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.     

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.     

What are human express saccades?

When a fixation point is removed 200 msec prior to target onset (the gap condition), human subjects are said to produce eye movements that have a short latency (80–120 msec), that form the early peak of a bimodal latency distribution, and that have been labeled “human express saccades” (see, e.g., Fischer, 1987; Fischer & Breitmeyer, 1987; Fischer & Ramsperger, 1984, 1986). In three experiments, we sought to obtain this express saccade diagnostic pattern in the gap condition, We orthogonally combined target location predictability with the presence versus absence of catch trials (Experiment 1). When target location was fixed and catch trials were not used, we found mostly anticipations. In the remaining conditions, where responses were under stimulus control, bimodality was not frequently observed, and, whether it was or not, latencies were not in the express saccade range. Using random target locations, we then varied stimulus luminance and the mode of stimulus presentation (LEDs vs. oscilloscope) in the gap and overlap (fixation is not removed) conditions (Experiment2). Bimodality was rarely observed, the gap effect (overlap minus gap reaction time) was additive with luminance, and only the brightest targets elicited saccades in the express range. When fixed locations and no catch trials were combined with latency feedback (Experiment 3), we observed many responses in the express saccade range and some evidence for bimodality, but the sudden introduction of catch trials revealed that many early responses were not under stimulus control. Humanscan make stimulus-controlled saccades that are initiated very rapidly (80–120 msec), but unless catch trials or choice reaction time is used, it is not possible to distinguish such saccades from anticipatory responses that are prepared in advance and timed to occur shortly after target onset. Because the express saccade diagnostic pattern is not a characteristic feature of human saccadic performance, we urge investigators to focus their attention on the robustgap effect     

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.     

Are microsaccades responsible for the gap effect?

Extinguishing a fixation point shortly before, or concurrently with, the onset of a peripheral visual target reduces the latency of saccades to that target. Saslow (1967) hypothesized that thisgap effect might occur because fixation point offsets reduce the incidence of corrective microsaccades with an associated saccadic refractory period. In the present study, a robust gap effect was obtained. However, using a Purkinje image eyetracker with 1 arcmin of resolution, we found that fixation point offsets had no effect on the occurrence of microsaccades and that the occurrence of microsaccades had no impact on the magnitude of the gap effect. Microsaccades therefore do not appear to play any part in the production of the gap effect.     

Presaccadic attention allocation and express saccades

Express saccades are visually-guided saccades that are characterized by an extremely short latency of about 100 ms. The present experiments tested the hypothesis that a disengagement of visual attention is necessary for the generation of express saccades. All subjects produced large numbers of express saccades in the gap paradigm, in which the fixation stimulus is removed 200 ms before target onset (Exp. 1), but not in the overlap paradigm, in which the fixation stimulus remains on during the entire trial (Exp. 2). By means of peripheral cues (Exps. 3–5) and central cues (Exps. 6–7), visual attention was directed at the target location for the saccade before the actual appearance of the saccade target. In all experiments, the location cues facilitated rather than abolished express saccades. The generation of express saccades was facilitated even when the currently fixated visual stimulus was not removed before target onset (fixation-overlap; Exps. 5–7). The results are explained by the hypothesis that a disengagement of a separate fixation system is necessary for the generation of express saccades, a hypothesis that is in line with current neurobiological findings.     

Response times and eye movements in feature and conjunction search as a function of target eccentricity

In four experiments, saccadic eye movements, reaction times (RTs), and accuracy were measured as observers searched for feature or conjunction targets presented at several eccentricities. A conjunction search deficit, evidenced by a large eccentricity effect on RTs, accuracy, and number of saccades, was seen in Experiments 1A and 1B. Experiment 2 indicated that, when saccades were precluded, there was an even larger eccentricity effect for conjunction search targets. In Experiment 3, practice in a conjunction search task allowed both RT and number of saccades to become independent of eccentricity. Additionally, there was evidence of feature-based selectivity in that observers were more likely to fixate distractors that had the same contrast as the target. Results are consistent with the view that the oculomotor and attentional systems are functionally linked and provide constraints for models of visual attention and search.     

Control of fixation duration in visual search and memory search: another look

In visual search a variable delay (up to 150 msec) between the beginning of each fixation and the onset of a search stimulus reduces the time (oculomotor latency) between stimulus onset and the subject's next saccadic eye movement. Two hypotheses for this effect of stimulus onset delay (SOD) were compared: first, process monitoring, that SOD simply serves as a warning interval to facilitate saccadic responses; and second, preprogramming, that saccades are preprogrammed at short SODs. In the first experiment SOD produced a decline in oculomotor latency in search similar to that seen in previous studies. In the second and third experiments, the size of the memory set in a Sternberg memory search paradigm was varied, or a mask flanking some of the search stimuli was used, to vary the processing time of each stimulus. Partial preprogramming of saccades at short delays would predict that increasing the processing time of individual stimuli would increase oculomotor latency at only short SODs. However, oculomotor latency increased equally at all SODs. In this search task, then, the SODs appeared to facilitate saccade initiation.     

Surface and Edge Information for Spatial Integration: A Saccadic-selection Search Task

Three experiments are described that investigate visual integration across space using a saccade selection paradigm. Subjects saccaded to a vertical target in the presence of a number of horizontal distractor items. Both horizontal and vertical items were composed of two identical elements, so, in order to locate the target, subjects had to integrate the pairs of elements together. We measured saccade accuracy, the proportion of saccades directed to the vertical target, together with saccade latency following display appearance. In Experiment 1 we found that saccade accuracy was improved by the items having either common surface properties or collinear edges. These effects were not a resultof increased display heterogeneity (Experiment 2), or a result of the introduction of a strong internal contour between the items (Experiment 3). These results show that for saccadic selection both surface and edge properties of items are processed. This in turn suggests thatearly visual processing encodes and exploits both types of information.     

Against a role for attentional disengagement in the gap effect: A friendly amendment to Tam and Stelmach (1993)

Saccadic reaction time (RT) is reduced when the fixation point is removed shortly before target onset. Although Tam and Stelmach (1993) argued that thisgap effect could not be explained solely by the idea that fixation offset disengaged visual attention and preferred an explanation based on disengagement of the oculomotor system, they felt that they could not rule out a hybrid model in which both oculomotorand attentional disengagement contribute to the gap effect. Our analysis of the dual response experiment (Experiment 4), upon which this hybrid model was based, shows that manual and saccadic responses were likely compromised by a grouping or delay strategy and that subjects may not have been attending as instructed. On these grounds, we argue that Tam and Stelmach (1993), like Kingstone and Klein (1990; 1993a) provide no evidence that attentional disengagement contributes to the gap effect. An alternative proposal (Klein & Kingstone, 1993), that motor preparation and oculomotor disengagement combine additively to produce the gap effect, is consistent with the data from Tam and Stelmach’s Experiments 1–3, is similar to the explanation that they prefer, and has been strongly supported when directly tested (Kingstone, Klein, & Taylor, 1994).     

Saccades to targets in three-dimensional space: Dependence of saccadic latency on target location

The latency of saccadic movements to targets appearing at various positions in three-dimensional visual space was measured in four experiments. The first experiment confirmed that latencies of saccades to visual targets are greater in the lower visual field and showed that the increase is not influenced by the vertical starting position of the eye in the orbit, nor by a time gap between the fixation offset and the target onset. A hypothesis that this visual field difference was caused by a link between downward saccades and convergence movements was tested by recording saccade latencies when the targets were in a different depth plane from that of the original fixation. We did not find any direct support for the vergence involvement hypothesis, although the lower/upper visual field effect was shown to decrease consistently in monocular viewing. It was also shown that saccades to targets positioned in a different depth plane have longer latencies. In a final experiment, the visual field effect was shown to depend on the egocentric rather than the gravitational vertical.     

Saccades to targets in three-dimensional space: dependence of saccadic latency on target location.

The latency of saccadic movements to targets appearing at various positions in three-dimensional visual space was measured in four experiments. The first experiment confirmed that latencies of saccades to visual targets are greater in the lower visual field and showed that the increase is not influenced by the vertical starting position of the eye in the orbit, nor by a time gap between the fixation offset and the target onset. A hypothesis that this visual field difference was caused by a link between downward saccades and convergence movements was tested by recording saccade latencies when the targets were in a different depth plane from that of the original fixation. We did not find any direct support for the vergence involvement hypothesis, although the lower/upper visual field effect was shown to decrease consistently in monocular viewing. It was also shown that saccades to targets positioned in a different depth plane have longer latencies. In a final experiment, the visual field effect was shown to depend on the egocentric rather than the gravitational vertical.     

Saccadic reaction times of dyslexic and age-matched normal subjects

Saccadic reaction times (SRT) were measured in a simple task: subjects had to make saccades from a central fixation point to peripheral targets, which appeared randomly 4 deg to the left or to the right. In the first test the fixation point went off before the target appeared (gap trials); in the second test it remained on the screen (overlap trials). The distribution of SRTs for trained normal adults (N = 4), untrained normal adults (N = 11), untrained normal children (aged 9-11 years, N = 9), untrained normal teenagers (aged 15-17 years, N = 8), dyslexic children (aged 9-11 years, N = 15), and dyslexic teenagers (aged 15-17 years, N = 5) were compared, with special emphasis on the number of express saccades, ie saccades with extremely short reaction times (100-120 ms, under the present conditions). In normal adults with the gap paradigm, the distribution of reaction times typically exhibits two or three modes (express saccades, fast regular saccades, and very few slow regular saccades), whereas in the overlap paradigm only one or two modes (few fast regular saccades and many slow regular saccades) are obtained. On average, normal children produce more express saccades than naive normal adults. Dyslexic children produce more express saccades than the normal age-matched controls. Among the dyslexic children, four different types of abnormalities in their reaction times were encountered. The group of dyslexic teenagers was characterized by a larger number of express saccades at the expense of fast regular saccades in gap trials and by fewer express saccades and fewer fast regular saccades in overlap trials when compared to the age-matched control group. It is concluded that the abnormal patterns of saccadic reaction times reflect defects in the system of visual attention and/or in its control over the oculomotor system, rather than indicating a defect in the oculomotor system itself. In this context, symptoms of dyslexia appear as a combination of attentional deficits and irregular timing of saccadic eye movements.     

Top-down control in time and space: Evidence from saccadic latencies and trajectories

Visual distractors disrupt the production of saccadic eye movements temporally, by increasing saccade latency, and spatially, by biasing the trajectory of the movement. The present research investigated the extent to which top-down control can be exerted over these two forms of oculomotor capture. In two experiments, people were instructed to make target directed saccades in the presence of distractors, and temporal and spatial capture were assessed simultaneously by measuring saccade latency and saccade trajectory curvature, respectively. In Experiment 1, an attentional control set manipulation was employed, resulting in the elimination of temporal capture, but only an attenuation of spatial capture. In Experiment 2, foreknowledge of the target location caused an attenuation of temporal capture but an enhancement of spatial capture. These results suggest that, whereas temporal capture is contingent on top-down control, the spatial component of capture is stimulus-driven.     

Spatial and temporal factors determine auditory-visual interactions in human saccadic eye movements

In this paper, we show that human saccadic eye movements toward a visual target are generated with a reduced latency when this target is spatially and temporally aligned with an irrelevant auditory nontarget. This effect gradually disappears if the temporal and/or spatial alignment of the visual and auditory stimuli are changed. When subjects are able to accurately localize the auditory stimulus in two dimensions, the spatial dependence of the reduction in latency depends on the actual radial distance between the auditory and the visual stimulus. If, however, only the azimuth of the sound source can be determined by the subjects, the horizontal target separation determines the strength of the interaction. Neither saccade accuracy nor saccade kinematics were affected in these paradigms. We propose that, in addition to an aspecific warning signal, the reduction of saccadic latency is due to interactions that take place at a multimodal stage of saccade programming, where theperceived positions of visual and auditory stimuli are represented in a common frame of reference. This hypothesis is in agreement with our finding that the saccades often are initially directed to the average position of the visual and the auditory target, provided that their spatial separation is not too large. Striking similarities with electrophysiological findings on multisensory interactions in the deep layers of the midbrain superior colliculus are discussed.     

Attention modulates saccadic inhibition magnitude

Visual transient events during ongoing eye movement tasks inhibit saccades within a precise temporal window, spanning from around 60–120 ms after the event, having maximum effect at around 90 ms. It is not yet clear to what extent this saccadic inhibition phenomenon can be modulated by attention. We studied the saccadic inhibition induced by a bright flash above or below fixation, during the preparation of a saccade to a lateralized target, under two attentional manipulations. Experiment 1 demonstrated that exogenous precueing of a distractor's location reduced saccadic inhibition, consistent with inhibition of return. Experiment 2 manipulated the relative likelihood that a distractor would be presented above or below fixation. Saccadic inhibition magnitude was relatively reduced for distractors at the more likely location, implying that observers can endogenously suppress interference from specific locations within an oculomotor map. We discuss the implications of these results for models of saccade target selection in the superior colliculus.     

Effect of stimulus intensity on manual and saccadic reaction time

Reaction time (RT) decreases with stimulus intensity. Hughes and Kesley (1984) demonstrated, however, that the effect of stimulus intensity on simple RT is larger for manual than for saccadic responses. We reexamined this relation under various conditions. The dissociation occurred when the task enabled the generation of exogenous saccades. We found, however, no dissociation if endogenous saccades had to be executed. It is hypothesized that the different effects of intensity result from the simplified neuronal processing of exogenous saccades performed in the direct route from the retina to the superior colliculus.     

Fixation-point offsets reduce the latency of saccades to acoustic targets.

If an observer's fixation point is extinguished just prior to the onset of a peripheral target, the latency to saccade to that target is reduced. We show that this "gap effect" is not specific to visual targets. Observers made saccades to a light flash or to a white-noise burst. A warning tone was presented on every trial to control for the possible warning effect of the fixation-point offset. For both target modalities, saccade latencies were significantly reduced when the fixation point was extinguished 200 msec prior to the target onset. Implications of this outcome for interpretations of the gap effect are considered. It is argued that the presence of a gap effect for tones, in conjunction with previous findings, is consistent with the hypothesis that the gap effect is produced by a facilitation of premotor processes in the superior colliculus.     

Fixation-point offsets reduce the latency of saccades to acoustic targets

If an observer’s fixation point is extinguished just prior to the onset of a peripheral target, the latency to saccade to that target is reduced. We show that this "gap effect" is not specific to visual targets. Observers made saccades to a light flash or to a white-noise burst. A warning tone was presented on every trial to control for the possible warning effect of the fixation-point offset. For both target modalities, saccade latencies were significantly reduced when the fixation point was extinguished 200 msec prior to the target onset. Implications of this outcome for interpretations of the gap effect are considered. It is argued that the presence of a gap effect for tones, in conjunction with previous findings, is consistent with the hypothesis that the gap effect is produced by a facilitation of premotor processes in the superior colliculus.