction in visual stability.

Visual mislocalization during double-step saccades.

Visual objects presented briefly at the time of saccade onset appear compressed toward the saccade target. Compression strength depends on the presentation of a visual saccade target signal and is strongly reduced during the second saccade of a double-step saccade sequence (Zimmermann et al., 2014b). Here, I tested whether perisaccadic compression is linked to saccade planning by contrasting two double-step paradigms. In the same-direction double-step paradigm, subjects were required to perform two rightward 10° saccades successively. At various times around execution of the saccade sequence a probe dot was briefly flashed. Subjects had to localize the position of the probe dot after they had completed both saccades. I found compression of visual space only at the time of the first but not at the time of the second saccade. In the reverse-direction paradigm, subjects performed first a rightward 10° saccade followed by a leftward 10° saccade back to initial fixation. In this paradigm compression was found in similar magnitude during both saccades. Analysis of the saccade parameters did not reveal indications of saccade sequence preplanning in this paradigm. I therefore conclude that saccade planning, rather than saccade execution factors, is involved in perisaccadic compression.

Peri-saccadic compression to two locations in a two-target choice saccade task.

When visual stimuli are presented at the onset of a saccadic eye movement they are seen compressed onto the target location of the saccade. This peri-saccadic compression is believed to result from internal feedback pathways between oculomotor and visual areas of the brain. This feedback enhances vision around the saccade target at the expense of localization ability in other regions of the visual field. Although saccades can be targeted at only one object at a time, often multiple potential targets are available in a visual scene, and the oculomotor system has to choose which target to look at. If two targets are available, preparatory activity builds-up at both target locations in oculomotor maps. Here we show that, in this situation, two foci of compression develop, independent of which of the two targets is eventually chosen for the saccade. Our results suggest that theories that use oculomotor feedback as efference copy signals for upcoming eye movements should take the possibility into account that multiple feedback signals from potential targets may occur in parallel before the execution of a saccade.

Effect of saccade automaticity on perisaccadic space compression.

Briefly presented stimuli occurring just before or during a saccadic eye movement are mislocalized, leading to a compression of visual space toward the target of the saccade. In most cases this has been measured in subjects over-trained to perform a stereotyped and unnatural task where saccades are repeatedly driven to the same location, marked by a highly salient abrupt onset. Here, we asked to what extent the pattern of perisaccadic mislocalization depends on this specific context. We addressed this question by studying perisaccadic localization in a set of participants with no prior experience in eye-movement research, measuring localization performance as they practiced the saccade task. Localization was marginally affected by practice over the course of the experiment and it was indistinguishable from the performance of expert observers. The mislocalization also remained similar when the expert observers were tested in a condition leading to less stereotypical saccadic behavior—with no abrupt onset marking the saccade target location. These results indicate that perisaccadic compression is a robust behavior, insensitive to the specific paradigm used to drive saccades and to the level of practice with the saccade task.

The visual component to saccadic compression.

Visual objects presented around the time of saccadic eye movements are strongly mislocalized towards the saccadic target, a phenomenon known as "saccadic compression." Here we show that perisaccadic compression is modulated by the presence of a visual saccadic target. When subjects saccaded to the center of the screen with no visible target, perisaccadic localization was more veridical than when tested with a target. Presenting a saccadic target sometime before saccade initiation was sufficient to induce mislocalization. When we systematically varied the onset of the saccade target, we found that it had to be presented around 100 ms before saccade execution to cause strong mislocalization: saccadic targets presented after this time caused progressively less mislocalization. When subjects made a saccade to screen center with a reference object placed at various positions, mislocalization was focused towards the position of the reference object. The results suggest that saccadic compression is a signature of a mechanism attempting to match objects seen before the saccade with those seen after.

Moving your head reduces perisaccadic compression.

Flashes presented around the time of a saccade appear to be closer to the saccade endpoint than they really are. The resulting compression of perceived positions has been found to increase with the amplitude of the saccade. In most studies on perisaccadic compression the head is static, so the eye-in-head movement is equal to the change in gaze. What if moving the head causes part of the change in gaze? Does decreasing the eye-in-head rotation by moving the head decrease the compression of perceived positions? To find out, we asked participants to shift their gaze between two positions, either without moving their head or with the head contributing to the change in gaze. Around the time of the saccades we flashed bars that participants had to localize. When the head contributed to the change in gaze, the duration of the saccade was shorter and compression was reduced. We interpret this reduction in compression as being caused by a reduction in uncertainty about gaze position at the time of the flash. We conclude that moving one's head can reduce the systematic mislocalization of flashes presented around the time of saccades.

Moving your head reduces perisaccadic compression

Stimuli that are presented briefly near the time of a saccadic gaze shift tend to be judged to have been closer to where the gaze ended than they really were. The resulting perisaccadic compression of perceived positions is probably related to uncertainty about the direction of gaze at the time of the flash. The larger the shift in gaze the stronger the compression, possibly because the uncertainty increases with the magnitude of the motor commands that give rise to the change in gaze. If so, then performing part of the gaze shift by moving one's head may decrease the compression, because the combined uncertainty of eye and head orientation in a combined eye-head gaze shift will be smaller than the uncertainty of the eye when it shifts gaze on its own. To test whether reducing eye-in-head movement by moving the head reduces perisaccadic compression for a given shift in gaze, we asked participants to shift their gaze between two positions, either without moving their head or with the head contributing to the change in gaze. We flashed targets around the time of the saccades and participants indicated where they saw these targets. Although allowing head movements did not affect the amplitude and velocity of the gaze shifts, the eye-in-head movements and their peak velocities were obviously larger when only the eyes moved, than when both the eyes and the head contributed to the change in gaze. There was also significantly more compression when only the eyes moved than when both the eyes and head moved. We conclude that moving one's head can reduce the magnitude of the systematic mislocalization that is observed near the time of rapid shifts of gaze. Meeting abstract presented at VSS 2014

The effect of perceptual grouping on perisaccadic spatial distortions

Perisaccadic spatial distortion (PSD) occurs when a target is flashed immediately before the onset of a saccade and it appears displaced in the direction of the saccade. In previous studies, the magnitude of PSD of a single target was affected by multiple experimental parameters, such as the target's luminance and its position relative to the central fixation target. Here we describe a contextual effect in which the magnitude of the PSD for a target was influenced by the synchronous presentation of another target: PSD for simultaneously presented targets was more uniform than when each was presented individually. Perisaccadic compression was ruled out as a causal factor, and the results suggest that both low- and high-level perceptual grouping mechanisms may account for the change in PSD magnitude. We speculate that perceptual grouping could play a key role in preserving shape constancy during saccadic eye movements.

Perisaccadic compression in two-saccade sequences

Around the onset of a saccade toward a target, localization judgments are systematically biased toward the saccade endpoint. This perisaccadic compression is thought to be related to transsaccadic reorganization and due to interfering motor signals in visual maps. It has, however, only been investigated for saccades targeting a single target. Here, we examined whether saccade-sequence programming to stationary target(s) would affect the pattern of localization judgments of a briefly flashed stimulus. We presented saccade targets that could induce either a single saccade or two-saccade sequences and we flashed a bar around saccade onsets. For all two-saccade-sequence conditions, we showed that localization judgments of the stimulus flashed around the first saccade onset are biased toward an intermediate position between the two-saccade landing position, indicating the influence of the second saccade's parallel planning. This implies that motor-planning signals are most likely responsible for perisaccadic compression than motor execution feedback.

Anticipatory Saccade Target Processing and the Presaccadic Transfer of Visual Features

As we shift our gaze to explore the visual world, information enters cortex in a sequence of successive snapshots, interrupted by phases of blur. Our experience, in contrast, appears like a movie of a continuous stream of objects embedded in a stable world. This perception of stability across eye movements has been linked to changes in spatial sensitivity of visual neurons anticipating the upcoming saccade, often referred to as shifting receptive fields (Duhamel et al., 1992; Walker et al., 1995; Umeno and Goldberg, 1997; Nakamura and Colby, 2002). How exactly these receptive field dynamics contribute to perceptual stability is currently not clear. Anticipatory receptive field shifts toward the future, postsaccadic position may bridge the transient perisaccadic epoch (Sommer and Wurtz, 2006; Wurtz, 2008; Melcher and Colby, 2008). Alternatively, a presaccadic shift of receptive fields toward the saccade target area (Tolias et al., 2001) may serve to focus visual resources onto the most relevant objects in the postsaccadic scene (Hamker et al., 2008). In this view, shifts of feature detectors serve to facilitate the processing of the peripheral visual content before it is foveated. While this conception is consistent with previous observations on receptive field dynamics and on perisaccadic compression (Ross et al., 1997; Morrone et al., 1997; Kaiser and Lappe, 2004), it predicts that receptive fields beyond the saccade target shift toward the saccade target rather than in the direction of the saccade. We have tested this prediction in human observers via the presaccadic transfer of the tilt-aftereffect (Melcher, 2007).

The spatial pattern of peri-saccadic compression for small saccades

In the temporal vicinity of a saccade onset, visual stability is transiently disrupted and briefly flashed visual stimuli undergo a systematic perceptual mislocalization. Specifically, when a stimulus is flashed around saccade onset, localization judgments are grossly biased toward the saccade endpoint. This peri-saccadic compression increases with saccade amplitude. Previous studies of peri-saccadic compression have typically used rather large saccade amplitudes. In the present study, we investigate systematically the pattern of errors for small saccade sizes (2°-10°), taking into account both the amplitude of the saccade and the position of the flashed stimulus (11 positions tested from 1 to 12°). Our results show a weaker compression effect for the smallest saccades. Moreover, we found that the strength of the compression depends on both stimulus side and relative distance to the saccade target.

Perisaccadic visual compression shown by target-flash mislocalization may be affected by flash visual persistence interacting with background stimuli

Perisaccadic visual compression shown by target-flash mislocalization may be affected by flash visual persistence interacting with background stimuli

Perisaccadic compression of space orthogonal to saccade direction

Perisaccadic compression of space orthogonal to saccade direction

The spatial distribution of receptive field changes in a model of peri-saccadic perception: Predictive remapping and shifts towards the saccade target

At the time of an impending saccade receptive fields (RFs) undergo dynamic changes, that is, their spatial profile is altered. This phenomenon has been observed in several monkey visual areas. Although their link to eye movements is obvious, neither the exact pattern nor their function is fully clear. Several RF shifts have been interpreted in terms of predictive remapping mediating visual stability. In particular, even prior to saccade onset some cells become responsive to stimuli presented in their future, post-saccadic RF. In visual area V4, however, the overall effect of RF dynamics consists of a shrinkage and shift of RFs towards the saccade target. These observations have been linked to a pre-saccadically enhanced processing of the future fixation. In order to better understand these seemingly different outcomes, we analyzed the RF shifts predicted by a recently proposed computational model of peri-saccadic perception ( Hamker, Zirnsak, Calow, & Lappe, 2008). This model unifies peri-saccadic compression, pre-saccadic attention shifts, and peri-saccadic receptive field dynamics in a common framework of oculomotor reentry signals in extrastriate visual cortical maps. According to the simulations that we present in the current paper, a spatially selective oculomotor feedback signal leads to RF dynamics which are both consistent with the observations made in studies aiming to investigate predictive remapping and saccade target shifts. Thus, the seemingly distinct experimental observations could be grounded in the same neural mechanism leading to different RF dynamics dependent on the location of the RF in visual space.