Perceived localizations and eye movements with action‐generated and computer‐generated vanishing points of moving stimuli

When observers localize the vanishing point of a moving target, localizations are reliably displaced beyond the final position, in the direction the stimulus was travelling just prior to its offset. We examined modulations of this phenomenon through eye movements and action control over the vanishing point. In Experiment 1 with pursuit eye movements, localization errors were in movement direction, but less pronounced when the vanishing point was self‐determined by a key press of the observer. In contrast, in Experiment 2 with fixation instruction, localization errors were opposite movement direction and independent from action control. This pattern of results points at the role of eye movements, which were gathered in Experiment 3. That experiment showed that the eyes lagged behind the target at the point in time, when it vanished from the screen, but that the eyes continued to drift on the targets' virtual trajectory. It is suggested that the perceived target position resulted from the spatial lag of the eyes and of the persisting retinal image during the drift.     

Saccadic suppression relies on luminance information

To determine whether saccadic suppression of image displacement uses information from luminance channels, we measured spatial displacement detection thresholds with equiluminant and nonequiluminant targets during saccades. We compared these saccadic thresholds with displacement thresholds measured during fixation by making ratios of saccadic thresholds to fixation thresholds. Ratios were lower in the equiluminant condition than in the nonequiluminant. This surprising result indicates that detection of equiluminant target displacements during saccades was better than detection of nonequiluminant targets, compared with the detection abilities during fixation. Thus, saccadic suppression of image displacement, which should increase displacement thresholds during saccades over fixation thresholds, was more effective with nonequiluminant targets. Because of target flicker, displacement thresholds were anisotropic in the nonequiluminant condition; thresholds were greater when target and eye moved in the same direction than when they moved in opposite directions, consistent with earlier results. These two effects (flicker-induced anisotropy and greater suppression in nonequiluminance) canceled when the eye moved opposite the displacement, yielding equal thresholds, and summed when eye and target moved in the same direction, yielding large threshold differences. We conclude that saccadic suppression of image displacement uses mechanisms sensitive to luminance contrast.     

Does gaze perception facilitate overt orienting?

Recent studies have demonstrated the facilitation of responses to peripheral targets cued by the direction of the gaze of a face. However, in the absence of data from an eye tracker, it has been unclear to what extent these effects are due to the participants making small saccades in response to the cue that bring them closer to the congruent location of the cued targets. We used an eye tracker to show that while such cue-driven saccades occur, they do not account for the main cueing effects observed. Additionally, by using the same general paradigm as has previously been used with infants, we suggest that different mechanisms underlie eye gaze cueing effects in infants and adults.     

Gaze direction and the extraction of egocentric distance

The angular declination of a target with respect to eye level is known to be an important cue to egocentric distance when objects are viewed or can be assumed to be resting on the ground. When targets are fixated, angular declination and the direction of the gaze with respect to eye level have the same objective value. However, any situation that limits the time available to shift gaze could leave to-be-localized objects outside the fovea, and, in these cases, the objective values would differ. Nevertheless, angular declination and gaze declination are often conflated, and the role for retinal eccentricity in egocentric distance judgments is unknown. We report two experiments demonstrating that gaze declination is sufficient to support judgments of distance, even when extraretinal signals are all that are provided by the stimulus and task environment. Additional experiments showed no accuracy costs for extrafoveally viewed targets and no systematic impact of foveal or peripheral biases, although a drop in precision was observed for the most retinally eccentric targets. The results demonstrate the remarkable utility of target direction, relative to eye level, for judging distance (signaled by angular declination and/or gaze declination) and are consonant with the idea that detection of the target is sufficient to capitalize on the angular declination of floor-level targets (regardless of the direction of gaze).     

Extraretinal information about eye position during involuntary eye movement: optokinetic afternystagmus

Despite importance for theories of perception, controversy exists as to whether information is available to the perceptual system about involuntary as well as voluntary eye movements. We measured the perceived direction of targets flashed briefly in an otherwise dark field during the primary phase of optokinetic afternystagmus (OKAN), an involuntary eye movement that persists in darkness following optokinetic stimulation. Perceived direction was measured by unseen pointing in one experiment and by pointing made under visual control in a second experiment. Pointing was essentially veridical in both experiments, indicating that accurate extra-retinal information about eye position (presumably, as efference copy) exists for OKAN. Illusory motion of visual targets, which can occur during involuntary oculomotor responses, therefore cannot be attributed to a lack of efference-copy signals for such eye movements.     

Human Eye Movements During Visually Guided Stepping

Visually guided locomotion was studied in an experiment in which human subjects (N = 8) had to accurately negotiate a series of irregularly spaced stepping-stones while infrared reflectometry and electrooculography were used to continuously record their eye movements. On average, 68% of saccades made toward the next target of footfall had been completed (visual target capture had occurred) while the foot to be positioned was still on the ground; the remainder were completed in the first 300 ms of the swing phase. The subjects' gaze remained fixed on a target, on average, until 51 ms after making contact with it, with little variation. A greater amount of variation was seen in the timing of trailing footlift relative to visual target capture. Assuming that subjects sampled the visual cues as and when they were required, visual information appeared most useful when the foot to be positioned was still on the ground.     

Perceptual localization of visual stimuli flashed during saccades

Subjects were asked to make a saccade to a visual target flashed in the dark during a prior primary saccade, and to report its apparent position by moving an adjustable light spot to that position. When targets were presented at the beginning of the primary saccade, subjects perceptually mislocated them in the direction of the saccade, whereas when targets were presented immediately before the end of the primary saccade, the flashed targets were mislocated in the opposite direction. The perceptually localized position of the target was primarily determined by its retinal position. However, at all actual and retinal positions of the target, the localized position shifted from the position that would be predicted if the location of the target was determined only by its retinal position to the prior primary saccade direction. The results were discussed in relation to extraretinal eye position signals. Subjects moved their eyes not to the actual position of the target, but to its apparent position. In some trials, there was a discrepancy between perceptual and oculomotor localization, which was interpreted as having been caused by the imprecise localization ability of the oculomotor system.     

The relationship between eye position and egocentric visual direction.

Hering's model of egocentric visual direction assumes implicitly that the effect of eye position on direction is both linear and equal for the two eyes; these two assumptions were evaluated in the present experiment. Five subjects pointed (open-loop) to the apparent direction of a target seen under conditions in which the position of one eye was systematically varied while the position of the other eye was held constant. The data were analyzed through examination of the relationship between the variations in perceived egocentric direction and variations in expected egocentric direction based on the positions of the varying eye. The data revealed that the relationship between eye position and egocentric direction is indeed linear. Further, the data showed that, for some subjects, variations in the positions of the two eyes do not have equal effects on egocentric direction. Both the between-eye differences and the linear relationship may be understood in terms of individual differences in the location of the cyclopean eye, an unequal weighting of the positions of the eyes in the processing of egocentric direction, or some combination of these two factors.     

The relationship between eye position and egocentric visual direction

Hering’s model of egocentric visual direction assumes implicitly that the effect of eye position on direction is both linear and equal for the two eyes; these two assumptions were evaluated in the present experiment. Five subjects pointed (open-loop) to the apparent direction of a target seen under conditions in which the position of one eye was systematically varied while the position of the other eye was held constant. The data were analyzed through examination of the relationship between the variations in perceived egocentric direction and variations inexpected egocentric direction based on the positions of the varying eye. The data revealed that the relationship between eye position and egocentric direction is indeed linear. Further, the data showed that, for some subjects, variations in the positions of the two eyes do not have equateffectsTjn egocentric direction. Both the between-eye differences and the linear relationship may be understood in terms of individual differences in the location of the cyclopean eye, an unequal weighting of the positions of the eyes in the processing of egocentric direction, or some combination of these two factors.     

The relationship between eye movements and spatial attention

Most previous studies of the attentional consequences of making saccadic eye movements have used peripheral stimuli to elicit eye movements. It is argued that in the light of evidence showing automatic “capture” of attention by peripheral stimuli, these experiments do not distinguish between attentional effects due to peripheral stimuli and those due to eye movements. In the present study, spatial attention was manipulated by varying the probability that peripheral probe stimuli would appear in different positions, while saccades were directed by a central arrow, enabling the effects of attention and eye movements to be separated. The results showed that the time to react to a peripheral stimulus could be shortened both by advance knowledge of its likely position and, separately, by preparing to make a saccade to that position. When the saccade was directed away from the most likely position of the probe, the targets for attention and eye movements were on opposite sides of the display. In this condition, the effects of preparing to make a saccade proved to be stronger than the effects of attentional allocation until well after the saccade had finished, suggesting that making a saccade necessarily involves the allocation of attention to the target position. The effects of probe stimuli on saccade latencies were also examined: probe stimuli that appeared before the saccade shortened saccade latencies if they appeared at the saccade target, and lengthened saccade latencies if they appeared on the opposite side of fixation. These facilitatory and inhibitory effects were shown to occur at different stages of saccade preparation and suggest that attention plays an important role in the generation of voluntary eye movements. The results of this study indicate that while it is possible to make attention movements without making corresponding eye movements, it is not possible to make an eye movement (in the absence of peripheral stimulation) without making a corresponding shift in the focus of attention.     

An investigation of the relationship between eye and retinal image movement in the perception of movement

The question investigated was whether or not eye movements accompanied by abnormal retinal image movements, movements that are either or both at a different rate or in a different direction than the eye movement, predictably lead to perceived movement. Os reported whether or not they saw a visual target move when the movement of the target was either dependent on and simultaneous with their eye movements or when the target movement was independent of their eye movements. In the main experiment, observations were made when the ratio between eye and target movement fem/tm) was 2/5, 1/5, 1/10, 1/20, and 0. All these ratios were tested when the direction of the target movement was in the same (H+), opposite (H?), and at right angles to (V+, V?) the movement of the eyes. Eye movements, target movements, and reports of target movement were recorded. Results indicate that a discrepancy between eye and target movement greater than 20% predictably leads to perceived target movement, whereas a discrepancy of 5% or less rarely leads to perceived movement. The results are interpreted as support for the operation of a compensatory mechanism during eye movements.     

“Avoiding or approaching eyes”? Introversion/extraversion affects the gaze-cueing effect

We investigated whether the extra-/introversion personality dimension can influence processing of others’ eye gaze direction and emotional facial expression during a target detection task. On the basis of previous evidence showing that self-reported trait anxiety can affect gaze-cueing with emotional faces, we also verified whether trait anxiety can modulate the influence of intro-/extraversion on behavioral performance. Fearful, happy, angry or neutral faces, with either direct or averted gaze, were presented before the target appeared in spatial locations congruent or incongruent with stimuli’s eye gaze direction. Results showed a significant influence of intra-/extraversion dimension on gaze-cueing effect for angry, happy, and neutral faces with averted gaze. Introverts did not show the gaze congruency effect when viewing angry expressions, but did so with happy and neutral faces; extraverts showed the opposite pattern. Importantly, the influence of intro-/extraversion on gaze-cueing was not mediated by trait anxiety. These findings demonstrated that personality differences can shape processing of interactions between relevant social signals.     

Top-down influences make saccades deviate away: the case of endogenous cues

We tested a recent hypothesis suggesting that the eye deviates away from a location when top-down preparation can influence target selection. Participants had to make an eye movement to a peripheral target. Before the upcoming target, a central cue indicated the likely target location. Results show that when the target was presented at a location different from that indicated by the cue, eye movements to the target deviated away from the cued location. Because central cues are under top-down control, the present results are in line with a determining role of top-down preparation on saccade direction. These results contrast with the findings reported in a similar paradigm executed with hand movements, in which the movements were mostly initiated in the direction of the cued location. Therefore, we conclude that inhibitory effects typically observed when executing eye movements may not be observed when executing hand movements in similar conditions.     

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.     

Effects of direct and averted gaze on the subsequent saccadic response

The saccadic latency to visual targets is susceptible to the properties of the currently fixated objects. For example, the disappearance of a fixation stimulus prior to presentation of a peripheral target shortens saccadic latencies (the gap effect). In the present study, we investigated the influences of a social signal from a facial fixation stimulus (i.e., gaze direction) on subsequent saccadic responses in the gap paradigm. In Experiment 1, a cartoon face with a direct or averted gaze was used as a fixation stimulus. The pupils of the face were unchanged (overlap), disappeared (gap), or were translated vertically to make or break eye contact (gaze shift). Participants were required to make a saccade toward a target to the left or the right of the fixation stimulus as quickly as possible. The results showed that the gaze direction influenced saccadic latencies only in the gaze shift condition, but not in the gap or overlap condition; the direct-to-averted gaze shift (i.e., breaking eye contact) yielded shorter saccadic latencies than did the averted-to-direct gaze shift (i.e., making eye contact). Further experiments revealed that this effect was eye contact specific (Exp. 2) and that the appearance of an eye gaze immediately before the saccade initiation also influenced the saccadic latency, depending on the gaze direction (Exp. 3). These results suggest that the latency of target-elicited saccades can be modulated not only by physical changes of the fixation stimulus, as has been seen in the conventional gap effect, but also by a social signal from the attended fixation stimulus.     

The role of the adjacency between background cues and objects in visual localization during ocular pursuit

Subjects used eye movements to pursue a light target that moved from left to right with a velocity of 15 deg s-1. The stimulus was a sudden five-fold decrease in target intensity during the movement. The subject's task was to localize the stimulus relative to either a single stationary background point or the midpoint between two points (28 deg apart) placed 0.5 deg above the target path. The stimulus was usually mislocated in the direction of eye movement; the mislocation was affected by the spatial adjacency between background and stimulus. When an auditory, rather than a visual, stimulus was presented during tracking, target position at the time of stimulus presentation was visually mislocated in the direction opposite to that of eye movement. The effect of adjacency between background and target remained the same. The involvement of processes of subject-relative and object-relative visual perception is discussed.     

Spatial but not temporal cueing influences the mislocalisation of a target flashed during smooth pursuit

Human subjects misjudge the position of a target that is flashed during a pursuit eye movement. Their judgments are biased in the direction in which the eyes are moving. We investigated whether this bias can be reduced by making the appearance of the flash more predictable. In the normal condition, subjects pursued a moving target that flashed somewhere along its trajectory. After the presentation, they indicated where they had seen the flash. The mislocalisations in this condition were compared to mislocalisations in conditions in which the subjects were given information about when or where the flash would come. This information consisted of giving two warning flashes spaced at equal intervals before the target flash, of giving two warning beeps spaced at equal intervals before the target flash, or of showing the same stimulus twice. Showing the same stimulus twice significantly reduced the mislocalisation. The other conditions did not. We interpret this as indicating that it is not predictability as such that influences the performance, but the fact that the target appears at a spatially cued position. This was supported by a second experiment, in which we examined whether subjects make smaller misjudgments when they have to determine the distance between a target flashed during pursuit and a reference seen previously, than when they have to determine the distance between the flashed target and a reference seen afterwards. This was indeed the case, presumably because the reference provided a spatial cue for the flash when it was presented first. We conclude that a spatial cue reduces the mislocalisation of targets that are flashed during pursuit eye movements. The cue does not have to be exactly at the same position as the flash.     

Fixational eye movements are not an index of covert attention

The debate about the nature of fixational eye movements has revived recently with the claim that microsaccades reflect the direction of attentional shifts. A number of studies have shown an association between the direction of attentional cues and the direction of microsaccades. We sought to determine whether microsaccades in attentional tasks are causally related to behavior. Is reaction time (RT) faster when microsaccades point toward the target than when they point in the opposite direction? We used a dual-Purkinje-image eyetracker to measure gaze position while 3 observers (2 of the authors, 1 naive observer) performed an attentional cuing task under three different response conditions: saccadic localization, manual localization, and manual detection. Critical trials were those on which microsaccades moved away from the cue. On these trials, RTs were slower when microsaccades were oriented toward the target than when they were oriented away from the target. We obtained similar results for direction of drift. Cues, not fixational eye movements, predicted behavior.     

Where a person with a squint is actually looking: Gaze-cued orienting in crossed eyes

In this study, we investigated gaze-cued attention orienting when the perceived eyes are not looking in the same direction. This condition occurs in strabismus (squint). Participants were asked to detect laterally presented reaction signals preceded by schematic faces in which the direction (left, straight, or right) of the left and right eye was independently manipulated. Consistent with earlier studies, the results showed a reliable cuing effect by two eyes with parallel gaze direction. Gaze-cued orienting was also shown in a situation when one eye was averted and the other eye was looking straight ahead. The gaze cuing was not significantly stronger in the former than in the latter situation. When both eyes were either nasally or temporally averted, no shifts of visual attention were observed. The results suggest that, if both eyes are visible, the direction of both eyes is computed and integrated for the gaze-cued orienting.