Saccadic Suppression of Displacement Does Not Reflect a Saccade-Specific Bias to Assume Stability.
ABSTRACT: Across saccades, small displacements of a visual target are harder to detect and their directions more difficult to discriminate than during steady fixation. Prominent theories of this effect, known as saccadic suppression of displacement, propose that it is due to a bias to assume object stability across saccades. Recent studies comparing the saccadic effect to masking effects suggest that suppression of displacement is not saccade-specific. Further evidence for this account is presented from two experiments where participants judged the size of displacements on a continuous scale in saccade and mask conditions, with and without blanking. Saccades and masks both reduced the proportion of correctly perceived displacements and increased the proportion of missed displacements. Blanking improved performance in both conditions by reducing the proportion of missed displacements. Thus, if suppression of displacement reflects a bias for stability, it is not a saccade-specific bias, but a more general stability assumption revealed under conditions of impoverished vision. Specifically, I discuss the potentially decisive role of motion or other transient signals for displacement perception. Without transients or motion, the quality of relative position signals is poor, and saccadic and mask-induced suppression of displacement reflects performance when the decision has to be made on these signals alone. Blanking may improve those position signals by providing a transient onset or a longer time to encode the pre-saccadic target position.
Project description:To establish a perceptually stable world despite the large retinal shifts caused by saccadic eye movements, the visual system reduces its sensitivity to the displacement of visual stimuli during saccades (e.g. saccadic suppression of displacement, SSD). Previous studies have demonstrated that inserting a temporal blank right after a saccade improves displacement detection performance. This 'blanking effect' suggests that visual information right after the saccade may play an important role in SSD. To understand the mechanisms underlying SSD, we here compare the effect of pre- and post-saccadic stimulus contrast on displacement detection during a saccade with and without inserting a blank. Our results show that observers' sensitivity to detect visual displacement was reduced by increasing post-saccadic stimulus contrast, but a blank relieves the impairment. We successfully explain the results with a model proposing that parvo-pathway signals suppress the magno-pathway processes responsible for detecting displacements across saccades. Our results suggest that the suppression of the magno-pathway by parvo-pathway signals immediately after a saccade causes SSD, which helps to achieve the perceptual stability of the visual world across saccades.
Project description:Humans do not notice small displacements to objects that occur during saccades, termed saccadic suppression of displacement (SSD), and this effect is reduced when a blank is introduced between the pre- and postsaccadic stimulus (Bridgeman, Hendry, & Stark, 1975; Deubel, Schneider, & Bridgeman, 1996). While these effects have been studied extensively in adults, it is unclear how these phenomena are characterized in children. A potentially related mechanism, saccadic suppression of contrast sensitivity-a prerequisite to achieve a stable percept-is stronger for children (Bruno, Brambati, Perani, & Morrone, 2006). However, the evidence for how transsaccadic stimulus displacements may be suppressed or integrated is mixed. While they can integrate basic visual feature information from an early age, they cannot integrate multisensory information (Gori, Viva, Sandini, & Burr, 2008; Nardini, Jones, Bedford, & Braddick, 2008), suggesting a failure in the ability to integrate more complex sensory information. We tested children 7 to 12 years old and adults 19 to 23 years old on their ability to perceive intrasaccadic stimulus displacements, with and without a postsaccadic blank. Results showed that children had stronger SSD than adults and a larger blanking effect. Children also had larger undershoots and more variability in their initial saccade endpoints, indicating greater intrinsic uncertainty, and they were faster in executing corrective saccades to account for these errors. Together, these results suggest that children may have a greater internal expectation or prediction of saccade error than adults; thus, the stronger SSD in children may be due to higher intrinsic uncertainty in target localization or saccade execution.
Project description:Although our eyes are in constant movement, we remain unaware of the high-speed stimulation produced by the retinal displacement. Vision is drastically reduced at the time of saccades. Here, I investigated whether the reduction of the unwanted disturbance could be established through a saccade-contingent habituation to intra-saccadic displacements. In more than 100 context trials, participants were exposed either to an intra-saccadic or to a post-saccadic disturbance or to no disturbance at all. After induction of a specific context, I measured peri-saccadic suppression. Displacement discrimination thresholds of observers were high after participants were exposed to an intra-saccadic disturbance. However, after exposure to a post-saccadic disturbance or a context without any intra-saccadic stimulation, displacement discrimination improved such that observers were able to see shifts as during fixation. Saccade-contingent habituation might explain why we do not perceive trans-saccadic retinal stimulation during saccades.
Project description:Visual perception is introspectively stable and continuous across eye movements. It has been hypothesized that displacements in retinal input caused by eye movements can be dissociated from displacements in the external world using extra-retinal information, such as a corollary discharge from the oculomotor system. The extra-retinal information can inform the visual system about an upcoming eye movement and accompanying displacements in retinal input. The parietal cortex has been hypothesized to be critically involved in integrating retinal and extra-retinal information. Two tasks have been widely used to assess the quality of this integration: double-step saccades and intra-saccadic displacements. Double-step saccades performed by patients with parietal cortex lesions seemed to show hypometric second saccades. However, recently idea has been refuted by demonstrating that patients with very similar lesions were able to perform the double step saccades, albeit taking multiple saccades to reach the saccade target. So, it seems that extra-retinal information is still available for saccade execution after a lesion to the parietal lobe. Here, we investigated whether extra-retinal signals are also available for perceptual judgements in nine patients with strokes affecting the posterior parietal cortex. We assessed perceptual continuity with the intra-saccadic displacement task. We exploited the increased sensitivity when a small temporal blank is introduced after saccade offset (blank effect). The blank effect is thought to reflect the availability of extra-retinal signals for perceptual judgements. Although patients exhibited a relative difference to control subjects, they still demonstrated the blank effect. The data suggest that a lesion to the posterior parietal cortex (PPC) alters the processing of extra-retinal signals but does not abolish their influence altogether.
Project description:Corollary discharge (CD) signals are "copies" of motor signals sent to sensory regions that allow animals to adjust sensory consequences of self-generated actions. Autism spectrum disorder (ASD) is characterized by sensory and motor deficits, which may be underpinned by altered CD signaling. We evaluated oculomotor CD using the blanking task, which measures the influence of saccades on visual perception, in 30 children with ASD and 35 typically developing (TD) children. Participants were instructed to make a saccade to a visual target. Upon saccade initiation, the presaccadic target disappeared and reappeared to the left or right of the original position. Participants indicated the direction of the jump. With intact CD, participants can make accurate perceptual judgements. Otherwise, participants may use saccade landing site as a proxy of the presaccadic target and use it to inform perception. We used multilevel modeling to examine the influence of saccade landing site on trans-saccadic perceptual judgements. We found that, compared with TD participants, children with ASD were more sensitive to target displacement and less reliant on saccade landing site when spatial uncertainty of the post-saccadic target was high. This pattern was driven by ASD participants with less severe restricted and repetitive behaviors. These results suggest a relationship between altered CD signaling and core ASD symptoms.
Project description:Saccades are rapid eye movements that change the direction of gaze, although the full-field image motion associated with these movements is rarely perceived. The attenuation of visual perception during saccades is referred to as saccadic suppression. The mechanisms that produce saccadic suppression are not well understood. We recorded from neurons in the dorsal medial superior temporal area (MSTd) of alert macaque monkeys and compared the neural responses produced by the retinal slip associated with saccades (active motion) to responses evoked by identical motion presented during fixation (passive motion). We provide evidence for a neural correlate of saccadic suppression and expand on two contentious results from previous studies. First, we confirm the finding that some neurons in MSTd reverse their preferred direction during saccades. We quantify this effect by calculating changes in direction tuning index for a large cell population. Second, it has been noted that neural activity associated with saccades can arrive in the parietal cortex <or=30 ms earlier than activity produced by similar visual stimulation during fixation. This led to the question of whether the saccade-related responses were visual in origin or were motor signals arising from saccade-planning areas of the brain. By comparing the responses to saccades made over textured backgrounds of different contrasts, we provide strong evidence that saccade-related responses were visual in origin. Refinements of the possible models of saccadic suppression are discussed.
Project description:We generally experience a stable visual world in spite of regular disruptions caused by our own movements (saccades, blinks) or by the visual input itself (flashes, occlusions). In trying to understand the mechanisms responsible for this stability, saccades have been particularly well-studied, and a number of peri-saccadic perceptual distortions (spatial and temporal compression, failure to detect target displacement) have been explored. It has been shown that some of these distortions are not saccade specific, but also arise when the visual input is instead abruptly and briefly masked. Here, we demonstrate that another peri-saccadic distortion, the reversal of the temporal order of a pair of brief events, may also be found with masking. Human participants performed a temporal order judgment task, and the timing of stimuli and mask was varied over trials. Perceptual order was reversed on ~25% of the trials at the shortest stimulus to mask intervals. This was not merely a failure of target detection, since participants often reported these reversals with high subjective confidence. These findings update the constraints on models of stability around disruptions.
Project description: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.
Project description:Visual sensitivity, probed through perceptual detectability of very brief visual stimuli, is strongly impaired around the time of rapid eye movements. This robust perceptual phenomenon, called saccadic suppression, is frequently attributed to active suppressive signals that are directly derived from eye movement commands. Here we show instead that visual-only mechanisms, activated by saccade-induced image shifts, can account for all perceptual properties of saccadic suppression that we have investigated. Such mechanisms start at, but are not necessarily exclusive to, the very first stage of visual processing in the brain, the retina. Critically, neural suppression originating in the retina outlasts perceptual suppression around the time of saccades, suggesting that extra-retinal movement-related signals, rather than causing suppression, may instead act to shorten it. Our results demonstrate a far-reaching contribution of visual processing mechanisms to perceptual saccadic suppression, starting in the retina, without the need to invoke explicit motor-based suppression commands.
Project description:The understanding of the subjective experience of a visually stable world despite the occurrence of an observer's eye movements has been the focus of extensive research for over 20 years. These studies have revealed fundamental mechanisms such as anticipatory receptive field (RF) shifts and the saccadic suppression of stimulus displacements, yet there currently exists no single explanatory framework for these observations. We show that a previously presented neuro-computational model of peri-saccadic mislocalization accounts for the phenomenon of predictive remapping and for the observation of saccadic suppression of displacement (SSD). This converging evidence allows us to identify the potential ingredients of perceptual stability that generalize beyond different data sets in a formal physiology-based model. In particular we propose that predictive remapping stabilizes the visual world across saccades by introducing a feedback loop and, as an emergent result, small displacements of stimuli are not noticed by the visual system. The model provides a link from neural dynamics, to neural mechanism and finally to behavior, and thus offers a testable comprehensive framework of visual stability.