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     

Neural correlates of inter- and intra-individual saccadic reaction time differences in the gap/overlap paradigm

To examine the neural correlates of contextually differing control mechanisms in saccade initiation, we studied 18 subjects who performed two saccade paradigms in a pseudo-random order, while their eye movements were recorded in the MRI scanner (1.5 T). In the gap task the fixation point was extinguished 200 ms before target onset, and in the overlap task the fixation point vanished 500 ms after target onset. Subjects were asked to maintain stable fixation in the fixation period and to quickly saccade to peripherally presented targets. Inter-individual activation differences were assessed using regression analyses at the second level, with mean saccadic reaction time (SRT) of subjects as a covariate. To identify brain regions varying with trial-by-trial changes in SRTs, we included SRTs as a parametric modulation regressor in the general linear model. All analyses were regions of interest based and were performed separately for the gap and overlap conditions. For the gap paradigm, we did not obtain activation in regions previously shown to be involved in preparatory processes with much longer gap periods. Interestingly, both inter- and intra-individual variability analyses revealed a positive correlation of activation in frontal and parietal eye-movement regions with SRTs, indicating that slower saccade performance is possibly associated with higher cortical control. For the overlap paradigm, the trial-by-trial variability analysis revealed a positive correlation of activation in the right opercular inferior frontal gyrus with SRTs, possibly linked to fixation-related processes that have to be overcome to perform a speeded saccade in presence of a fixation point.     

Saccadic eye movements to visual and auditory targets

 Recent neurophysiological studies of the saccadic ocular motor system have lent support to the hypothesis that this system uses a motor error signal in retinotopic coordinates to direct saccades to both visual and auditory targets. With visual targets, the coordinates of the sensory and motor error signals will be identical unless the eyes move between the time of target presentation and the time of saccade onset. However, targets from other modalities must undergo different sensory-motor transformations to access the same motor error map. Because auditory targets are initially localized in head-centered coordinates, analyzing the metrics of saccades from different starting positions allows a determination of whether the coordinates of the motor signals are those of the sensory system. We studied six human subjects who made saccades to visual or auditory targets from a central fixation point or from one at 10° to the right or left of the midline of the head. Although the latencies of saccades to visual targets increased as stimulus eccentricity increased, the latencies of saccades to auditory targets decreased as stimulus eccentricity increased. The longest auditory latencies were for the smallest values of motor error (the difference between target position and fixation eye position) or desired saccade size, regardless of the position of the auditory target relative to the head or the amplitude of the executed saccade. Similarly, differences in initial eye position did not affect the accuracy of saccades of the same desired size. When saccadic error was plotted as a function of motor error, the curves obtained at the different fixation positions overlapped completely. Thus, saccadic programs in the central nervous system compensated for eye position regardless of the modality of the saccade target, supporting the hypothesis that the saccadic ocular motor system uses motor error signals to direct saccades to auditory targets. Received: 8 September 1995 / Accepted: 22 November 1996     

Characteristics of contralesional and ipsilesional saccades in hemianopic patients

In order to further our understanding of action-blindsight, four hemianopic patients suffering from visual field loss contralateral to a unilateral occipital lesion were compared to six healthy controls during a double task of verbally reported target detection and saccadic responses toward the target. Three oculomotor tasks were used: a fixation task (i.e., without saccade) and two saccade tasks (eliciting reflexive and voluntary saccades, using step and overlap 600 ms paradigms, respectively), in separate sessions. The visual target was briefly presented at two different eccentricities (5° and 8°), in the right or left visual hemifield. Blank trials were interleaved with target trials, and signal detection theory was applied. Despite their hemifield defect, hemianopic patients retained the ability to direct a saccade toward their contralesional hemifield, whereas verbal detection reports were at chance level. However, saccade parameters (latency and amplitude) were altered by the defect. Saccades to the contralesional hemifield exhibited longer latencies and shorter amplitudes compared to those of the healthy group, whereas only the latencies of reflexive saccades to the ipsilesional hemifield were altered. Furthermore, healthy participants showed the expected latency difference between reflexive and voluntary saccades, with the latter longer than the former. This difference was not found in three out of four patients in either hemifield. Our results show action-blindsight for saccades, but also show that unilateral occipital lesions have effects on saccade generation in both visual hemifields.     

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.     

Relationship between directed visual attention and saccadic reaction times

Summary Saslow (1967) and Fischer and Ramsperger (1984) found that saccadic reaction time (SRT) depends on the interval between the fixation point offset and the target onset. Using a continuously visible fixation point, we asked whether a similar function would be obtained if subjects attended to a peripherally viewed point extinguished at variable intervals before or after the target onset. The interval was varied between -500ms (i.e., attention stimulus offset after saccade target onset = overlap trials) and 500ms (i.e., attention stimulus offset before saccade target onset = gap trials). The results show a constant mean SRT of about 240 ms for overlap trials, and a U-shaped function with a minimum of 140 ms, at a gap duration of 200 ms, for gap trials. These findings suggest that saccadic latencies do not depend on the cessation of fixation per se, but rather on the disengagement of attention from any location in the visual field. The time required for subjects to disengage their attention is approximately 100 ms. This disengaged state of attention — during which short latency (express) saccades can be made — can be sustained only for a gap duration of 300 ms. At longer gap durations mean SRTs increase again.     

Influence of previous visual stimulus or saccade on saccadic reaction times in monkey.

Influence of previous visual stimulus or saccade on saccadic reaction times in monkey. Saccadic reaction times (SRTs) to suddenly appearing targets are influenced by neural processes that occur before and after target presentation. The majority of previous studies have focused on how posttarget factors, such as target attributes or changes in task complexity, affect SRTs. Studies of pretarget factors have focused on how prior knowledge of the timing or location of the impending target, gathered through cueing or probabilistic information, affects SRTs. Our goal was to investigate additional pretarget factors to determine whether SRTs can also be influenced by the history of saccadic and visual activity even when these factors are spatially unpredictive as to the location of impending saccadic targets. Monkeys were trained on two paradigms. In the saccade-saccade paradigm, monkeys were required to follow a saccadic target that stepped from a central location, to an eccentric location, back to center, and finally to a second eccentric location. The stimulus-saccade paradigm was similar, except the central fixation target remained illuminated during presentation of the first eccentric stimulus; the monkey was required to maintain central fixation and to make a saccade to the second eccentric stimulus only on disappearance of the fixation point. In both paradigms, the first eccentric stimulus was presented at the same, opposite, or orthogonal location with respect to the final target location in a given trial. We measured SRTs to the final target under conditions in which all parameters were identical except for the location of the first eccentric stimulus. In the saccade-saccade paradigm, we found that the SRT to the final target was slowest when it was presented opposite to the initial saccadic target, whereas in the stimulus-saccade paradigm the SRT to the final target was slowest when it was presented at the same location as the initial stimulus. In both paradigms, these increases in SRTs were greatest during the shortest intervals between presentation of successive eccentric stimuli, yet these effects remained present for the longest intervals employed in this study. SRTs became faster as the direction and eccentricity of the two successive stimuli became increasingly misaligned from that which produced the maximal SRT slowing in each paradigm. The results of the stimulus-saccade paradigm are similar to the phenomenon of inhibition of return (IOR) in which human subjects are slower to respond to stimuli that are presented at previously cued locations. We interpret these findings in terms of overlapping representations of visuospatial and oculomotor activity in the same neural structures.     

Fixation disengagement enhances peripheral perceptual processing: evidence for a perceptual gap effect

Temporal gaps between the offset of a central fixation stimulus and the onset of an eccentric target typically reduce saccade latencies (saccadic gap effect). Here, we test whether temporal gaps also affect perceptual performance in peripheral vision. In Experiment 1, subjects executed saccades to briefly presented peripheral target letters and reported letter identity afterwards. A central fixation stimulus either remained visible throughout the trial (overlap) or disappeared 200 ms before letter onset (gap). Experiment 2 tested perceptual performance without saccade execution, whereas Experiment 3 tested saccade execution without perceptual demands. Peripheral letter perception performance was enhanced in gap as compared to overlap conditions (perceptual gap effect) irrespective of concurrent oculomotor demands. Furthermore, the saccadic gap effect was modulated by concurrent perceptual demands. Experiment 4 ruled out a general warning explanation of the perceptual gap effect. These findings extend recent theories assuming a strong coupling between the preparation of goal-directed saccades and shifts of visual attention from the spatial to the temporal domain.     

Auditory-visual interactions subserving goal-directed saccades in a complex scene

This study addresses the integration of auditory and visual stimuli subserving the generation of saccades in a complex scene. Previous studies have shown that saccadic reaction times (SRTs) to combined auditory-visual stimuli are reduced when compared with SRTs to either stimulus alone. However, these results have been typically obtained with high-intensity stimuli distributed over a limited number of positions in the horizontal plane. It is less clear how auditory-visual interactions influence saccades under more complex but arguably more natural conditions, when low-intensity stimuli are embedded in complex backgrounds and distributed throughout two-dimensional (2-D) space. To study this problem, human subjects made saccades to visual-only (V-saccades), auditory-only (A-saccades), or spatially coincident auditory-visual (AV-saccades) targets. In each trial, the low-intensity target was embedded within a complex auditory-visual background, and subjects were allowed over 3 s to search for and foveate the target at 1 of 24 possible locations within the 2-D oculomotor range. We varied systematically the onset times of the targets and the intensity of the auditory target relative to background [i.e., the signal-to-noise (S/N) ratio] to examine their effects on both SRT and saccadic accuracy. Subjects were often able to localize the target within one or two saccades, but in about 15% of the trials they generated scanning patterns that consisted of many saccades. The present study reports only the SRT and accuracy of the first saccade in each trial. In all subjects, A-saccades had shorter SRTs than V-saccades, but were more inaccurate than V-saccades when generated to auditory targets presented at low S/N ratios. AV-saccades were at least as accurate as V-saccades but were generated at SRTs typical of A-saccades. The properties of AV-saccades depended systematically on both stimulus timing and S/N ratio of the auditory target. Compared with unimodal A- and V-saccades, the improvements in SRT and accuracy of AV-saccades were greatest when the visual target was synchronous with or leading the auditory target, and when the S/N ratio of the auditory target was lowest. Further, the improvements in saccade accuracy were greater in elevation than in azimuth. A control experiment demonstrated that a portion of the improvements in SRT could be attributable to a warning-cue mechanism, but that the improvements in saccade accuracy depended on the spatial register of the stimuli. These results agree well with earlier electrophysiological results obtained from the midbrain superior colliculus (SC) of anesthetized preparations, and we argue that they demonstrate multisensory integration of auditory and visual signals in a complex, quasi-natural environment. A conceptual model incorporating the SC is presented to explain the observed data.     

Visuomotor dependency on an initial fixation target involved in the disorder of visually-guided manual movement in Parkinson's disease

Role of a central fixation target on the latencies of visually guided manual movement was analyzed on young healthy subjects, age-matched control subjects and patients with Parkinson's disease (Hoehn and Yahr stages II, III, and IV). Two paradigms were used: overlap paradigm where a central fixation target was lighted throughout the test, and gap paradigm where a central fixation target was turned off 200 ms before a peripheral target was lighted. The subject was first asked to fixate the central target then instructed to locate a peripheral target with a laser beam spot, operated with wrist flexion or extension as quickly as possible. Latencies of gap paradigm are always shorter than those of overlap task in all the groups. Latencies of both overlap and gap tasks prolonged from young to elder, from elder to PD II, from PD II to PD III and from PD III to PD IV. Also latencies were extremely prolonged in the overlap tasks and correlated with disease severity. Latencies in the gap tasks were less prolonged as compared with those in the overlap tasks. The visual fixation target prolonged the visuo-motor latency in association with severity of Parkinson's disease.     

Observing human interaction with physical devices

Triggering of saccades depends on the task: in the gap task, fixation point switches off and target appears after a gap period; in the overlap task, target appears while fixation point is still on. Saccade latencies are shorter in the gap task, due to fixation disengagement and advanced movement preparation during the gap. The two modes of initiation are also hypothesized to be subtended by different cortical-subcortical circuits. This study tested whether interleaving the two tasks modifies latencies, due to switching between different modes of triggering. Two groups of healthy participants (21–29 vs. 39–55 years) made horizontal and vertical saccades in gap, overlap, and mixed tasks; saccades were recorded with the Eyelink. Both groups showed shorter latencies in the gap task, i.e. a robust gap effect and systematic differences between directions. For young adults, interleaving tasks made the latencies shorter or longer depending on direction, while for middle-age adults, latencies became longer for all directions. Our observations can be explained in the context of models such as that of Brown et al. (Neural Netw 17:471–510, 2004), which proposed that different combinations of frontal eye field (FEF) layers, interacting with cortico-subcortical areas, control saccade triggering in gap and overlap trials. Moreover, we suggest that in early adulthood, the FEF is functioning optimally; frequent changes of activity in the FEF can be beneficial, leading to shorter latencies, at least for some directions. However, for middle-age adults, frequent changes of activity of a less optimally functioning FEF can be time consuming. Studying the alternation of gap and overlap tasks provides a fine tool to explore development, aging and disease. M. Vernet and Z. Kapoula contributed equally to this work.     

Discharge patterns of neurons in the rostral superior colliculus of cat: activity related to fixation of visual and auditory targets

Neurons in the rostral superior colliculus (SC) of alert cats exhibit quasi-sustained discharge patterns related to the fixation of visual targets. Because some SC neurons also respond to auditory stimuli, we investigated whether there is a population of neurons in the rostral SC which is active in relation to fixation of both auditory and visual targets. We identified cells which were active with visual fixation and which continued to discharge if the fixation stimulus was briefly extinguished. The population of neurons exhibited similar discharge characteristics when the fixation stimulus was auditory. Few neurons were significantly more active during fixation of visual targets than during fixation of auditory targets. Most fixation neurons showed a diminished discharge rate during spontaneous (self-generated) saccadic eye movements away from a visual fixation stimulus, regardless of the direction of the saccade. this diminished discharge rate (or pause) typically began, on average, 12.2 ms before saccade onset and the duration of the pause was Ionger than the duration of the saccade. These observations are consistent with the hypothesis that increased discharge of these neurons is related to active fixation and that reductions in their activity are important for the generation of saccades. However, the lack of a precise relationship between pause duration and saccade duration implies that these neurons would be unlikely to project directly to the saccadic burst generator. The mean interval from the beginning of the pauses of fixation neurons to be beginning of the saccades away from fixation targets is also shorter than has been found in brainstem omnipause neurons. By analogy with the concept of a receptive field, agaze position error field depicts the range of gaze position error for which a cell is active. Although fixation neurons appear to encode the magnitude and direction of the error between visual targets and the visual axis, visual error fields at the end of fixating eye movements were significantly larger than those at stimulus onset. For auditory stimuli, this difference was not significant. These observations are compatible with a number of recent experiments indicating that neural signals of eye position are damped or delayed with respect to current eye position.     

Influence of removal of invisible fixation on the saccadic and manual gap effect

Saccadic and manual reactions to a peripherally presented target are facilitated by removing a central fixation stimulus shortly before a target onset (the gap effect). The present study examined the effects of removal of a visible and invisible fixation point on the saccadic gap effect and the manual gap effect. Participants were required to fixate a central fixation point and respond to a peripherally presented target as quickly and accurately as possible by making a saccade (Experiment 1) or pressing a corresponding key (Experiment 2). The fixation point was dichoptically presented, and visibility was manipulated by using binocular rivalry and continuous flash suppression technique. In both saccade and key-press tasks, removing the visible fixation strongly quickened the responses. Furthermore, the invisible fixation, which remained on the display but suppressed, significantly delayed the saccadic response. Contrarily, the invisible fixation had no effect on the manual task. These results indicate that partially different processes mediate the saccadic gap effect and the manual gap effect. In particular, unconscious processes might modulate an oculomotor-specific component of the saccadic gap effect, presumably via subcortical mechanisms.     

The control of vertical saccades in aged subjects

In real life we produce vertical saccades at different distances and eccentricities, and while our fixation is more or less actively engaged. The goal of this study is to examine vertical saccades in aged and young subjects, taking into consideration all these parameters. Eleven adults (20–28 years) and 11 aged subjects (63–83 years) were recruited. We used LED targets at 7.5° or 15°, up or down in four conditions: gap and overlap tasks, each done at two distances—at near (40 cm) and at far (150 cm). In the gap task fixation target extinguishes prior to target onset, while in the overlap condition it stays on after target onset; consequently, visual attention and fixation are employed differently in the two tasks. Eye movements were recorded with the Chronos video eye tracker. Results showed that vertical saccades were longer for aged subjects than for young adults under almost all conditions. For both aged and young subjects, latencies were shorter under the gap than under the overlap task. Latencies for eccentric targets at 15° were significantly longer than those at 7.5° but for aged subjects only; this effect was more pronounced for upward saccades under the overlap condition. Express type of latencies (80–120 ms) occurred frequently in the gap task and at similar rates for young adults (16%) and aged subjects (12%); in the overlap task express latencies were scarce in young adults (0.4%) and aged subjects (1.8%). Age deteriorates the ability to trigger regular volitional saccades but not the ability to produce express type of saccades. Latency increase with aging is attributed to the degeneration of central areas, e.g. oculomotor cortical areas involved in the initiation of vertical saccades.Grant/financial support: European Union (QLK6-CT-2002-00151: EUROKINESIS) and CNRS/CTI, Handicap contract CR:N.     

Saccade reaction time asymmetries during task-switching in pursuit tracking

We investigate how smooth pursuit eye movements affect the latencies of task-switching saccades. Participants had to alternate their foveal vision between a continuous pursuit task in the display center and a discrete object discrimination task in the periphery. The pursuit task was either carried out by following the target with the eyes only (ocular) or by steering an on-screen cursor with a joystick (oculomanual). We measured participants’ saccadic reaction times (SRTs) when foveal vision was shifted from the pursuit task to the discrimination task and back to the pursuit task. Our results show asymmetries in SRTs depending on the movement direction of the pursuit target: SRTs were generally shorter in the direction of pursuit. Specifically, SRTs from the pursuit target were shorter when the discrimination object appeared in the motion direction. SRTs to pursuit were shorter when the pursuit target moved away from the current fixation location. This result was independent of the type of smooth pursuit behavior that was performed by participants (ocular/oculomanual). The effects are discussed in regard to asymmetries in attention and processes that suppress saccades at the onset of pursuit.     

The gap effect and express saccades in the auditory modality

Latencies of eye movements to peripheral targets are reduced when there is a short delay (typically 200 ms) between the offset of a central visual fixation point and the target onset. This has been termed the gap effect. In addition, some subjects, usually with practice, exhibit a separate population of very short latency saccades, called express saccades. Both these phenomena have been attributed to disengagement of visual attention when the fixation point is extinguished. A competing theory of the gap effect attributes it to disengagement of oculomotor fixation during the temporal gap. It is known that auditory targets are effective in eliciting saccadic eye movements, and also that covert attention operates in the auditory modality. If the gap effect and express saccades are due to disengagement of spatial attention, both should persist in the auditory modality. However, fixation of gaze is largely under visual control. If the gap effect results from disengagement of fixation, then at least a reduced effect should be seen in the auditory modality. Human subjects performed the gap task and a control task in the dark, using auditory fixation points and saccadic targets, on five successive days. Despite this practice, express saccades were not observed. There was a reliable gap effect, but the reduction in saccadic latency was only 17 ms, compared with 32 ms for the same subjects in the visual modality. This suggests that about half the gap effect is due to disengagement of visual fixation. The remainder was not due to non-specific warning effects and could be attributed to offset of the auditory fixation stimulus. Received: 1 March 1996 / Accepted: 11 July 1997     

Combined auditory and visual stimuli facilitate head saccades in the barn owl (Tyto alba)

The barn owl naturally responds to an auditory or visual stimulus in its environment with a quick head turn toward the source. We measured these head saccades evoked by auditory, visual, and simultaneous, co-localized audiovisual stimuli to quantify multisensory interactions in the barn owl. Stimulus levels ranged from near to well above saccadic threshold. In accordance with previous human psychophysical findings, the owl's saccade reaction times (SRTs) and errors to unisensory stimuli were inversely related to stimulus strength. Auditory saccades characteristically had shorter reaction times but were less accurate than visual saccades. Audiovisual trials, over a large range of tested stimulus combinations, had auditory-like SRTs and visual-like errors, suggesting that barn owls are able to use both auditory and visual cues to produce saccades with the shortest possible SRT and greatest accuracy. These results support a model of sensory integration in which the faster modality initiates the saccade and the slower modality remains available to refine saccade trajectory.     

Task-dependent effects of social attention on saccadic reaction times

Previous research has shown that saccadic reaction times (SRTs) are shorter when a stimulus is flashed on the same side as the observed gaze direction of another individual. The gaze imitation hypothesis contends that observed gaze evokes the preparation of a saccade toward the same direction. Previous studies of this phenomenon have employed pro-saccade tasks in which the instructed saccade is directed toward the stimulus. In agreement with previous findings, we found that SRTs on pro-saccade trials were shorter when the stimulus appeared in the same direction as observed gaze. Here we also included anti-saccade trials in which subjects were required to look-away from a stimulus and toward its mirror position in the opposite visual field. The gaze imitation hypothesis predicts that subjects will have shorter SRTs on anti-saccade trials in which the stimulus appears opposite the observed gaze direction because they will have prepared already a saccade in that direction. However, contrary to the prediction of the gaze imitation hypothesis, we found that subjects had shorter SRTs on anti-saccade trials when the stimulus appeared in the same direction as observed gaze. Moreover, subjects also made more pro-saccade errors on anti-saccade trials in which the stimulus was presented opposite the observed gaze direction. The results of our study indicate that subjects prepared a saccade in the same direction as observed gaze on pro-saccade trials but opposite the observed gaze direction on anti-saccade trials. These findings suggest that the effect of social gaze cues on SRTs is task dependent.     

Separate populations of visually guided saccades in humans: reaction times and amplitudes

Summary The saccadic eye movements of 20 naive adults, 7 naive teenagers, 12 naive children, and 4 trained adult subjects were measured using two single target saccade tasks; the gap and the overlap task. In the gap task, the fixation point was switched off before the target occurred; in the overlap task it remained on until the end of each trial. The target position was randomly selected 4° to the left or 4° to the right of the fixation point. The subjects were instructed to look at the target when it appeared, not to react as fast as possible. They were not given any feedback about their performance. The results suggest that, in the gap task, most of the naive subjects exhibit at least two (the teenagers certainly three) clearly separated peaks in the distribution of the saccadic reaction times. The first peak occurs between 100 and 135 ms (express saccades), the second one between 140 and 180 ms (fast regular), and a third peak may follow at about 200 ms (slow regular). Other subjects did not show clear signs of two modes in the range of 100 to 180 ms, and still others did not produce any reaction times below 135 ms. In the overlap task as well three or even more peaks were obtained at about the same positions along the reaction time scale of many, but not all subjects. Group data as well as those of individual subjects were fitted by the superposition of three gaussian functions. Segregating the reaction time data into saccades that over- or undershoot the target indicated that express saccades almost never overshoot. The results are discussed in relation to the different neural processes preceding the initiation of visually-guided saccades.     

The effect of frontal eye field and superior colliculus lesions on saccadic latencies in the rhesus monkey

Rhesus monkeys were trained to make saccadic eye movements to visual targets using detection and discrimination paradigms in which they were required to make a saccade either to a solitary stimulus (detection) or to that same stimulus when it appeared simultaneously with several other stimuli (discrimination). The detection paradigm yielded a bimodal distribution of saccadic latencies with the faster mode peaking around 100 ms (express saccades); the introduction of a pause between the termination of the fixation spot and the onset of the target (gap) increased the frequency of express saccades. The discrimination paradigm, on the other hand, yielded only a unimodal distribution of latencies even when a gap was introduced, and there was no evidence for short-latency "express" saccades. In three monkeys either the frontal eye field or the superior colliculus was ablated unilaterally. Frontal eye field ablation had no discernible long-term effects on the distribution of saccadic latencies in either the detection or discrimination tasks. After unilateral collicular ablation, on the other hand, express saccades obtained in the detection paradigm were eliminated for eye movements contralateral to the lesion, leaving only a unimodal distribution of latencies. This deficit persisted throughout testing, which in one monkey continued for 9 mo. Express saccades were not observed again for saccades contralateral to the lesion, and the mean latency of the contralateral saccades was longer than the mean latency of the second peak for the ipsiversive saccades. The latency distribution of saccades ipsiversive to the collicular lesion was unaffected except for a few days after surgery, during which time an increase in the proportion of express saccades was evident. Saccades obtained with the discrimination paradigm yielded a small but reliable increase in saccadic latencies following collicular lesions, without altering the shape of the distribution. Unilateral muscimol injections into the superior colliculus produced results similar to those obtained immediately after collicular lesions: saccades contralateral to the injection site were strongly inhibited and showed increased saccadic latencies. This was accompanied by a decrease of ipsilateral saccadic latencies and an increase in the number of saccades falling into the express range. The results suggest that the superior colliculus is essential for the generation of short-latency (express) saccades and that the frontal eye fields do not play a significant role in shaping the distribution of saccadic latencies in the paradigms used in this study.(ABSTRACT TRUNCATED AT 400 WORDS)