Visual perception of direction when voluntary saccades occur. I. Relation of visual direction of a fixation target extinguished before a saccade to a flash presented during the saccade

In experiments designed to clarify the mechanisms underlying the normal stability of visual direction for stationary objects when voluntary saccades occur, Ss reported on the horizontal visual direction of a brief test [lash presented when the eye was at a specific point in the saccade (the trigger point) relative to a fixation target viewed and extinguished prior to the saccade. From these reports, PSEs (points of subjective equality) were calculated for the fixation target as measured by the test [lashes. The distance of the trigger point from the previous fixation position was systematically varied in each experiment. Different experiments required saccades of different lengths and directions. With the exception of the presentation of the test [lash the saccades were carried out in complete darkness so that the possible utilization of an extraretinal signal regarding the eye movement (change in eye position, the intention to turn the eye, or a change of attention related to the eye movement) in the determination of visual direction could be observed uncomplicated by a continuing visual context. According to classical theories, an extraretinal signal proportional to the change in eye position acts to maintain direction constancy by compensating for the Shift of the retinal image resulting from the movement of the eye. In general, direction constancy was not preserved in the present experiments, and thus the data would not be predicted by classical theories. However, the PSE varied with distance of the trigger point from the fixation target. Since this displacement of PSE from the trigger point was in the correct direction for compensation, the presence of an extraretinal signal was confirmed. However, the growth of this signal appears to be time-locked to the saccade rather than locked to eye position; it is suggested that this growth takes place over a time period which is longer than the duration of the saccade itself.     

A novel evaluation of two related and two independent algorithms for eye movement classification during reading

Nystr?m and Holmqvist have published a method for the classification of eye movements during reading (ONH) (Nyström & Holmqvist, 2010). When we applied this algorithm to our data, the results were not satisfactory, so we modified the algorithm (now the MNH) to better classify our data. The changes included: (1) reducing the amount of signal filtering, (2) excluding a new type of noise, (3) removing several adaptive thresholds and replacing them with fixed thresholds, (4) changing the way that the start and end of each saccade was determined, (5) employing a new algorithm for detecting PSOs, and (6) allowing a fixation period to either begin or end with noise. A new method for the evaluation of classification algorithms is presented. It was designed to provide comprehensive feedback to an algorithm developer, in a time-efficient manner, about the types and numbers of classification errors that an algorithm produces. This evaluation was conducted by three expert raters independently, across 20 randomly chosen recordings, each classified by both algorithms. The MNH made many fewer errors in determining when saccades start and end, and it also detected some fixations and saccades that the ONH did not. The MNH fails to detect very small saccades. We also evaluated two additional algorithms: the EyeLink Parser and a more current, machine-learning-based algorithm. The EyeLink Parser tended to find more saccades that ended too early than did the other methods, and we found numerous problems with the output of the machine-learning-based algorithm.     

Strategic control over saccadic eye movements: studies of the fixation offset effect

We studied the strategic (presumably cortical) control of ocular fixation in experiments that measured the fixation offset effect (FOE) while manipulating readiness to make reflexive or voluntary eye movements. The visual grasp reflex, which generates reflexive saccades to peripheral visual signals, reflects an opponent process in the superior colliculus (SC) between fixation cells at the rostral pole, whose activity helps maintain ocular position and increases when a stimulus is present at fixation, and movement cells, which generate saccades and are inhibited by rostral fixation neurons. Voluntary eye movements are controlled by movement and fixation cells in the frontal eye field (FEF). The FOE--a decrease in saccade latency when the fixation stimulus is extinguished--has been shown to reflect activity in the collicular eye movement circuitry and also to have an activity correlate in the FEF. Our manipulation of preparatory set to make reflexive or voluntary eye movements showed that when reflexive saccades were frequent and voluntary saccades were infrequent, the FOE was attenuated only for reflexive saccades. When voluntary saccades were frequent and reflexive saccades were infrequent, the FOE was attenuated only for voluntary saccades. We conclude that cortical processes related to task strategy are able to decrease fixation neuron activity even in the presence of a fixation stimulus, resulting in a smaller FOE. The dissociation in the effects of a fixation stimulus on reflexive and voluntary saccade latencies under the same strategic set suggests that the FOEs for these two types of eye movements may reflect a change in cellular activity in different neural structures, perhaps in the SC for reflexive saccades and in the FEF for voluntary saccades.     

An adaptive algorithm for fixation,saccade, and glissade detection in eyetracking data

Event detection is used to classify recorded gaze points into periods of fixation, saccade, smooth pursuit, blink, and noise. Although there is an overall consensus that current algorithms for event detection have serious flaws and that a de facto standard for event detection does not exist, surprisingly little work has been done to remedy this problem. We suggest a new velocity-based algorithm that takes several of the previously known limitations into account. Most important, the new algorithm identifies so-called glissades, a wobbling movement at the end of many saccades, as a separate class of eye movements. Part of the solution involves designing an adaptive velocity threshold that makes the event detection less sensitive to variations in noise level and the algorithm settings-free for the user. We demonstrate the performance of the new algorithm on eye movements recorded during reading and scene perception and compare it with two of the most commonly used algorithms today. Results show that, unlike the currently used algorithms, fixations, saccades, and glissades are robustly identified by the new algorithm. Using this algorithm, we found that glissades occur in about half of the saccades, during both reading and scene perception, and that they have an average duration close to 24 msec. Due to the high prevalence and long durations of glissades, we argue that researchers must actively choose whether to assign the glissades to saccades or fixations; the choice affects dependent variables such as fixation and saccade duration significantly. Current algorithms do not offer this choice, and their assignments of each glissade are largely arbitrary.     

Programming of endogenous and exogenous saccades: evidence for a competitive integration model

Participants were required to make a saccade to a uniquely colored target while ignoring the presentation of an onset distractor. The results provide evidence for a competitive integration model of saccade programming that assumes endogenous and exogenous saccades are programmed in a common saccade map. The model incorporates a lateral interaction structure in which saccade-related activation at a specific location spreads to neighboring locations but inhibits distant locations. In addition, there is top-down, location-specific inhibition of locations to which the saccade should not go. The time course of exogenous and endogenous activation in the saccade map can explain a variety of eye movement data, including endpoints, latencies, and trajectories of saccades and the well-known global effect.     

Saccade generation during reading: Are words necessary?

Most current theories of eye movement control during reading are word based in multiple ways: They assume that saccade onset times result from word‐based processes, and that words are involved in selecting a saccade target. In the current study the role of words was examined by occasionally replacing the text with one of five alternate stimulus patterns for a single fixation during reading, and observing the effects on the time, direction, and length of the saccade that ends that fixation. The onset times of many saccades are unaffected by replacing spaces with random letters, thus removing visible word‐units; also, the effects of this removal on saccade length is not different than that of having space‐delimited nonwords. It does not appear that words play a critical role in generating saccades. The results are compatible with the Competition/Interaction theory of eye movement control during reading (Yang & McConkie, 2001).     

Memory representations guide targeting eye movements in a natural task

The change blindness phenomenon suggests that visual representations retained across saccades are very limited. In this paper we sought to specify the kind of information that is in fact retained. We investigated targeting performance for saccadic eye movements, since one need for visual representations across eye and body positions may be to guide coordinated movements. We examined saccades in the context of an ongoing sensory motor task in order to make stronger generalizations about natural visual functioning and deployment of attention. Human subjects copied random patterns of coloured blocks on a computer display. Their eye movement pattern was consistent from block to block, including a precise saccade to a previously-placed, neighbouring block during each additional block placement. This natural, consistent eye movement allowed the previously-placed, neighbouring block to serve as an implicit target without instructions to the subject. On random trials, we removed the target object from the display during a preceding saccade, so that observers were required to make the targeting saccade without a currently visible target. Targeting performance was excellent, and appeared to be influenced by spatial information that was not visible during the preceding fixation. Subjects were generally unaware of the disappearance and reappearance of the target. We conclude that spatial information about visual targets is retained across eye movements and used to guide subsequent movements.     

Saccadic suppression of displacement is strongest in central vision

Perception of target displacement is severely degraded if the displacement occurs during a saccadic eye movement, but the variation of this effect across the visual field is unknown. A small target was displaced from a starting point at the midline, or 10 deg to the right or left, while the eye made a saccade from the 10 deg right position to the 10 deg left position. Saccades were detected and the target displaced on line. Assessed with a signal detection measure, suppression was stronger in central vision than in more peripheral locations for all three subjects. Leftward and rightward displacements yielded equal thresholds. The results complement the findings of others to reveal a picture of perceptual events during saccades, with both deeper saccadic suppression and faster correction of spatial values (the correspondences between retinal position and perceived egocentric direction), favouring more accurate spatial processing in central vision than in the periphery.     

One algorithm to rule them all? An evaluation and discussion of ten eye movement event-detection algorithms

Almost all eye-movement researchers use algorithms to parse raw data and detect distinct types of eye movement events, such as fixations, saccades, and pursuit, and then base their results on these. Surprisingly, these algorithms are rarely evaluated. We evaluated the classifications of ten eye-movement event detection algorithms, on data from an SMI HiSpeed 1250 system, and compared them to manual ratings of two human experts. The evaluation focused on fixations, saccades, and post-saccadic oscillations. The evaluation used both event duration parameters, and sample-by-sample comparisons to rank the algorithms. The resulting event durations varied substantially as a function of what algorithm was used. This evaluation differed from previous evaluations by considering a relatively large set of algorithms, multiple events, and data from both static and dynamic stimuli. The main conclusion is that current detectors of only fixations and saccades work reasonably well for static stimuli, but barely better than chance for dynamic stimuli. Differing results across evaluation methods make it difficult to select one winner for fixation detection. For saccade detection, however, the algorithm by Larsson, Nyström and Stridh (IEEE Transaction on Biomedical Engineering, 60(9):2484–2493,2013) outperforms all algorithms in data from both static and dynamic stimuli. The data also show how improperly selected algorithms applied to dynamic data misestimate fixation and saccade properties.     

An automated eye movement laboratory for on-line electrooculography

This paper describes an automated eye movement laboratory that uses electrooculography (EOG) to study people’s eye movements while they read. An on-line minicomputer processes bioelectric potentials that correspond to saccadic eye movements. Horizontal saccades larger than 1.5 deg of visual angle are detected and analyzed in real-time as they occur. The laboratory is designed for prolonged yet unobtrusive observation of human eye movements during sustained reading periods of minutes or hours. All important functions regarding data collection and data reduction are performed automatically, according to simple procedures that can be applied uniformly and without bias to nearly all subjects that we study. Results from three experiments are cited in order to quantify the performance of the laboratory with respect to four criteria: saccade detection accuracy, measurement accuracy, sensitivity, and the uniformity of these measures over different subjects.     

Minicomputer processing of dual Purkinje image eye-tracker data

Eye movement data analysis often entails reliable detection of the onset of saccades. Saccades are most easily separated from fixations on the basis of movement velocity and acceleration. Convolution integers are used to estimate the velocity and acceleration of eye movements, and these serve as the basis of an eye movement data-classification program. The technique of derivative estimation by convolution integers has the advantage of being easily calculated and the disadvantage of returning results shifted in phase. The disadvantage is overcome by using ring buffers for the synchronization of data and results.     

Endogenous saccades are preceded by shifts of visual attention: evidence from cross-saccadic priming effects

The present study examines whether endogenous saccades are preceded by shifts of attention. Three experiments are reported in which participants were required to execute a saccadic eye movement to a certain location and to subsequently identify the orientation of a target triangle. Prior to the execution of the saccade a prime was presented, which was compatible or incompatible with the target. A priming effect (faster responses in the compatible condition than in the incompatible condition) occurred only when the prime was presented at the saccade destination, and this effect was larger when the prime was presented during oculomotor programming than when it was presented prior to oculomotor programming. The results indicate that an endogenous shift of attention precedes endogenous saccades, providing further support for theories of visual selection that assume a tight coupling between attention and saccades.     

Programming saccadic eye movements

This article addresses questions about the preparatory processes that immediately precede saccadic eye movements. Saccade latencies were measured in a task in which subjects were provided partial advance information about the spatial location of a target fixation. In one experiment, subjects were faster in initiating saccades when they knew either the direction or amplitude of the required movement in advance compared to a condition with equal uncertainty about the number of potential saccade targets but without knowledge of the parameters required to execute the movement. These results suggest that the direction and amplitude for an upcoming saccade were calculated separately, and not in a fixed serial order. In another experiment, subjects appear to have programmed the saccades more holistically--with computations of direction and amplitude parameters occurring simultaneously. The implications of these results for models of eye movement preparation are discussed.     

Expedited suppression signal in ocular go/no-go decision

When scrutinizing the visual world, complex and unexpected stimuli often lead to prolonged eye fixations to enhance cognitive processing, likely by temporarily suppressing a planned saccade. The present study examined whether the suppression signal is tightly linked to a specific planned saccade and if it conforms to the viewer's intention. A novel Go/No-go task was devised where participants made consecutive saccades to fixate a stimulus appearing across the screen horizontal meridian in 4° steps. At times, the features of the stimulus (colour and/or shape) were altered when it reappeared at a new location. Participants had to suppress the saccade that would otherwise leave the stimulus if its features matched instructed criteria. Saccade suppression was determined by the reduced probability for saccades towards and away from a target stimulus. Results show both correct suppression to saccades leaving the target and erroneous suppression to saccades towards it. The erroneous suppression was initially observed for any change in features but later lifted. The suppression shortened the length of saccades leaving a target but not those towards it. The initial suppression during previewing the target appears to be based on expedited but incomplete evaluation of visual stimulus, and is not linked to any specific saccade. These properties might reflect the stage of ocular decision based on which the suppression signal is generated. They also account for the phenomenon of “peripheral-to-foveal” effect on eye movements in reading.     

Adaptive modification of saccadic eye movements.

Experiments are reported in which the target for a saccadic eye movement was displaced during the saccade. Subjects adapted to the displacement by altering the amplitudes of subsequent saccades to compensate for it. Analysis of kinematic details of the saccade trajectories revealed that the adaptation did not arise from a simple remapping of perceived target locations. Instead, the adaptation appeared to be accomplished by a change in the gain of the saccadic system. The gain change arose primarily from a change in the magnitude of the force pulse for the saccade, not a change in the duration of the pulse. These results have implications for the mechanisms that underlie saccades in normal situations. In particular, people can separately adjust the magnitudes and durations of the force pulses used to produce saccades.     

Impaired initiation but not execution of contralesional saccades in hemispatial neglect

Patients with unilateral neglect are impaired at making saccades to contralesional targets. Whether this problem arises from a deficit in perception, in planning the saccade or in executing the eye movement or some combination thereof remains unclear. We measured several variables related to the initiation and execution of saccades in an experiment which crossed two factors: target side (left, right) and direction of saccade (leftwards, rightwards). Relative to control subjects, patients with left-sided neglect were impaired in planning but not executing the contralesional saccade; while the latency to move their eyes following the onset of the target was increased, the duration and velocity to reach the target were normal. In addition, there were also no directional differences for saccades that were hypometric or inaccurate in the patients, further ruling out an execution impairment. Interestingly, this directional initiation deficit was exaggerated for leftward saccades to left targets, compared with all other conditions. We suggest that the disadvantage for contralesional saccades in neglect patients is attributable to a deficit not only in perceiving contralateral targets but also in planning leftward saccades. Once the saccade is initiated, however, execution apparently proceeds unimpaired.     

Age-group differences in inhibiting an oculomotor response

Age-group differences were examined in the delayed oculomotor response task, which requires that observers delay the execution of a saccade (eye movement) toward an abrupt-onset visual cue. This task differs from antisaccade and attentional capture in that inhibition causes saccades to be postponed, not redirected. Older adults executed more premature saccades than young adults, but there were no age-group differences in latency or accuracy of saccades executed at the proper time. The results suggest that older adults are less capable of inhibiting a prepotent saccadic response, but that other aspects of visual working memory related to the task are preserved.     

Backward and forward masking associated with saccadic eye movement

Visual masking effects on test flash thresholds were measured under real and simulated eye movement conditions to determine whether visual masking is primarily responsible for elevations in threshold that are sometimes associated with saccadic eye movements. Brief luminous flashes presented to the central retina before, during, and after saccades were masked by stimuli presented either pre- or postsaccadically. The amount and time course of masking were quantitatively dependent on stimulus parameters of intensity and temporal separation and were unaffected by eye movement parameters (amplitude, velocity, direction) as long as retinal stimulus conditions were constant. The duration of forward masking was longer than that of backward masking. When retinal conditions during saccades were mimicked while the eyes were held steady, masking interactions were identical to those obtained during real saccades. These results indicate that masking effects during saccades in ordinary environments are determined solely by the stimulus situation at the retina. Putative nonvisual, centrally originating saccadic suppression suggested by other authors is evidently not additive with visually determined masking during saccades.     

Target localization after saccades and at fixation: Nontargets both facilitate and bias responses

ABSTRACT

The image on our retina changes every time we make an eye movement. To maintain visual stability after saccades, specifically to locate visual targets, we may use nontarget objects as “landmarks”. In the current study, we compared how the presence of nontargets affects target localization after saccades and during sustained fixation. Participants fixated a target object, which either maintained its location on the screen (sustained-fixation trials), or displaced to trigger a saccade (saccade trials). After the target disappeared, participants reported the most recent target location with a mouse click. We found that the presence of nontargets decreased response error magnitude and variability. However, this nontarget facilitation effect was not larger for saccade trials than sustained-fixation trials, indicating that nontarget facilitation might be a general effect for target localization, rather than of particular importance to post-saccadic stability. Additionally, participants’ responses were biased towards the nontarget locations, particularly when the nontarget-target relationships were preserved in relative coordinates across the saccade. This nontarget bias interacted with biases from other spatial references, e.g., eye movement paths, possibly in a way that emphasized non-redundant information. In summary, the presence of nontargets is one of several sources of reference that combine to influence (both facilitate and bias) target localization.     

Enhanced saccadic control in young people with Tourette syndrome despite slowed pro‐saccades

Tourette syndrome (TS) is a neurodevelopmental disorder characterized by motor and vocal tics. Tics are repetitive and uncontrolled behaviours that have been associated with basal ganglia dysfunction. We investigated saccadic eye movements in a group of young people with TS but without co‐morbid ADHD. Participants performed two tasks. One required them to perform only pro‐saccade responses (pure pro‐saccade task). The other involved shifting, unpredictably, between executing pro‐ and anti‐saccades (mixed saccade task). We show that in the mixing saccade task, the TS group makes significantly fewer errors than an age‐matched control group, while responding equally fast. By contrast, on the pure pro‐saccade task, the TS group were shown to be significantly slower to initiate and to complete the saccades (longer movement duration and decreased peak velocity) than controls, while movement amplitude and direction accuracy were not different. These findings demonstrate enhanced shifting ability despite slower reflexive responding in TS and are discussed with respect to a disorder‐related adaptation for increased cognitive regulation of behaviour.