Saccade Landing Position Research Articles

Saccades to partially occluded objects: Perceptual completion mediates oculomotor control.

Oculomotor behavior typically consists of directing gaze to objects in complex scenes for the purpose of extracting detailed perceptual information. Here, we probed the nature of the visual representations over which saccades to objects are computed. We contrasted an image-based oculomotor control hypothesis, holding that saccades are computed solely over information explicit in the retinal image, and an object-based oculomotor control hypothesis, holding that saccades are computed over object representations reflecting the three-dimensional structure of the scene. We recorded saccade landing positions to partially occluded objects in a naturalistic search task. In Experiment 1, saccade landing positions were biased toward the center of the perceptually completed object. Experiment 2 demonstrated that the bias held even when it would have been strategically advantageous to avoid it. Experiment 3 demonstrated that the bias was not due to image-level differences generated by the presence of occluders. The results indicate that saccade motor programs are computed, at least in part, over object-level representations reflecting the completion of occluded surfaces.

Eye movements are made to the centre of gravity of texture-defined targets

Saccadic localisation of targets of various properties has been extensively studied, but rarely for texture-defined figures. In this paper, three experiments that investigate the way information from a texture target is processed in order to provide a signal for eye movement control are presented. Participants made saccades to target regions embedded in a background structure, and the saccade landing position and latency were measured. The textures comprised line elements, with orientations of the lines configured to form the figure and ground. Various orientation profile configurations (Block, Blur, and Cornsweet), were used in order to measure the role of edge profiles in driving eye movements and producing salience. We found that in all cases the visual system is in fact able to effectively segregate a texture figure from the ground in order to accurately plan a saccade to the target-figure. While saccadic latency was the highest for the Blur profile, the mean saccadic landing position was mostly unaffected by the various profiles (Experiment 1). More specifically, we showed that saccades were directed to the centre-of-gravity of the target (Experiment 2). We also found that figures with information of orientation contrast at both the edge and centre of figure (i.e. Block) produced the highest level of saliency in attracting eye movements (Experiment 3). Overall, the results show that saccades are planned on the representation of the whole target shape rather than a local salient region based on orientation contrast cues, and that the various texture profiles were important only to the extent that they affected the time to programme a saccade.

Practical Saccade Prediction for Head-Mounted Displays: Towards a Comprehensive Model

Eye-tracking technology has started to become an integral component of new display devices such as virtual and augmented reality headsets. Applications of gaze information range from new interaction techniques that exploit eye patterns to gaze-contingent digital content creation. However, system latency is still a significant issue in many of these applications because it breaks the synchronization between the current and measured gaze positions. Consequently, it may lead to unwanted visual artifacts and degradation of the user experience. In this work, we focus on foveated rendering applications where the quality of an image is reduced towards the periphery for computational savings. In foveated rendering, the presence of system latency leads to delayed updates to the rendered frame, making the quality degradation visible to the user. To address this issue and to combat system latency, recent work proposes using saccade landing position prediction to extrapolate gaze information from delayed eye tracking samples. Although the benefits of such a strategy have already been demonstrated, the solutions range from simple and efficient ones, which make several assumptions about the saccadic eye movements, to more complex and costly ones, which use machine learning techniques. However, it is unclear to what extent the prediction can benefit from accounting for additional factors and how more complex predictions can be performed efficiently to respect the latency requirements. This paper presents a series of experiments investigating the importance of different factors for saccades prediction in common virtual and augmented reality applications. In particular, we investigate the effects of saccade orientation in 3D space and smooth pursuit eye-motion (SPEM) and how their influence compares to the variability across users. We also present a simple, yet efficient post-hoc correction method that adapts existing saccade prediction methods to handle these factors without performing extensive data collection. Furthermore, our investigation and the correction technique may also help future developments of machine-learning-based techniques by limiting the required amount of training data.

Saccadic landing positions reveal that eye movements are affected by distractor-based retrieval

Binding theories assume that stimulus and response features are integrated into short-lasting episodes and that upon repetition of any feature the whole episode is retrieved, thereby affecting performance. Such binding theories are nowadays the standard explanation for a wide range of action control tasks and aim to explain all simple actions, without making assumptions of effector specificity. Yet, it is unclear if eye movements are affected by integration and retrieval in the same way as manual responses. We asked participants to discriminate letters framed by irrelevant shapes. In Experiment 1, participants gave their responses with eye movements. Saccade landing positions showed a spatial error pattern consistent with predictions of binding theories. Saccadic latencies were not affected. In Experiment 2 with an increased interval between prime and probe, the error pattern diminished, again congruent with predictions of binding theories presuming quickly decaying retrieval effects. Experiment 3 used the same task as in Experiment 1, but participants executed their responses with manual key presses; again, we found a binding pattern in response accuracy. We conclude that eye movements and manual responses are affected by the same integration and retrieval processes, supporting the tacit assumption of binding theories to apply to any effector.

Characteristic fixation biases in Super-Recognizers.

Neurotypical observers show large and reliable individual differences in gaze behavior along several semantic object dimensions. Individual gaze behavior toward faces has been linked to face identity processing, including that of neurotypical observers. Here, we investigated potential gaze biases in Super-Recognizers (SRs), individuals with exceptional face identity processing skills. Ten SRs, identified with a novel conservative diagnostic framework, and 43 controls freely viewed 700 complex scenes depicting more than 5000 objects. First, we tested whether SRs and controls differ in fixation biases along four semantic dimensions: faces, text, objects being touched, and bodies. Second, we tested potential group differences in fixation biases toward eyes and mouths. Finally, we tested whether SRs fixate closer to the theoretical optimal fixation point for face identification. SRs showed a stronger gaze bias toward faces and away from text and touched objects, starting from the first fixation onward. Further, SRs spent a significantly smaller proportion of first fixations and dwell time toward faces on mouths but did not differ in dwell time or first fixations devoted to eyes. Face fixation of SRs also fell significantly closer to the theoretical optimal fixation point for identification, just below the eyes. Our findings suggest that reliable superiority for face identity processing is accompanied by early fixation biases toward faces and preferred saccadic landing positions close to the theoretical optimum for face identification. We discuss future directions to investigate the functional basis of individual fixation behavior and face identity processing ability.

Understanding the collinear masking effect in visual search through eye tracking.

Recent research has reported that, while both orientation contrast and collinearity increase target salience in visual search, a combination of the two counterintuitively masks a local target. Through eye-tracking and eye-movement analysis with hidden Markov models (EMHMM), here we showed that this collinear masking effect was associated with reduced eye-fixation consistency (as measured in entropy) at the central fixation cross prior to the search display presentation. As a decreased precision of saccade landing position is shown to be related to attention shift away from the saccadic target, our result suggested that the collinear masking effect may be related to attention shift to a non-saccadic-goal location in expectation of the search display before saccading to the central fixation cross. This attention shift may consequently interfere with attention capture by the collinear distractor containing the target, resulting in the masking effect. In contrast, although older adults had longer response times, more dispersed eye-movement pattern, and lower eye-movement consistency than young adults during visual search, the two age groups did not differ in the masking effect, suggesting limited contribution from ageing-related cognitive decline. Thus, participants' pre-saccadic attention shift prior to search may be an important factor influencing their search behavior.

Eye-movement patterns to social and non-social cues in early deaf adults.

Previous research on covert orienting to the periphery suggested that early profound deaf adults were less susceptible to uninformative gaze-cues, though were equally or more affected by non-social arrow-cues. The aim of this work was to investigate whether spontaneous eye movement behaviour helps explain the reduced impact of the social cue in deaf adults. We tracked the gaze of 25 early profound deaf and 25 age-matched hearing observers performing a peripheral discrimination task with uninformative central cues (gaze vs arrow), stimulus-onset asynchrony (250 vs 750 ms), and cue validity (valid vs invalid) as within-subject factors. In both groups, the cue effect on reaction time (RT) was comparable for the two cues, although deaf observers responded significantly slower than hearing controls. While deaf and hearing observers' eye movement pattern looked similar when the cue was presented in isolation, deaf participants made significantly more eye movements than hearing controls once the discrimination target appeared. Notably, further analysis of eye movements in the deaf group revealed that independent of the cue type, cue validity affected saccade landing position, while latency was not modulated by these factors. Saccade landing position was also strongly related to the magnitude of the validity effect on RT, such that the greater the difference in saccade landing position between invalid and valid trials, the greater the difference in manual RT between invalid and valid trials. This work suggests that the contribution of overt selection in central cueing of attention is more prominent in deaf adults and helps determine the manual performance, irrespective of the cue type.

Using eye movements to detect visual field loss: a pragmatic assessment using simulated scotoma

Glaucoma is a leading cause of irreversible sight-loss and has been shown to affect natural eye-movements. These changes may provide a cheap and easy-to-obtain biomarker for improving disease detection. Here, we investigated whether these changes are large enough to be clinically useful. We used a gaze-contingent simulated visual field (VF) loss paradigm, in which participants experienced a variable magnitude of simulated VF loss based on longitudinal data from a real glaucoma patient (thereby controlling for other variables, such as age and general health). Fifty-five young participants with healthy vision were asked to view two short videos and three pictures, either with: (1) no VF loss, (2) moderate VF loss, or (3) advanced VF loss. Eye-movements were recorded using a remote eye tracker. Key eye-movement parameters were computed, including saccade amplitude, the spread of saccade endpoints (bivariate contour ellipse area), location of saccade landing positions, and similarity of fixations locations among participants (quantified using kernel density estimation). The simulated VF loss caused some statistically significant effects in the eye movement parameters. Yet, these effects were not capable of consistently identifying simulated VF loss, despite it being of a magnitude likely easily detectable by standard automated perimetry.

The parallel programming of landing position in saccadic eye movement sequences.

Saccadic eye movements occur in sequences, gathering new information about the visual environment to support successful task completion. Here, we examine the control of these saccadic sequences and specifically the extent to which the spatial aspects of the saccadic responses are programmed in parallel. We asked participants to saccade to a series of visual targets and, while they shifted their gaze around the display, we displaced select targets. We found that saccade landing position was deviated toward the previous location of the target suggesting that partial parallel programming of target location information was occurring. The saccade landing position was also affected by the new target location, which demonstrates that the saccade landing position was also partially updated following the shift. This pattern was present even for targets that were the subject of the next fixation. Having a greater preview about the sequence path influenced saccade accuracy with saccades being less affected by relocations when there is less preview information. The results demonstrate that landing positions from a saccade sequence are programmed in parallel and combined with more immediate visual signals.

Gaze direction as equilibrium: more evidence from spatial and temporal aspects of small-saccade triggering in the rhesus macaque monkey.

Rigorous behavioral studies made in human subjects have shown that small-eccentricity target displacements are associated with increased saccadic reaction times, but the reasons for this remain unclear. Before characterizing the neurophysiological foundations underlying this relationship between the spatial and temporal aspects of saccades, we tested the triggering of small saccades in the male rhesus macaque monkey. We also compared our results to those obtained in human subjects, both from the existing literature and through our own additional measurements. Using a variety of behavioral tasks exercising visual and nonvisual guidance of small saccades, we found that small saccades consistently require more time than larger saccades to be triggered in the nonhuman primate, even in the absence of any visual guidance and when valid advance information about the saccade landing position is available. We also found a strong asymmetry in the reaction times of small upper versus lower visual field visually guided saccades, a phenomenon that has not been described before for small saccades, even in humans. Following the suggestion that an eye movement is not initiated as long as the visuo-oculomotor system is within a state of balance, in which opposing commands counterbalance each other, we propose that the longer reaction times are a signature of enhanced times needed to create the symmetry-breaking condition that puts downstream premotor neurons into a push-pull regime necessary for rotating the eyeballs. Our results provide an important catalog of nonhuman primate oculomotor capabilities on the miniature scale, allowing concrete predictions on underlying neurophysiological mechanisms.NEW & NOTEWORTHY Leveraging a multitude of neurophysiological investigations in the rhesus macaque monkey, we generated and tested hypotheses about small-saccade latencies in this animal model. We found that small saccades always take longer, on average, than larger saccades to trigger, regardless of visual and cognitive context. Moreover, small downward saccades have the longest latencies overall. Our results provide an important documentation of oculomotor capabilities of an indispensable animal model for neuroscientific research in vision, cognition, and action.

Experimental test of Bayesian saccade targeting under reversed reading direction.

During reading, rapid eye movements (saccades) shift the reader's line of sight from one word to another for high-acuity visual information processing. While experimental data and theoretical models show that readers aim at word centers, the eye-movement (oculomotor) accuracy is low compared to other tasks. As a consequence, distributions of saccadic landing positions indicate large (i) random errors and (ii) systematic over- and undershoot of word centers, which additionally depend on saccade lengths (McConkie et al. Visual Research, 28(10), 1107-1118, 1988). Here we show that both error components can be simultaneously reduced by reading texts from right to left in German language (N = 32). We used our experimental data to test a Bayesian model of saccade planning. First, experimental data are consistent with the model. Second, the model makes specific predictions of the effects of the precision of prior and (sensory) likelihood. Our results suggest that it is a more precise sensory likelihood that can explain the reduction of both random and systematic error components.

Dynamic combination of position and motion information when tracking moving targets.

To accurately foveate a moving target, the oculomotor system needs to estimate the position of the target at the saccade end, based on information about its position and ongoing movement, while accounting for neuronal delays and execution time of the saccade. We investigated human interceptive saccades and pursuit responses to moving targets defined by high and low luminance contrast or by chromatic contrast only (isoluminance). We used step-ramps with perpendicular directions between vertical target steps of 10 deg/s and horizontal ramps of 2.5 to 20 deg/s to separate errors with respect to the position step of the target in the vertical dimension, and errors related to target motion in the horizontal dimension. Interceptive saccades to targets of high and low luminance contrast landed close to the actual target positions, suggesting relatively accurate estimates of the amount of target displacement. Interceptive saccades to isoluminant targets were less accurate. They landed at positions the target had on average 100 ms before saccade onset. One account of this finding is that the integration of target motion is compromised for isoluminant targets moving in the periphery. In this case, the oculomotor system can use an accurate, but delayed position component, but cannot account for target movement. This deficit was also present for the postsaccadic pursuit speed. For the two luminance conditions, pursuit direction and speed were adjusted depending on the saccadic landing position. The rapid postsaccadic pursuit adjustments suggest shared position- and motion-related signals of target and eye for saccade and pursuit control.

The role of eye movements in manual responses to social and nonsocial cues.

Gaze and arrow cues cause covert attention shifts even when they are uninformative. Nonetheless, it is unclear to what extent oculomotor behavior influences manual responses to social and nonsocial stimuli. In two experiments, we tracked the gaze of participants during the cueing task with nonpredictive gaze and arrow cues. In Experiment 1, the discrimination task was easy and eye movements were not necessary, whereas in Experiment 2 they were instrumental in identifying the target. Validity effects on manual response time (RT) were similar for the two cues in Experiment 1 and in Experiment 2, though in the presence of eye movements observers were overall slower to respond to the arrow cue compared with the gaze cue. Cue direction had an effect on saccadic performance before the discrimination was presented and throughout the duration of the trial. Furthermore, we found evidence of a distinct impact of the type of cue on diverse oculomotor components. While saccade latencies were affected by the type of cue, both before and after the target onset, saccade landing positions were not. Critically, the manual validity effect was predicted by the landing position of the initial eye movement. This work suggests that the relationship between eye movements and attention is not straightforward. In the presence of overt selection, saccade latency related to the overall speed of manual response, while eye movements landing position was closely related to manual performance in response to different cues.

Spatial and temporal dynamics of presaccadic attentional facilitation before pro- and antisaccades.

The premotor theory of attention and the visual attention model make different predictions about the temporal and spatial allocation of presaccadic attentional facilitation. The current experiment investigated the spatial and temporal dynamics of presaccadic attentional facilitation during pro- and antisaccade planning; we investigated whether attention shifts only to the saccade goal location or to the target location or elsewhere, and when. Participants performed a dual-task paradigm with blocks of either anti- or prosaccades and also discriminated symbols appearing at different locations before saccade onset (measure of attentional allocation). In prosaccades blocks, correct prosaccade discrimination was best at the target location, while during errors, discrimination was best at the location opposite to the target location. This pattern was inversed in antisaccades blocks, although discrimination remained high opposite to the target location. In addition, we took the benefit of a large range of saccadic landing positions and showed that performance across both types of saccades was best at the actual saccade goal location (where the eye will actually land) rather than at the instructed position. Finally, temporal analyses showed that discrimination remained highest at the saccade goal location, from long before to closer to saccade onset, increasing slightly for antisaccades closer to saccade onset. These findings are in line with the premises of the premotor theory of attention, showing that attentional allocation is primarily linked both temporally and spatially to the saccade goal location.

The faster you decide, the more accurate localization is possible: Position representation of "curveball illusion" in perception and eye movements.

When the inside texture of a moving object moves, the perceived motion of the object is often distorted toward the direction of the texture’s motion (motion-induced position shift), and such perceptual distortion accumulates while the object is watched, causing what is known as the curveball illusion. In a recent study, however, the accumulation of the position error was not observed in saccadic eye movements. Here, we examined whether the position of the illusory object is represented independently in the perceptual and saccadic systems. In the experiments, the stimulus of the curveball illusion was adopted to examine the temporal change in the position representation for saccadic eye movements and for perception by varying the elapsed time from the input of visual information to saccade onset and perceptual judgment, respectively. The results showed that the temporal accumulation of the motion-induced position shift is observed not only in perception but also in saccadic eye movements. In the saccade tasks, the landing positions of saccades gradually shifted to the illusory perceived position as the elapsed time from the target offset to the saccade “go” signal increased. Furthermore, in the perception task, shortening the time between the target offset and the perceptual judgment reduced the size of the illusion effect. Therefore, these results argue against the idea of dissociation between saccadic and perceptual localization of a moving object suggested in the previous study, in which saccades were measured in a rushed way while perceptual responses were measured without time constraint. Instead, the similar temporal trends of these effects imply a common or similar target representation for perception and eye movements which dynamically changes over the course of evidence accumulation.

Saccade landing position prediction for gaze-contingent rendering

Gaze-contingent rendering shows promise in improving perceived quality by providing a better match between image quality and the human visual system requirements. For example, information about fixation allows rendering quality to be reduced in peripheral vision, and the additional resources can be used to improve the quality in the foveal region. Gaze-contingent rendering can also be used to compensate for certain limitations of display devices, such as reduced dynamic range or lack of accommodation cues. Despite this potential and the recent drop in the prices of eye trackers, the adoption of such solutions is hampered by system latency which leads to a mismatch between image quality and the actual gaze location. This is especially apparent during fast saccadic movements when the information about gaze location is significantly delayed, and the quality mismatch can be noticed. To address this problem, we suggest a new way of updating images in gaze-contingent rendering during saccades. Instead of rendering according to the current gaze position, our technique predicts where the saccade is likely to end and provides an image for the new fixation location as soon as the prediction is available. While the quality mismatch during the saccade remains unnoticed due to saccadic suppression, a correct image for the new fixation is provided before the fixation is established. This paper describes the derivation of a model for predicting saccade landing positions and demonstrates how it can be used in the context of gaze-contingent rendering to reduce the influence of system latency on the perceived quality. The technique is validated in a series of experiments for various combinations of display frame rate and eye-tracker sampling rate.

Saccades to Explicit and Virtual Features in the Poggendorff Figure Show Perceptual Biases.

Human participants made saccadic eye movements to various features in a modified vertical Poggendorff figure, to measure errors in the location of key geometrical features. In one task, subjects (n = 8) made saccades to the vertex of the oblique T-intersection between a diagonal pointer and a vertical line. Results showed both a small tendency to shift the saccade toward the interior of the angle, and a larger bias in the direction of a shorter saccade path to the landing line. In a different kind of task (visual extrapolation), the same subjects fixated the tip of a 45° pointer and made a saccade to the implicit point of intersection between pointer and a distant vertical line. Results showed large errors in the saccade landing positions and the saccade polar angle, in the direction predicted from the perceptual Poggendorff bias. Further experiments manipulated the position of the fixation point relative to the implicit target, such that the Poggendorff bias would be in the opposite direction from a bias toward taking the shortest path to the landing line. The bias was still significant. We conclude that the Poggendorff bias in eye movements is in part due to the mislocation of visible target features but also to biases in planning a saccade to a virtual target across a gap. The latter kind of error comprises both a tendency to take the shortest path to the landing line, and a perceptual error that overestimates the vector component orthogonal to the gap.

The influence of distractors on express saccades.

It is well known that regular target-driven saccades are affected by the presence of close and remote distractors. Distractors close to the target affect the saccade landing position (known as the global effect), while remote distractors prolong saccade latencies to the target (known as the remote-distractor effect). Little is known about whether a different population of saccades known as express saccades (saccades with very short latencies between 80 and 130 ms) is similarly affected by close and remote distractors, as these saccades are considered to be the result of advanced preparation of an oculomotor program toward the target. We designed a task in which we were able to generate a large number of express saccades, as evidenced by a separate and very early peak in the saccade-latency distribution-a distribution that was different from that of regular saccades. Our results show that irrelevant and unexpected visual input had a large effect on express saccades. We found a global and a remote-distractor effect which were similar to those seen in regular saccades. Even though our findings confirm the existence of very-short-latency saccades in humans, it is questionable whether they represent a different population of saccades, as they were equally affected by the presence of distractors as are regular saccades.

Adaptation of Saccadic Sequences with and without Remapping.

It is relatively easy to adapt visually-guided saccades because the visual vector and the saccade vector match. The retinal error at the saccade landing position is compared to the prediction error, based on target location and efference copy. If these errors do not match, planning processes at the level(s) of the visual and/or motor vector processing are assumed to be inaccurate and the saccadic response is adjusted. In the case of a sequence of two saccades, the final error can be attributed to the last saccade vector or to the entire saccadic displacement. Here, we asked whether and how adaptation can occur in the case of remapped saccades, such as during the classic double-step saccade paradigm, where the visual and motor vectors of the second saccade do not coincide and so the attribution of error is ambiguous. Participants performed saccades sequences to two targets briefly presented prior to first saccade onset. The second saccade target was either briefly re-illuminated (sequential visually-guided task) or not (remapping task) upon first saccade offset. To drive adaptation, the second target was presented at a displaced location (backward or forward jump condition or control—no jump) at the end of the second saccade. Pre- and post-adaptation trials were identical, without the re-appearance of the target after the second saccade. For the 1st saccade endpoints, there was no change as a function of adaptation. For the 2nd saccade, there was a similar increase in gain in the forward jump condition (52% and 61% of target jump) in the two tasks, whereas the gain decrease in the backward condition was much smaller for the remapping task than for the sequential visually-guided task (41% vs. 94%). In other words, the absolute gain change was similar between backward and forward adaptation for remapped saccades. In conclusion, we show that remapped saccades can be adapted, suggesting that the error is attributed to the visuo-motor transformation of the remapped visual vector. The mechanisms by which adaptation takes place for remapped saccades may be similar to those of forward sequential visually-guided saccades, unlike those involved in adaptation for backward sequential visually-guided saccades.

Revealing the time course of signals influencing the generation of secondary saccades using Aalen’s additive hazards model

Saccadic eye movements are frequently followed by smaller secondary saccades which are generally assumed to correct for the error in primary saccade landing position. However, secondary saccades can also occur after accurate primary saccades and they are often as small as microsaccades, therefore raising the need to further scrutinize the processes involved in secondary saccade generation. Following up a previous study, we analyzed secondary saccades using rate analysis which allows us to quantify experimental effects as shifts in distributions, therefore going beyond comparisons of mean differences. We use Aalen’s additive hazards model to delineate the time course of key influences on the secondary saccade rate. In addition to the established effect of primary saccade error, we observed a time-varying influence of under- vs. overshooting – with a higher risk of generating secondary saccades following undershoots. Moreover, increasing target eccentricity influenced the programming of secondary saccades, therefore demonstrating that error-unrelated variables co-determine secondary saccade programs. Our results provide new insights into the generative mechanisms of small saccades during postsaccadic fixation that need to be accounted for by secondary saccade models.