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.     

Delay of stimulus presentation after the saccade in visual search

Delay of stimulus onset after each saccade in visual search decreased oculomotor and manual reaction times, with a greater effect occurring for the oculomotor response. The saccadic oculomotor reaction might have been facilitated in three ways: by the facilitation of reaction with a foreperiod warning stimulus, by the attenuation of saccadic suppression effects due to the stimulus onset delay, or by the use of a strategy of preprogramming fixation durations. The results support a model of visual search using preprogrammed control of visual fixation durations.     

Saccadic facilitation by modulation of microsaccades in natural backgrounds

Saccades move objects of interest into the center of the visual field for high-acuity visual analysis. White, Stritzke, and Gegenfurtner (Current Biology, 18, 124-128, 2008) have shown that saccadic latencies in the context of a structured background are much shorter than those with an unstructured background at equal levels of visibility. This effect has been explained by possible preactivation of the saccadic circuitry whenever a structured background acts as a mask for potential saccade targets. Here, we show that background textures modulate rates of microsaccades during visual fixation. First, after a display change, structured backgrounds induce a stronger decrease of microsaccade rates than do uniform backgrounds. Second, we demonstrate that the occurrence of a microsaccade in a critical time window can delay a subsequent saccadic response. Taken together, our findings suggest that microsaccades contribute to the saccadic facilitation effect, due to a modulation of microsaccade rates by properties of the background.     

Illusory motion produced by dichoptic stimuli during saccades

A new motion illusion is reported in which saccadic eye movements can produce a perceived jump of a static stimulus presented dichoptically. In three experiments, observers made saccades while viewing a stationary stimulus consisting of a disk and random dots presented separately to the two eyes. In experiments 1 and 2, by measuring the strength of the perceived motion and the velocity of binocular eye movements, we found that (a) motion ratings were high for the stimulus that contained a large interocular difference in luminance, and (b) the saccadic strategy of the observer was virtually identical across different stimulus conditions. In experiment 3, by measuring the detectability of a short temporal gap introduced into the stimulus around saccades, we found that saccadic suppression was normal in the dichoptic presentation. We discuss possible mechanisms underlying the illusory motion.     

Fixation disengagement and eye-movement latency

We examined eye-movement latencies to a target that appeared during visual fixation of a stationary stimulus, a moving stimulus, or an extrafoveal stimulus. The stimulus at fixation was turned off either before target onset (gap condition) or after target onset (overlap condition). Consistent with previous research, saccadic latencies were shorter in gap conditions than they were in overlap conditions (the gap effect). In Experiment 1, a gap effect was observed for vergence eye movements. In Experiment 2, a gap effect was observed for saccades directed at a target that appeared during visual pursuit of a moving stimulus. In Experiment 3, a gap effect was observed for saccades directed at a target that appeared during extrafoveal fixation. The present results extend reports of the gap effect for saccadic shifts during visual fixation to (a) vergence shifts during visual fixation, (b) saccadic shifts during smooth visual pursuit, and (c) saccadic shifts during extrafoveal fixation. The present findings are discussed with respect to the incompatible goals of fixation-locking and fixation-shifting oculomotor responses.     

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.     

Attention and saccadic eye movements

Four threshold detection experiments addressed three issues concerning the relationship between movements of spatial attention and saccadic eye movements: (a) the time course of attention shifts wit saccades, (b) the response of the two systems to changes in stimulus parameters, and (c) the relationship of attention to saccadic suppression. These issues bear on the more general question of the degree of independence between the saccadic and attentional movement systems. The results of these experiments support the contention that the mechanisms that shift attention are separate from those that control saccadic eye movements. Relevant events in the visual field periphery, however, will trigger both a saccade and attention shift. The attentional response to such events does not appear to be under subjects' control. The implication of these results for theories of saccadic suppression is discussed.     

Probing the influence of unconscious fear-conditioned visual stimuli on eye movements

Efficient threat detection from the environment is critical for survival. Accordingly, fear-conditioned stimuli receive prioritized processing and capture overt and covert attention. However, it is unknown whether eye movements are influenced by unconscious fear-conditioned stimuli. We performed a classical fear-conditioning procedure and subsequently recorded participants’ eye movements while they were exposed to fear-conditioned stimuli that were rendered invisible using interocular suppression. Chance-level performance in a forced-choice-task demonstrated unawareness of the stimuli. Differential skin conductance responses and a change in participants’ fearfulness ratings of the stimuli indicated the effectiveness of conditioning. However, eye movements were not biased towards the fear-conditioned stimulus. Preliminary evidence suggests a relation between the strength of conditioning and the saccadic bias to the fear-conditioned stimulus. Our findings provide no strong evidence for a saccadic bias towards unconscious fear-conditioned stimuli but tentative evidence suggests that such an effect may depend on the strength of the conditioned response.     

The gap effect for eye and hand movements

A temporal gap between fixation point offset and stimulus onset typically yields shorter saccadic latencies to the stimulus than if the fixation stimulus remained on. Several researchers have explored the extent to which this gap also reduces latencies of other responses but have failed to find a gap effect isolated from general warning effects. Experiment 1, however, showed a robust gap effect for aimed hand movements (which required determination of a precise spatial location), regardless of whether the hand moved alone or was accompanied by a saccadic eye movement. Experiment 2 replicated this aimed hand gap effect and also showed a smaller effect for choice manual keypress responses (which required determination of the direction of response only). Experiment 3 showed no gap effect for simple manual keypress responses (which required no spatial determination). The results are consistent with an interpretation of the gap effect in terms of facilitation of spatially oriented responses.     

Asymmetry in infant hemispheric readiness after exposure to a visual stimulus

This experiment assessed whether prior exposure to one visual stimulus could result in differential hemispheric responsiveness to a subsequent visual stimulus. The latency of saccadic orientation to a star‐shaped stimulus in the left or right visual field was assessed for 24 infants (mean age 22 weeks, SD 4.5) after exposure to either an upright or inverted facial pattern in the central visual field. The response to the lateral stimulus was equivalent in either visual field after exposure to the inverted facial pattern, but was significantly slower (p = 0.043) in the LVF (RH) than the RVF (LH) following presentation of the upright facial pattern. The outcome confirms that the processing of one visual stimulus may lead to differential hemispheric readiness to engage with a subsequent visual stimulus.     

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.     

Conditioned suppression, punishment, and aversion

Three experiments were conducted to assess the aversive properties of a visual stimulus in the presence of which one group of birds received response-contingent shock (discriminated punishment) while a yoked group of birds received non-contingent shocks (conditioned suppression). In Experiment 1, presentation of the visual stimulus contingent on key pecking reduced the response rate (conditioned punishment effect) for birds under the conditioned suppression procedure but did not reduce the response rate of birds under the discriminative punishment procedure. Non-contingent shocks also produced greater suppression of responding maintained by positive reinforcement in the presence of a visual stimulus than did response-contingent shocks. In Experiment 2, a greater shock intensity (2 mA) was used. All the differences between the two groups found in Experiment 1 were also found in Experiment 2. Experiment 3 demonstrated that response-contingent shock did not result in a conditioned punishment effect even when positive reinforcers were unavailable during the discriminative punishment schedule. The exteroceptive stimulus that was paired with shock in the conditioned suppression procedure acquired the ability to punish behavior. The exteroceptive stimulus in the discriminative punishment schedule did not acquire this ability.     

The reduction of saccadic latency by prior offset of the fixation point: an analysis of the gap effect

The latency to initiate a saccade (saccadic reaction time) to an eccentric target is reduced by extinguishing the fixation stimulus prior to the target onset. Various accounts have attributed this latency reduction (referred to as the gap effect) to facilitated sensory processing, oculomotor readiness, or attentional processes. Two experiments were performed to explore the relative contributions of these factors to the gap effect. Experiment 1 demonstrates that the reduction in saccadic reaction time (RT) produced by fixation point offset is additive with the effect of target luminance. Experiment 2 indicates that the gap effect is specific for saccades directed toward a peripheral target and does not influence saccades directed away from the target (i.e., antisaccades) or choice-manual RT. The results are consistent with an interpretation of the gap effect in terms of facilitated premotor processing in the superior colliculus.     

Evidence for visual persistence during saccadic eye movements

Summary 1. The persistence of visual perception was investigated under conditions of visual fixation as well as eye movement. The Ss' task was to discriminate brief double light impulses; their responses were recorded as a function of the duration of the interstimulus interval. Based on these data the critical interstimulus interval was calculated, which yielded equal response frequencies for the perception of one or two stimuli upon presentation of double light pulses.2. In the condition of visual fixation the two stimuli could not be discriminated until the mean value of interstimulus interval exceeded 73 msec. In the condition with eye movements, when the first stimulus was presented in the parafoveal region of the retina before the beginning of the saccade and the second stimulus in the foveal region just after termination of the eye movement, this duration was shown to be statistically of the same magnitude (76 msec).3. Possible alternative interpretations of this latter result, e.g., that it could be explained in terms of masking or saccadic suppression rather than visual persistence was discussed; it was attempted to invalidate such explanations by means of three control experiments.4. The main result, the persistence of visual perception during voluntary eye movements, was discussed in relation to the problem of spatial and temporal stability of visual perception.I thank Prof. Dr. H.W. Wendt for support in correcting the English translation.     

眼跳的双相调节理论的行为证据

为了研究眼跳的双相调节理论是否适用于人类的视觉系统,本研究测量了人类被试对分别呈现在三种眼跳时间段（基线、眼跳抑制和眼跳增强）内的光栅的朝向辨别准确率。研究发现：相对于光栅呈现在基线时间段内,被试对呈现在抑制（或增强）时间段内的光栅的朝向辨别准确率显著地更低（或更高）（实验1）;另外,只有使用低或中等空间频率光栅作为测试刺激时,才有这种双相调节作用（实验2）。这些结果表明：人类的视觉系统在眼跳过程中存在双相调节机制,并且这种双相调节机制具有刺激选择性。     

Giving subjects the eye and showing them the finger: socio-biological cues and saccade generation in the anti-saccade task

Pointing with the eyes or the finger occurs frequently in social interaction to indicate direction of attention and one's intentions. Research with a voluntary saccade task (where saccade direction is instructed by the colour of a fixation point) suggested that gaze cues automatically activate the oculomotor system, but non-biological cues, like arrows, do not. However, other work has failed to support the claim that gaze cues are special. In the current research we introduced biological and non-biological cues into the anti-saccade task, using a range of stimulus onset asynchronies (SOAs). The anti-saccade task recruits both top-down and bottom-up attentional mechanisms, as occurs in naturalistic saccadic behaviour. In experiment 1 gaze, but not arrows, facilitated saccadic reaction times (SRTs) in the opposite direction to the cues over all SOAs, whereas in experiment 2 directional word cues had no effect on saccades. In experiment 3 finger pointing cues caused reduced SRTs in the opposite direction to the cues at short SOAs. These findings suggest that biological cues automatically recruit the oculomotor system whereas non-biological cues do not. Furthermore, the anti-saccade task set appears to facilitate saccadic responses in the opposite direction to the cues.     

The role of the gap effect in the orienting of attention: Evidence for express attentional shifts

The “gap effect” refers to the finding that saccadic latencies are typically reduced when the fixation point is removed just prior to the presentation of a target. One explanation for this effect is that the removal of the fixation point causes the disengagement of covert attention and allows for extremely rapid movements of attention (express attentional shifts). However, previous research regarding express attentional shifts has yielded equivocal results. The present study used a variation of a peripheral cueing paradigm with a discrimination task (Experiment 1) and a detection task (Experiment 2) to further examine this issue. The results from eye movement and keypress latencies indicated that there were express attentional shifts with the discrimination task but not in the detection task. This pattern of results may have been due to differences in how attention was allocated between the two tasks. Thus, evidence for express attentional shifts was found, but only under certain conditions.     

Control of reflexive and voluntary saccades in the gap effect

In two experiments, we examined whether voluntary and reflexive saccades shared a common fixation disengagement mechanism, Participants were required to perform a variety of tasks, each requiring a different level of information processing of the display prior to execution of the saccade. In Experiment 1, participants executed either a prosaccade or an antisaccade upon detecting a stimulus array. In Experiment 2, participants executed a prosaccade to a stimulus array only if the array contained a target item. The target could be a line (easy search) or a digit (difficult search). The critical manipulation in both experiments was the relative timing between the removal of the fixation stimulus and the onset of the stimulus array. In both experiments, it was found that saccadic latencies were shortest when the fixation stimulus was removed before the onset of the stimulus array—a gap effect. It was concluded that reflexive and voluntary saccades share a common fixation disengagement mechanism that is largely independent of higher level cognitive processes.     

Control' of reflexive and voluntary saccades in the gap effect.

In two experiments, we examined whether voluntary and reflexive saccades shared a common fixation disengagement mechanism. Participants were required to perform a variety of tasks, each requiring a different level of information processing of the display prior to execution of the saccade. In Experiment 1, participants executed either a prosaccade or an antisaccade upon detecting a stimulus array. In Experiment 2, participants executed a prosaccade to a stimulus array only if the array contained a target item. The target could be a line (easy search) or a digit (difficult search). The critical manipulation in both experiments was the relative timing between the removal of the fixation stimulus and the onset of the stimulus array. In both experiments, it was found that saccadic latencies were shortest when the fixation stimulus was removed before the onset of the stimulus array--a gap effect. It was concluded that reflexive and voluntary saccades share a common fixation disengagement mechanism that is largely independent of higher level cognitive processes.     

Infant attention and the development of smooth pursuit tracking.

The effect of attention on smooth pursuit and saccadic tracking was studied in infants at 8, 14, 20, and 26 weeks of age. A small rectangle was presented moving in a sinusoidal pattern in either the horizontal or vertical direction. Attention level was distinguished with a recording of heart rate. There was an increase across age in overall tracking, the gain of the smooth pursuit eye movements, and an increase in the amplitude of compensatory saccades at faster tracking speeds. One age change was an increase in the preservation of smooth pursuit tracking ability as stimulus speed increased. A second change was the increasing tendency during attentive tracking to shift from smooth pursuit to saccadic tracking when the stimulus speed increased to the highest velocities. This study shows that the development of smooth pursuit and targeted saccadic eye movements is closely related to the development of sustained attention in this age range.