Smooth pursuit in schizophrenia: Abnormalities of open- and closed-loop responses

A sample of 29 schizophrenia patients and 27 nonpsychiatric subjects were tested on measures of open- and closed-loop smooth-pursuit performance. Rashbass step-ramps were used to measure pursuit latency and open-loop gain. Regular ramps were used to calculate frequency and amplitude of both catch-up saccades and square-wave jerks, frequency of anticipatory saccades, and steady-state gain. Schizophrenia patients demonstrated lower open-loop gain than did nonpsychiatric subjects, an effect that was accentuated at faster target velocities. They also showed reduced steady-state gain, but only to 30%s right-moving targets. There was no evidence of saccadic abnormalities during smooth pursuit among the schizophrenia patients. These patients generated fewer square-wave jerks than did nonpsychiatric subjects for 10% left-moving targets. These results suggest an abnormality of smooth-pursuit initiation among patients with schizophrenia.     

Quantitative analysis of catch-up saccades during sustained pursuit

During visual tracking of a moving stimulus, primates orient their visual axis by combining two very different types of eye movements, smooth pursuit and saccades. The purpose of this paper was to investigate quantitatively the catch-up saccades occurring during sustained pursuit. We used a ramp-step-ramp paradigm to evoke catch-up saccades during sustained pursuit. In general, catch-up saccades followed the unexpected steps in position and velocity of the target. We observed catch-up saccades in the same direction as the smooth eye movement (forward saccades) as well as in the opposite direction (reverse saccades). We made a comparison of the main sequences of forward saccades, reverse saccades, and control saccades made to stationary targets. They were all three significantly different from each other and were fully compatible with the hypothesis that the smooth pursuit component is added to the saccadic component during catch-up saccades. A multiple linear regression analysis was performed on the saccadic component to find the parameters determining the amplitude of catch-up saccades. We found that both position error and retinal slip are taken into account in catch-up saccade programming to predict the future trajectory of the moving target. We also demonstrated that the saccadic system needs a minimum period of approximately 90 ms for taking into account changes in target trajectory. Finally, we reported a saturation (above 15 degrees /s) in the contribution of retinal slip to the amplitude of catch-up saccades.     

Eye movement disorders after frontal eye field lesions in humans

Eye movements were recorded electro-oculographically in three patients with a small ischemic lesion affecting the left frontal eye field (FEF) and in 12 control subjects. Reflexive visually guided saccades (gap and overlap tasks), antisaccades, predictive saccades, memory-guided saccades, smooth pursuit and optokinetic nystagmus (OKN) were studied in the three patients. Staircase saccades and double step saccades were also studied in one of the three patients. For both leftward and rightward saccades, latency in the overlap task (but not in the gap task) and that of correct antisaccades and of memory-guided saccades was significantly increased, compared with the results of controls. There was a significant decrease in the amplitude gain of all rightward saccades programmed using retinotopic coordinates (gap and overlap tasks, predictive and memory-guided saccades), whereas the amplitude gain of corresponding leftward saccades was preserved. Such an asymmetry between leftward and rightward saccades was significant. In the staircase paradigm as well as for the first saccade in the double step paradigm (with the use of retinotopic coordinates in both cases), the amplitude gain of rightward saccades was also significantly lower than that of leftward saccades. Moreover, in the double step paradigm, the amplitude gain of the first rightward saccade was significantly lower than that of the second rightward saccade (programmed using extraretinal signals), which was preserved. The percentage of errors in the antisaccade task did not differ significantly from that of normal subjects. In the predictive saccade paradigm, the percentage of predictive rightward saccades was significantly decreased. The left smooth pursuit gain for all tested velocities, the right smooth pursuit gain for higher velocities, and the left OKN gain were significantly decreased. The results show, for the first time in humans, that the FEF plays an important role in (1) the disengagement from central fixation, (2) the control of contralateral saccades programmed using retinotopic coordinates, (3) saccade prediction and (4) the control of smooth pursuit and OKN, mainly ipsilaterally. In contrast, the left FEF did not appear to be crucial for the control of the only type of saccades programmed using extraretinal signals studied here.     

Vestibular catch-up saccades in labyrinthine deficiency

During rapid head rotations, saccades ipsiversive with compensatory vestibulo-ocular reflex (VOR) slow phases may augment the deficient VOR and assist gaze stabilization in space. The present experiments compared these vestibular catch-up saccades (VCUSs) with visually and memory-guided saccades. To characterize VCUSs and their relationship to deficiency of the initial VOR, we delivered random, whole-body transients of 1000 and 2800 degrees/s2 peak yaw acceleration around four different eccentric vertical axes in eight unilaterally and one bilaterally vestibulopathic subjects, as well as nine age-matched normal subjects. Eye and head movements were sampled at 1200 Hz using magnetic search coils. Subjects fixed targets at either 500 or 15 cm distance immediately before unpredictable onset of rotation in darkness. Under all testing conditions, normal subjects exhibited only compensatory vestibular slow phases and occasional anticompensatory quick phases. This behavior was also typical of unilaterally vestibulopathic subjects rotated contralesionally. When rotated ipsilesionally, however, vestibulopathic subjects had deficient slow-phase VOR gain with prolonged latency, and six of the nine exhibited saccadic movements in the compensatory direction (VCUSs). Higher head accelerations preferentially evoked VCUSs, but there were no preferred combinations of target distances and eccentric rotation axes. Peak velocities and durations of VCUSs increased with saccade amplitude. The latency distribution for VCUSs peaked around 70 ms, substantially shorter than reported for either visually guided express saccades or vestibular memory contingent saccades. The latency of each VCUS was highly correlated with the gaze error prior to that VCUS. The amplitude of VCUSs was calibrated to gaze position error, such that VCUSs reduced gaze error by an average of 37%. Thus when VOR slow-phase responses cannot compensate fully for head rotation, vestibular gaze position error can nevertheless calibrate the programming of VCUSs to augment the deficient VOR, much like catch-up saccades substitute for deficient visual pursuit.     

Event-related potentials before saccades and antisaccades and their relation to reaction time

In the present study, reaction time (RT) was measured in 12 healthy subjects in a saccade and antisaccade task while recording electroencephalographic activity (EEG) from 62 electrodes on the scalp. Event-related potentials averaged both on target appearance and on saccade onset were larger in amplitude (increased negativity) for the antisaccade task compared to the saccade task. The relation of RT variability to EEG amplitude was studied by averaging stimulus-aligned and movement-aligned individual trials for each subject into four RT quartile groups. The analysis showed a relation of EEG amplitude to RT for both saccades and antisaccades. More specifically, the ERP negativity at 100–120 ms after stimulus onset in the saccade task and at 160–200 ms after stimulus onset in the antisaccade task for stimulus-aligned ERPs decreased monotonically with increasing RT as would be expected if this signal would be related to the eye movement preparation processes. This was much more pronounced and wide spread for the antisaccades than for visually triggered saccades. The peak negativity before movement onset for movement-aligned ERPs also covaried with RT suggesting no relation of this activity to movement preparation processes. This study then confirmed that only a particular ERP signal variation was related to the saccadic eye movement preparatory processes while other components of the ERP have no specific relation to the movement preparation. This particular signal was more prominent for antisaccades compared to visually triggered saccades supporting previous evidence for the cortical involvement in the preparation of these voluntary eye movements. In conclusion, this study validates the use of ERPs in the study of the planning and execution of saccadic eye movements.     

Amplitude transition function of human express saccades

The gap paradigm often promotes the occurrence of express saccades, which are supposed to be short latency, visually guided saccades, often forming a separate peak in saccadic latency distribution. We designed six experiments in which we compared the amplitudes of anticipatory, express and regular saccades, for various conditions of target eccentricities, target direction, and predictability. Then, saccadic amplitude was expressed as a continuous function of latency, for the various target eccentricities. From the obtained results, it is proposed that a saccade of a given amplitude is prepared during the gap period, on the basis of internal cues. The latency range of express saccades is a transition zone when the target begins to influence the already prepared saccade. The resulting amplitude will be a weighted average of the value determined during the gap and of the value defined by the target, the weighting being determined by the latency of the saccade. If the preprogrammed saccade is wrongly directed, the target will not be able to correct the saccadic amplitude and the express saccade will have the same amplitude as anticipatory saccades. Regular saccades are delayed sufficiently so that a wrongly directed preprogrammed saccade can be canceled or the amplitude of a rightly directed saccade can be adjusted according to the exact position of the visual target.     

The mirror antisaccade task: direction–amplitude interaction and spatial accuracy characteristics

In this study we examined the performance of human subjects in three oculomotor tasks: a visually guided saccade task (VST), a simple antisaccade task (SAT) and a mirror antisaccade task (MAT). The stimulus presentation was identical for all three tasks, and the differentiation of the tasks was based on the instruction given to the subjects. Subjects were instructed to either look at a visually presented target location (visually triggered saccade task), or to look at the opposite direction of the visually presented target (simple antisaccade task) or finally to look at the mirror location opposite to the location of the visually presented target (mirror antisaccade task). The loading of the simple antisaccade task with the addition of the amplitude requirement did not affect the percentage of directional errors but slowed down the onset of antisaccade execution by 19 ms. The amplitude of the directionally correct antisaccade in the mirror antisaccade task showed a significant distortion of the true mirror target location. This dysmetria followed the same qualitative pattern to that observed for the visually guided saccades, i.e., a near-target hypermetria together with a far-target hypometria, but both these features were exaggerated in the mirror antisaccade task. This distorted amplitude modulation of mirror antisaccade amplitude was completely lost in corrected antisaccades that followed a directional error.     

Accuracies of saccades to moving targets during pursuit initiation and maintenance

The overall goals of the studies presented here were to compare (1) the accuracies of saccades to moving targets with either a novel or a known target motion, and (2) the relationships between the measures of target motion and saccadic amplitude during pursuit initiation and maintenance. Since resampling of position error just prior to saccade initiation can confound the interpretation of results, the target ramp was masked during the planning and execution of the saccade. The results suggest that saccades to moving targets were significantly more accurate if the target motion was known from the early part of the trial (e.g., during pursuit maintenance) than in the case of novel target motion (e.g., during pursuit initiation); both these types of saccades were more accuate than those when target motion information was not available. Using target velocity in space as a rough estimate of the magnitude of the extra-retinal signal during pursuit maintenance, the saccadic amplitude was significantly associated with the extra-retinal target motion information after accounting for the position error. In most subjects, this association was stronger than the one between retinal slip velocity and saccadic amplitude during pursuit initiation. The results were similar even when the smooth eye motion prior to the saccade was controlled. These results suggest that different sources of target motion information (retinal image velocity vs internal representation of previous target motion in space) are used in planning saccades during different stages of pursuit. The association between retinal slip velocity and saccadic amplitude is weak during initiation, thus explaining poor saccadic accuracy during this stage of pursuit.     

Saccadic disinhibition in schizophrenia patients and their first-degree biological relatives

Several studies have reported that patients with schizophrenia and their relatives perform poorly on antisaccade tasks and have suggested that this deficit represents saccadic disinhibition. If this proposition is correct, then varying task parameters that specifically increase the difficulty with which unwanted saccades can be inhibited should exacerbate deficits. Forty-two schizophrenia patients, 42 of their first-degree biological relatives, 21 psychotic affective disorder patients, and 38 nonpsychiatric comparison subjects were given fixation and antisaccade tasks. The introduction of distracters and the presence of visible fixation stimuli were parameters used to vary the difficulty in suppressing unwanted saccades (inhibitory load). It is known that the presence of a fixation stimulus at the time when a saccade must be inhibited results in fewer reflexive errors on antisaccade tasks. Performance on fixation tasks without (low load) vs with distracters (high load) and antisaccade tasks that had fixation stimuli still visible (low load) vs already extinguished (high load) at the time when the reflexive saccade must be inhibited was compared. The schizophrenia patients and their first-degree biological relatives showed evidence of increased saccadic disinhibition that was most pronounced during high inhibitory load conditions. These data indicate that dysfunctional inhibitory processes, at least in the oculomotor domain, are associated with the liability to schizophrenia. Results also suggest that this genetic liability may be related to dysfunctional prefrontal cortical areas that provide top-down inhibitory control over reflexive saccade generation.     

Deficits of cortical oculomotor mechanisms in cerebellar atrophy patients

Commonly, the cerebellum is not associated with cortical components of saccadic eye movement programming. The present study investigates cerebellar effects on visually guided saccades in reflexive tasks (step, gap, overlap) and on internally driven saccades in intentional tasks (anti, memory, short memory sequences of four targets) in five patients with isolated cerebellar atrophy. The cerebellar dysfunction led to impairments in both reflexive and intentional saccades. Cerebellar atrophy patients showed an increase in the gain variability and an increase in the saccade latency. Furthermore, in the memory and anti task, suppression and pro-saccade errors were more frequent in the atrophy group compared to the control group. In the sequence task, patients had difficulties reproducing all four target locations in the order of the displayed sequence. The high variability of the saccade gain is a common observation in cerebellar atrophy patients and can be explained by the general variability present in the saccadic system. The increase in the saccade latency could be due to a cerebellar contribution to cortical processes related to fixation and target selection preceding the initiation of a saccade. Furthermore, the frequent occurrence of saccade errors in the memory and anti task suggests a cerebellar involvement in frontal inhibition of unwanted reflexive saccades. The impaired reproduction of saccade sequences in atrophy patients points to a deficit in short-term memory processes. Thus, this study provides further evidence that the cerebellum is involved in different cortical mechanisms related to the control of saccadic eye movements.     

Saccadic and smooth pursuit eye movements: computational modeling of a common inhibitory mechanism in brainstem

The oculomotor system coordinates different types of eye movements in order to orient the visual axis, including saccade and smooth pursuit,. It was traditionally thought that the premotor pathways for these different eye movements are largely separate. In particular, a group of midline cells in the pons called omnipause neurons were considered to be part of only the saccadic system. Recent experimental findings have shown activity modulation of these brainstem premotor neurons during both kinds of eye movements. In this study, we propose a new computational model of the brainstem circuitry underlying the generation of saccades and smooth pursuit eye movements. Similar models have been developed earlier, but mainly looking at pure saccades. Here, we integrated recent neurophysiological findings on omnipause neuron activity during smooth pursuit. Our computational model can mimic some new experimental findings as the similarity of "eye velocity profile" with "omnipause neuron pattern of activity" in pursuit movement. We showed that pursuit neuron activity is augmented during catch-up saccades; this increment depends on the initial pursuit velocity in catch-up saccade onset. We conclude that saccadic and pursuit components of catch-up saccades are added to each other nonlinearly.     

Interaction between smooth anticipation and saccades during ocular orientation in darkness

A saccade triggered during sustained smooth pursuit is programmed using retinal information about the relative position and velocity of the target with respect to the eye. Thus the smooth pursuit and saccadic systems are coordinated by using common retinal inputs. Yet, in the absence of retinal information about the relative motion of the eye with respect to the target, the question arises whether the smooth and saccadic systems are still able to be coordinated possibly by using extraretinal information to account for the saccadic and smooth eye movements. To address this question, we flashed a target during smooth anticipatory eye movements in darkness, and the subjects were asked to orient their visual axis to the remembered location of the flash. We observed multiple orientation saccades (typically 2-3) toward the memorized location of the flash. The first orienting saccade was programmed using only the position error at the moment of the flash, and the smooth eye movement was ignored. However, subsequent saccades executed in darkness compensated gradually for the smooth eye displacement (mean compensation congruent with 70%). This behavior revealed a 400-ms delay in the time course of orientation for the compensation of the ongoing smooth eye displacement. We conclude that extraretinal information about the smooth motor command is available to the saccadic system in the absence of visual input. There is a 400-ms delay for smooth movement integration, saccade programming and execution.     

Reaction times of vertical prosaccades and antisaccades in gap and overlap tasks

Horizontal saccadic reaction times (SRTs) have been extensively studied over the past 3 decades, concentrating on such topics as the gap effect, express saccades, training effects, and the role of fixation and attention. This study investigates some of these topics with regard to vertical saccades. The reaction times of vertical saccades of 13 subjects were measured using the gap and the overlap paradigms in the prosaccade task (saccade to the stimulus) and the antisaccade task (saccade in the direction opposite to the stimulus). In the gap paradigm, the initial fixation point (FP) was extinguished 200 ms before stimulus onset, while, in the overlap paradigm, the FP remained on during stimulus presentation. With the prosaccade overlap task, it was found that most subjects (10/13) — whether they were previously trained making horizontal saccades or naive — had significantly faster upward saccades compared with their downward saccades. One subject was faster in the downward direction and two were symmetrical. The introduction of the gap reduced the reaction times of the prosaccades, and express saccades were obtained in some naive and most trained subjects. This gap effect was larger for saccades made to the downward target. The strength of the updown asymmetry was more pronounced in the overlap as compared to the gap paradigm. With the antisaccade task, up-down asymmetries were much reduced. Express antisaccades were absent even with the gap paradigm, but reaction times were reduced as compared to the antisaccade overlap paradigm. There was a slight tendency for a larger gap effect of downward saccades. All subjects produced a certain number of erratic prosaccades in the antitaks, more with the gap than with the overlap paradigm. There was a significantly larger gap effect for the erratic prosaccades made to the downward, as compared to the upward, target, due to increased downward SRTs in the overlap paradigm. Three subjects trained in both the horizontal and the vertical direction showed faster SRTs and more express saccades in the horizontal directions as compared to the vertical. It is concluded that different parts of the visual field are differently organized with both directional and nondirectional components in saccade preparation.     

Adaptation of catch-up saccades during the initiation of smooth pursuit eye movements

Reduction of retinal speed and alignment of the line of sight are believed to be the respective primary functions of smooth pursuit and saccadic eye movements. As the eye muscles strength can change in the short-term, continuous adjustments of motor signals are required to achieve constant accuracy. While adaptation of saccade amplitude to systematic position errors has been extensively studied, we know less about the adaptive response to position errors during smooth pursuit initiation, when target motion has to be taken into account to program saccades, and when position errors at the saccade endpoint could also be corrected by increasing pursuit velocity. To study short-term adaptation (250 adaptation trials) of tracking eye movements, we introduced a position error during the first catch-up saccade made during the initiation of smooth pursuit—in a ramp-step-ramp paradigm. The target position was either shifted in the direction of the horizontally moving target (forward step), against it (backward step) or orthogonally to it (vertical step). Results indicate adaptation of catch-up saccade amplitude to back and forward steps. With vertical steps, saccades became oblique, by an inflexion of the early or late saccade trajectory. With a similar time course, post-saccadic pursuit velocity was increased in the step direction, adding further evidence that under some conditions pursuit and saccades can act synergistically to reduce position errors.     

Saccadic instabilities and voluntary saccadic behaviour

Primary gaze fixation is never perfectly stable but can be interrupted by involuntary, conjugate saccadic intrusions (SI). SI have a high prevalence in the normal population and are characterised by a horizontal fast eye movement away from the desired eye position, followed, after a variable duration, by a return saccade or drift. Amplitudes are usually below 1° and they often exhibit a directional bias. The aim of the present study was to investigate the aetiology of SI in relation to saccadic behaviour. It was hypothesised that if SI resulted from deficits in the saccadic system (i.e. reduced inhibitory mechanisms), changes in voluntary saccade behaviour may be apparent and related to SI frequency. To examine this, synchrony (no gap), gap, overlap and antisaccade tasks were conducted on ten normal subjects. No significant correlations were found between SI frequency and voluntary saccade latencies, the percentage of express saccades, or the percentage of antisaccade errors. In addition, no significant correlations were found between SI directional biases and saccade latency directional biases, express saccade biases or antisaccade error biases. These results suggest that an underlying alteration to saccadic behaviour is unlikely to be involved in SI production, and that the SI command signal may arise from the influence of attention on an intact saccadic system. Specifically, descending corticofugal signals relating to attention level and orientation may alter the balance between fixation and saccade generation, so determining SI characteristics.     

Role of retinal slip in the prediction of target motion during smooth and saccadic pursuit

Visual tracking of moving targets requires the combination of smooth pursuit eye movements with catch-up saccades. In primates, catch-up saccades usually take place only during pursuit initiation because pursuit gain is close to unity. This contrasts with the lower and more variable gain of smooth pursuit in cats, where smooth eye movements are intermingled with catch-up saccades during steady-state pursuit. In this paper, we studied in detail the role of retinal slip in the prediction of target motion during smooth and saccadic pursuit in the cat. We found that the typical pattern of pursuit in the cat was a combination of smooth eye movements with saccades. During smooth pursuit initiation, there was a correlation between peak eye acceleration and target velocity. During pursuit maintenance, eye velocity oscillated at approximately 3 Hz around a steady-state value. The average gain of smooth pursuit was approximately 0.5. Trained cats were able to continue pursuing in the absence of a visible target, suggesting a role of the prediction of future target motion in this species. The analysis of catch-up saccades showed that the smooth-pursuit motor command is added to the saccadic command during catch-up saccades and that both position error and retinal slip are taken into account in their programming. The influence of retinal slip on catch-up saccades showed that prediction about future target motion is used in the programming of catch-up saccades. Altogether, these results suggest that pursuit systems in primates and cats are qualitatively similar, with a lower average gain in the cat and that prediction affects both saccades and smooth eye movements during pursuit.     

Abnormalities of internally generated saccades in obsessive-compulsive disorder

BACKGROUND: We aimed to utilize tests of saccadic function to investigate whether cognitive abnormalities in obsessive-compulsive disorder (OCD) arise from a dysfunction of inhibitory processes or whether they reflect a more general difficulty in guiding behaviour on the basis of an internal representation of task goal. METHODS: Twelve patients with OCD and 12 matched controls performed a visually-guided saccade task, a volitional prosaccade task and an antisaccade task. The latency and gain of saccades was compared between groups for the three saccade tasks. The number of antisaccade errors was also calculated and compared between groups. RESULTS: There was no difference for antisaccade error rates between the groups. The latency of visually guided saccades did not differ between groups, however the latency of both volitional prosaccades and antisaccades was significantly slower in the patients with OCD than in controls. The difference in latency between volitional prosacades and antisaccades, however, was equal between groups. CONCLUSIONS: These results suggest that patients with OCD have an abnormality in guiding behaviour on the basis of an internal representation of the task goal, rather than a problem with inhibiting reflexive behaviour.     

Effects of age on latency and error generation in internally mediated saccades

Most studies on the effects of ageing on saccades have examined reflexive saccades; the only commonly studied volitional task has been the antisaccade task, with contradictory results. We examined in both young and elderly normal subjects the latency of anti-, memory-guided, and predictable saccades and the timing of self-paced saccades; we also evaluated errors made on the first two tasks. We expected errors to be correlated between tasks; we also expected antisaccade latencies and errors to be inversely correlated. We also expected antisaccade and memory-guided saccade latencies to be longer in individuals with a high self-paced rate. Except for predictable saccades, mean latencies were significantly higher in the elderly. However, their performance was more variable. Errors were also significantly more frequent on anti- and memory-guided saccade tasks. Most of the hypothesised correlations were not observed. Analysis of error latencies showed that whilst most antisaccade errors were reflexive, for memory-guided saccades both express errors and errors with latencies between 0.4 and 2.5 s were observed. The latter appeared to be a premature release of what would otherwise have been a properly planned response. Age thus impaired all but the predictable saccade task; nevertheless, there were few relationships between measures across tasks. This suggests that a range of processes mediate peoples' performance on these saccade paradigms.     

Switch performance in peripherally and centrally triggered saccades

A common hypothesis is that the switch cost measured when switching between prosaccades and antisaccades mainly reflects the inhibition of the saccadic system after the execution of an antisaccade, which requires the inhibition of a gaze response. The present study further tested this hypothesis by comparing switch performance between peripherally triggered saccades and centrally triggered saccades with the latter type of saccades not requiring inhibition of a gaze response. For peripherally triggered saccades, a switch cost was present for prosaccades but not for antisaccades. For centrally triggered saccades, a switch cost was present both for prosaccades and for antisaccades. The difference between both saccade tasks further supports the hypothesis that the switch performance observed for peripherally triggered saccades is related to the inhibition of a gaze response that is required when executing a peripherally triggered antisaccade and the persisting inhibition in the saccadic system this entails. Furthermore, the switch costs observed for centrally triggered saccades indicate that more general processes besides the persisting inhibition in the saccadic system, such as reconfiguration and interference control, also contribute to the switch performance in saccades.     

The Saccadic system more readily co-processes orthogonal than co-linear saccades

Real-life visual tasks such as tracking jumping objects and scanning visual scenes often require a sequence of saccadic eye movements. The ability of the ocular motor system to parallel process saccades has been previously demonstrated. We recorded the monocular eye movements of five normal human subjects using the magnetic search coil technique in a double step paradigm. Initial target jumps were always purely horizontal or purely vertical. We were interested in the latency to onset of the second saccade as a function of direction in relation to the first saccade. When the inter stimulus interval (ISI) was 150 or 180 ms orthogonal second saccades were of significantly shorter latency than second co-linear saccades. When the ISI was 250 ms the latencies of orthogonal and co-linear second saccades were statistically indistinguishable. Based on these findings it is postulated that the ocular motor system can more readily co-process orthogonal than co-linear saccades.