Effect of stimulus probability on anti-saccade error rates

Subjects sometimes fail to suppress a reflexive saccade towards the flashed stimulus in an anti-saccade task. Here, we studied how error rates in the anti-saccade task varied as a function of saccadic probability. Ten subjects performed 200 anti-saccade trials for each of three saccade-direction probability conditions (20%, 50%, and 80%). We found that as the likelihood of a saccade in a given direction increased, the percentage of pro-saccade errors also increased for stimulus presentations in this direction. These results provide support for the hypothesis that errors in the anti-saccade task are the result of an increased level of motor preparation.     

Effects of pre-cues on voluntary and reflexive saccade generation I. Anti-cues for pro-saccades

Experiments on visual attention have employed both physical cues and verbal instructions to enable subjects to allocate attention at a location that becomes relevant within a perceptual or motor task some time later (cue lead time, CLT). In this study we have used valid visual peripheral cues (CLT between 100 and 700 ms) to indicate the direction and location of the next saccade. A cue is considered valid or invalid if its meaning with respect to the next saccade is correct or incorrect. A cue is called an anti- or pro-cue if the side of its presentation is opposite to or the same as the direction of the saccade required on a given trial. Correspondingly, a saccade is called an anti- or pro-saccade if it is directed to the side opposite to or the same as the stimulus presentation. A condition in which the cue and the stimulus are presented on opposite sides provides a simple way of dissociating voluntary attention allocation from automatic orienting. This paper considers the anti-cue pro-saccade task: the subjects were instructed to use the cue to direct attention to the opposite side, i.e. the location, where on valid trials the saccade target would occur. In the companion paper we have used the same physical condition, but we have reversed the instructions as to saccade direction and we have reversed the meaning of the cue, i.e. we designed a pro-cue anti-saccade task. In this first paper, the saccadic reaction times (SRTs) of pro-saccades of five adult subjects were measured in the gap paradigm (fixation point offset precedes target onset by 200 ms). With a CLT of 100 ms, valid anti-cues reduced the number of express saccades (i.e. saccades with SRTs in the range 80–120 ms) significantly compared with the control values (no cues). Valid anti-cues with increasingly long CLTs (100–700 ms) resulted in an increasing incidence of anticipatory saccades and saccades with longer SRTs (more than 120 ms), while the frequency of express saccades remained below the control value. When cue and saccade target were dissociated in location or in both location and direction, the effects of the cueing revealed a much lower spatial selectivity as compared to the effects that have been described for voluntary attention allocation by means of central cues. The results suggest that voluntary allocation of attention and cue-induced automatic orienting not only have different time courses but also have opposite effects on the generation of express saccades, and different spatial selectivities. A possible neuronal basis of these results is discussed considering related findings from electrophysiological studies in monkeys. Received: 27 March 1997 / Accepted: 17 December 1997     

Are somatosensory saccades voluntary or reflexive?

The present study examines whether the distinction between voluntary (endogenous) and reflexive (stimulus-elicited) saccades made in the visual modality can be applied to the somatosensory modality. The behavioural characteristics of putative reflexive pro-saccades and voluntary anti-saccades made to visual and somatosensory stimuli were examined. Both visual and somatosensory pro-saccades had much shorter latency than voluntary anti-saccades made in the direction opposite to a peripheral stimulus. Furthermore, erroneous pro-saccades were made towards both visual and somatosensory stimuli on approximately 11-13% of anti-saccade trials. The observed difference in pro- and anti-saccade latency and the presence of pro-saccade errors in the anti-saccade task indicates that a somatosensory stimulus can elicit a form of reflexive saccade comparable to pro-saccades made in the visual modality. It is proposed that a peripheral somatosensory stimulus can elicit a form of reflexive saccade and that somatosensory saccades do not depend exclusively on higher level endogenous control processes for their generation. However, a comparison of the underlying latency distributions and of peak-velocity profiles of saccades made to visual and somatosensory stimuli showed that this distinction may be less clearly defined for the somatosensory modality and that modality-specific differences (such as differences in neural conduction rates) in the underlying oculomotor structures involved in saccade target selection also need to be considered. It is further suggested that a broader conceptualisation of saccades and saccade programming beyond the simple voluntary and reflexive dichotomy, that takes into account the control processes involved in saccade generation for both modalities, may be required.     

Effects of pre-cues on voluntary and reflexive saccade generation II. Pro-cues for anti-saccades

The reaction times of saccades (SRT) to a suddenly presented visual stimulus (pro-saccade) can be decreased and a separate mode of express saccades can occur when a gap paradigm is used (i.e. fixation-point offset precedes target onset by 200 ms). A valid peripheral cue, presented briefly (100 ms) before target onset, has been found to facilitate the generation of saccades to the target, thereby increasing the frequency of express saccades and decreasing the mean latency. This facilitation occurs only for cues that correctly indicate the direction of the subsequent target presentation (valid cues). The present study investigates the effects of valid cues on SRTs and error rate in the anti-saccade task (saccades in the direction opposite to the stimulus) by systematically varying the cue lead time (CLT) and using the gap and overlap conditions, i.e. fixation point remains on throughout the trial. For a CLT of 100 ms, both reaction times and error rates were significantly increased. With increasing CLT (200–500 ms), both the reaction times of the anti-saccades and the error rates returned to approximately control level, with CLT more than 200 ms in both the gap and the overlap condition. Additional experiments using non-informative cues in the overlap task showed that the reaction times of correct anti-saccades and the error rate were decreased when cue and stimulus appeared at the same side. Analysis of the erratic pro-saccades revealed that almost all of them were corrected, i.e. they were followed by a second saccade towards the required location. It is found that the correction times were usually very short, with intersaccadic intervals between 0 and 150 ms. We suggest that the orienting mechanism, elicited by a transient peripheral cue, relates to the command and the decision to make a pro- rather than an anti-saccade. The cue elicits pro-orienting towards its position when a pro-saccade is required, and anti-orienting when an anti-saccade is required. The orienting effect is transient and decays with CLTs of more than 200 ms; this result holds for both anti-saccades and pro-saccades. Since subjects reported that they could not prevent the erratic pro-saccades or were often not aware of them, we conclude that this orienting mechanism occurs automatically, beyond voluntary control.     

Perception of visual space at the time of pro- and anti-saccades

The localization of peri-saccadically flashed objects shows two types of errors: first, a uniform shift in saccade direction, and second, a compression of visual space around the saccade target. Whereas the uniform shift occurs when the experiment is performed in complete darkness compression occurs when additional visual references are available. Thus peri-saccadic mislocalization contains motor and visual components. To distinguish between both factors we compared peri-saccadic localization errors during pro- and anti-saccades. In the case of anti-saccades, the visual cue that elicits the saccade and the actual eye movement are in opposite directions. We asked whether peri-saccadic compression can be observed with anti-saccades, and if so, whether the compression is directed toward the visual cue or follows the actual eye movement. In blocked trials, subjects performed saccades either toward a visual cue (pro-saccade) or to the mirrored position opposite to a visual cue (anti-saccade). Peri-saccadically, we flashed a thin vertical bar at one of four possible locations. Subjects had to indicate the perceived position of the bar with a mouse pointer about 500 ms after the saccade. Experiments were performed in complete darkness and with visual references. Peri-saccadic mislocalizations occurred during anti-saccades. The mislocalizations were very similar for pro- and anti-saccades in magnitude and direction. For both, pro- and anti-saccades, mislocalizations were directed toward the actual eye movement and not the visual cue.     

Age-related trends in saccade characteristics among the elderly

Eye movement recordings are useful for assessing neurological disorders, the prevalence of which increases with age. However, there is little rigorous quantitative data on describing oculomotor changes that occur during healthy aging. Here, we measured the ability of 81 normal elderly subjects (60-85 years) to perform two saccadic eye movement tasks: a pro-saccade task, requiring an automatic response to look towards a stimulus and an anti-saccade task, requiring inhibition of the automatic response to instead initiate a voluntary saccade away from the stimulus. Saccadic ability decreased with age: the oldest subjects were slower to initiate saccades and they made more direction errors (i.e., erroneous pro-saccades) in the anti-saccade task. Intra-subject variability in reaction time also correlated positively with age in both saccade tasks. Voluntary saccade control, as assessed by the anti-saccade task, was far more affected by aging than automatic control, as assessed by the pro-saccade task, suggesting that the mechanisms driving voluntary and automatic saccade performance deteriorate at different rates in the aging brain, and therefore likely involves different neural substrates. Our data provide insight into deficits due to normal brain changes in aging as well as a baseline to evaluate deficits caused by neurological disorders common in this age range.     

Saccadic impairments in Huntington's disease

Huntington’s disease (HD), a progressive neurological disorder involving degeneration in basal ganglia structures, leads to abnormal control of saccadic eye movements. We investigated whether saccadic impairments in HD (N = 9) correlated with clinical disease severity to determine the relationship between saccadic control and basal ganglia pathology. HD patients and age/sex-matched controls performed various eye movement tasks that required the execution or suppression of automatic or voluntary saccades. In the “immediate” saccade tasks, subjects were instructed to look either toward (pro-saccade) or away from (anti-saccade) a peripheral stimulus. In the “delayed” saccade tasks (pro-/anti-saccades; delayed memory-guided sequential saccades), subjects were instructed to wait for a central fixation point to disappear before initiating saccades towards or away from a peripheral stimulus that had appeared previously. In all tasks, mean saccadic reaction time was longer and more variable amongst the HD patients. On immediate anti-saccade trials, the occurrence of direction errors (pro-saccades initiated toward stimulus) was higher in the HD patients. In the delayed tasks, timing errors (eye movements made prior to the go signal) were also greater in the HD patients. The increased variability in saccadic reaction times and occurrence of errors (both timing and direction errors) were highly correlated with disease severity, as assessed with the Unified Huntington’s Disease Rating Scale, suggesting that saccadic impairments worsen as the disease progresses. Thus, performance on voluntary saccade paradigms provides a sensitive indicator of disease progression in HD. A. Peltsch and A. Hoffman contributed equally.     

Preparatory set associated with pro-saccades and anti-saccades in humans investigated with event-related FMRI

Previous studies have shown that the BOLD functional MRI (fMRI) signal is increased in several cortical areas when subjects perform anti-saccades compared with pro-saccades. It remains unknown, however, whether this increase is due to an increased cortical motor signal for anti-saccades or due to differences in preparatory set between pro- and anti-saccade trials. To address this question, we measured event-related fMRI in a paradigm that allowed us to separate instruction-related brain activity from saccade-related brain activity. In this paradigm, the instruction to either generate a pro-saccade or an anti-saccade was conveyed by a switch in the color of the central fixation stimulus and preceded the presentation of a peripheral stimulus by either 6, 10, or 14 s. Cortical areas were functionally mapped using the general linear model comparing standard pro- and anti-saccade blocks with fixation blocks. When the trials were aligned on the onset of the instruction stimulus, bilateral frontal eye fields and right hemisphere dorsolateral prefrontal cortex showed an increased signal during the instruction period on anti-saccade trials as compared with pro-saccade trials. When the trials were aligned on the movement stimulus and the instruction period activity was subtracted, there were no differences between pro- and anti-saccades. This finding suggests that the increased cortical activation found in previous blocked designs originates predominately from differences in preparatory set and not from differences in the motor signal between pro- and anti-saccades.     

Altered control of visual fixation and saccadic eye movements in attention-deficit hyperactivity disorder

Attention-deficit hyperactivity disorder (ADHD) is characterized by the overt symptoms of impulsiveness, hyperactivity, and inattention. A frontostriatal pathophysiology has been hypothesized to produce these symptoms and lead to reduced ability to inhibit unnecessary or inappropriate behavioral responses. Oculomotor tasks can be designed to probe the ability of subjects to generate or inhibit reflexive and voluntary responses. Because regions of the frontal cortex and basal ganglia have been identified in the control of voluntary responses and saccadic suppression, we hypothesized that children and adults diagnosed with ADHD may have specific difficulties in oculomotor tasks requiring the suppression of reflexive or unwanted saccadic eye movements. To test this hypothesis, we measured eye movement performance in pro- and anti-saccade tasks of 114 ADHD and 180 control participants ranging in age from 6 to 59 yr. In the pro-saccade task, participants were instructed to look from a central fixation point toward an eccentric visual target. In the anti-saccade task, stimulus presentation was identical, but participants were instructed to suppress the saccade to the stimulus and instead look from the central fixation point to the side opposite the target. The state of fixation was manipulated by presenting the target either when the central fixation point was illuminated (overlap condition) or at some time after it disappeared (gap condition). In the pro-saccade task, ADHD participants had longer reaction times, greater intra-subject variance, and their saccades had reduced peak velocities and increased durations. In the anti-saccade task, ADHD participants had greater difficulty suppressing reflexive pro-saccades toward the eccentric target, increased reaction times for correct anti-saccades, and greater intra-subject variance. In a third task requiring prolonged fixation, ADHD participants generated more intrusive saccades during periods when they were required to maintain steady fixation. The results suggest that ADHD participants have reduced ability to suppress unwanted saccades and control their fixation behavior voluntarily, a finding that is consistent with a fronto-striatal pathophysiology. The findings are discussed in the context of recent neurophysiological data from nonhuman primates that have identified important control signals for saccade suppression that emanate from frontostriatal circuits.     

An early transient 40 Hz activity discriminates a following pro-saccade from a no-move and anti-saccade choice

We studied the oscillatory activity of the scalp-recorded EEG in healthy humans performing a task that required a particular eye-movement response choice according to the shape of a visual target. We observed a significant stimulus-aligned activity at the 40 Hz frequency band 100 ms after the appearance of the target only when that target was the end point for the subsequent eye movement (pro-saccade). This activity was most prominent over the central-parietal area of the right hemisphere. When the target indicated a movement to the opposite direction (anti-saccade) or indicated that no movement was required (no-move), this 40 Hz activity was nearly absent. This difference in activity between the pro-saccade and the other two tasks was evident in the single subject ERPs for four of the six subjects studied. In contrast, the movement-aligned 40 Hz activity for the pro-saccade and anti-saccade was almost identical. We speculate that this early stimulus-aligned 40 Hz activity might reflect a fast transformation of a visual stimulus to a motor response (eye movement) that can be performed for the pro-saccade task where stimulus-response compatibility is strong compared to the anti-saccade and no-move tasks. The movement-aligned 40 Hz activity might be related to the motor response preparation per se. We conclude that this task specific transient oscillatory activity could be used as a probe in the study of the temporal dynamics of visuomotor transformations.     

Age-related performance of human subjects on saccadic eye movement tasks

We measured saccadic eye movements in 168 normal human subjects, ranging in age from 5 to 79 years, to determine age-related changes in saccadic task performance. Subjects were instructed to look either toward (pro-saccade task) or away from (anti-saccade task) an eccentric target under different conditions of fixation. We quantified the percentage of direction errors, the time to onset of the eye movement (saccadic reaction time: SRT), and the metrics and dynamics of the movement itself (amplitude, peak velocity, duration) for subjects in different age groups. Young children (5–8 years of age) had slow SRTs, great intra-subject variance in SRT, and the most direction errors in the anti-saccade task. Young adults (20–30 years of age) typically had the fastest SRTs and lowest intra-subject variance in SRT. Elderly subjects (60–79 years of age) had slower SRTs and longer duration saccades than other subject groups. These results demonstrate very strong age-related effects in subject performance, which may reflect different stages of normal development and degeneration in the nervous system. We attribute the dramatic improvement in performance in the anti-saccade task that occurs between the ages of 5–15 years to delayed maturation of the frontal lobes. Received: 27 October 1997 / Accepted: 27 February 1998     

Express saccades in cat: effects of task and target modality

Saccadic eye movements to visual, auditory, and bimodal targets were measured in four adult cats. Bimodal targets were visual and auditory stimuli presented simultaneously at the same location. Three behavioral tasks were used: a fixation task and two saccadic tracking tasks (gap and overlap task). In the fixation task, a sensory stimulus was presented at a randomly selected location, and the saccade to fixate that stimulus was measured. In the gap and overlap tasks, a second target (hereafter called the saccade target) was presented after the cat had fixated the first target. In the gap task, the fixation target was switched off before the saccade target was turned on; in the overlap task, the saccade target was presented before the fixation target was switched off. All tasks required the cats to redirect their gaze toward the target (within a specified degree of accuracy) within 500 ms of target onset, and in all tasks target positions were varied randomly over five possible locations along the horizontal meridian within the cat's oculomotor range. In the gap task, a significantly greater proportion of saccadic reaction times (SRTs) were less than 125 ms, and mean SRTs were significantly shorter than in the fixation task. With visual targets, saccade latencies were significantly shorter in the gap task than in the overlap task, while, with bimodal targets, saccade latencies were similar in the gap and overlap tasks. On the fixation task, SRTs to auditory targets were longer than those to either visual or bimodal targets, but on the gap task, SRTs to auditory targets were shorter than those to visual or bimodal targets. Thus, SRTs reflected an interaction between target modality and task. Because target locations were unpredictable, these results demonstrate that cats, as well as primates, can produce very short latency goal-directed saccades.     

The influence of stimulus direction and eccentricity on pro- and anti-saccades in humans

We examined the sensory and motor influences of stimulus eccentricity and direction on saccadic reaction times (SRTs), direction-of-movement errors, and saccade amplitude for stimulus-driven (prosaccade) and volitional (antisaccade) oculomotor responses in humans. Stimuli were presented at five eccentricities, ranging from 0.5° to 8°, and in eight radial directions around a central fixation point. At 0.5° eccentricity, participants showed delayed SRT and increased direction-of-movement errors consistent with misidentification of the target and fixation points. For the remaining eccentricities, horizontal saccades had shorter mean SRT than vertical saccades. Stimuli in the upper visual field trigger overt shifts in gaze more easily and faster than in the lower visual field: prosaccades to the upper hemifield had shorter SRT than to the lower hemifield, and more anti-saccade direction-of-movement errors were made into the upper hemifield. With the exception of the 0.5° stimuli, SRT was independent of eccentricity. Saccade amplitude was dependent on target eccentricity for prosaccades, but not for antisaccades within the range we tested. Performance matched behavioral measures described previously for monkeys performing the same tasks, confirming that the monkey is a good model for the human oculomotor function. We conclude that an upper hemifield bias lead to a decrease in SRT and an increase in direction errors.     

Event-related potentials and saccadic reaction times: effects of fixation point offset or change

Previous studies have shown that saccadic reaction times (SRTs) are reduced if the initial fixation point (FP) disappears 200 ms (gap period) before a peripheral target is presented. This gap saccade task is associated with a negative cortical potential at the end of the gap period. To determine whether the neural processes underlying this potential account for the reduction of SRTs during gap saccade tasks, we recorded event-related potentials (ERPs) in 19 subjects performing a gap saccade task (gap duration 200 ms), a warning saccade task (the color of the FP changed 200 ms prior to target appearance) and an overlap task (the FP remained visible during the trial). SRTs were shortest during the gap task, longest during the overlap task and intermediate during the warning task. The gap and warning tasks were accompanied by the same widespread negative cortical potential with a maximum at the time of stimulus presentation. These findings indicate that the warning effect mediated by the disappearance of the FP during gap saccade tasks is responsible for the gap negativity which was observed by several authors. Our findings of shorter SRTs during the gap task than the warning task, however, suggest that the gap has an additional effect that probably depends on subcortical mechanisms. Received: 01 June 1998 / Accepted: 12 March 1999     

Visuomotor mental rotation of saccade direction

This investigation studied the latencies of saccadic eye movements that were directed away from a target by a variable angular distance, which was given by instruction. Such a movement presumably requires an intentional, visuomotor mental rotation of the saccade vector, resulting in prolonged reaction times. From a study on the control of directed hand movements, it has been hypothesized that all visuomotor and visual mental rotation tasks share a common processing stage. We tested this hypothesis with a saccade task in which subjects shifted their gaze either towards (0°, pro-saccade), or 30, 60, 90, 120, 150, or 180° (anti-saccade) away from a randomly cued position on an imaginary clock face. With four different cueing conditions, latencies increased monotonically with required gaze shift from 0–150°, thus exhibiting a mental rotation latency pattern. However, we also found anti-saccades faster than 150° gaze shift and slower rotation speeds with peripheral cues than with central cues. Together with the overall shallower latency increase compared with previous findings with mental rotation tasks, these results cast doubt on the notion of a common, central processing mechanism for the different types of tasks. Received: 11 August 1998 / Accepted: 9 February 1999     

Developmental fractionation and differential discrimination of the anti-saccadic direction error

This partial re-analysis of the data of a total of 433 experimental sessions transfers the distinction of "express" and "regular" latency ranges from the pro-saccade task to the latencies of the anti-saccadic direction errors. Express errors and express saccades (ES) loaded on one and the same PCA factor, and both variables were subject to only minor developmental changes from childhood to young adulthood. Regular direction errors loaded, with opposite signs, on the same factor as pro-saccadic reaction times and were unrelated to express errors; these direction errors showed substantial developmental changes. Our data contribute with new evidence from the anti-saccade task to the long standing debate on whether ES should be considered as a separate type of saccade. Anti-saccadic direction errors with express latencies can be distinguished from those with regular latencies. Future anti-saccade task research should therefore analyse these error types separately in order to look for further evidence in favour of their conceptual and statistical distinction.     

Effects of ethanol on anti-saccade task performance

It has been shown that saccade-related neurons in the superior colliculus (SC) display an increased level of prestimulus activity and a higher stimulus-related burst in action potentials preceding direction errors in the anti-saccade task compared with correct anti-saccades. From this, it has been hypothesized that errors occur when the incoming visual signal in the SC passes a threshold and triggers a reflexive saccade. This hypothesis predicts that an attenuated visual signal will reduce the number of direction errors. Since ethanol has been shown to have a suppressive effect on cortical visual event-related potentials (ERPs), the purpose of the present study was to investigate the effects of moderate ethanol consumption on anti-saccade performance. Under both placebo and ethanol conditions, we recorded ERPs and measured eye movements in male subjects during the performance of an anti-saccade task in which the fixation point disappeared 200 ms prior to stimulus presentation. Compared with the placebo condition, we found in the ethanol condition: (1) a decrease in ERP amplitudes during the gap period and after stimulus presentation, (2) an increase in the latencies of anti-saccades, and (3) a decrease in the percentage of direction errors. These data demonstrate the effects of ethanol on anti-saccade task performance and provide further support for the hypothesis that errors in the anti-saccade task are triggered by the incoming visual signal. Electronic Publication     

Remapping the remembered target location for anti-saccades in human posterior parietal cortex

We used functional magnetic resonance imaging (fMRI) to investigate the role of the human posterior parietal cortex (PPC) in anti-saccades. To do so, we exploited the laterality of a subregion of the PPC for remembered target location. Using an event-related design, we tracked fMRI signal changes in this region while subjects remembered the location of a flashed target, then were instructed to plan either an anti- or pro-saccade to that location, and finally were instructed to execute the movement. At first, the region responded preferentially to the memory of a target location presented in the contralateral visual field. However, when an anti-cue specified a saccadic response into the opposite visual field, we observed a dynamic shift in cortical activity from one hemisphere to the other. This shows that this region within the human posterior parietal cortex codes the target location for an upcoming saccade, rather than the location of the remembered visual stimulus in an anti-saccade task.     

Dynamic dissociation of visual selection from saccade programming in frontal eye field.

Previous studies of visually responsive neurons in the frontal eye fields have identified a selection process preceding saccades during visual search. The goal of this experiment was to determine whether the selection process corresponds to the selection of a conspicuous stimulus or to preparation of the next saccade. This was accomplished with the use of a novel task, called search-step, in which the target of a singleton visual search array switches location with a distracter on random trials. The target step trials created a condition in which the same stimulus yielded saccades either toward or away from the target. Visually responsive neurons in frontal eye field selected the current location of the conspicuous target even when gaze shifted to the location of a distractor. This dissociation demonstrates that the selection process manifest in visual neurons in the frontal eye field may be an explicit interpretation of the image and not an obligatory saccade command.     

Control of eye movement reflexes

We investigated the effect of strategic suppression of reflexive eye movements on external control over fixation using a fixation offset paradigm. A visual signal at fixation facilitates the fixation reflex and inhibits eye movements. Certain preparatory states render the fixation reflex less reactive to visual stimulation at fixation, as evidenced by a reduction in the fixation offset effect (FOE). For example, past studies have suggested that the reduced FOE during anti-saccade tasks results from the requirement to inhibit reflexive eye movements. We tested whether suppressing reflexive saccades reduces external control over ocular fixation using a go-nogo saccade paradigm. During each trial, one of two targets appeared in the periphery. Participants were instructed to saccade to one target (go), but when the other target appeared they either had to maintain fixation (nogo) or move their eyes in the direction opposite the target (anti). When nogo trials were admixed with target-directed saccades a large FOE was observed compared to when target-directed saccades occurred alone (experiment 1); however, when anti-saccades were mixed with target-directed saccades, a small FOE was observed for both types of eye movements (experiment 2). We conclude that suppressing reflexive eye movements does not reduce external control over fixation. Further research is necessary to elucidate which other component of preparing to make an anti-saccade diminishes the FOE.