Frontal eye field lesions impair predictive and visually-guided pursuit eye movements

Summary The study initially explored the frontal eye field's (FEF) control of predictive eye movements, i.e., eye movements driven by previous rather than current sensory signals. Five monkeys were trained to pursue horizontal target motion, including sinusoidal targets and random-walk targets which sometimes deviated from a sine motion. Some subjects also tracked other target trajectories and optokinetic motion. FEF ablations or cold lesions impaired predictive pursuit, but also degraded visually guided foveal pursuit of all targets. Unilateral lesions impaired pursuit of targets moving in both horizontal orbital fields and in both directions of movement. Saccadic estimates of target motion were generally accurate. The slow-phase velocity of optokinetic pursuit (collected after 54 s of OKN) also appeared normal. Pursuit recovered over 1–3 weeks after surgery but the deficits were then reinstated by removal of FEF in the other hemisphere. Thereafter, a slight deficit persisted for up to 10 weeks of observation in two subjects. The pattern of symptoms suggests that FEF lies subsequent to parietal area MST and prior to the pontine nuclei in controlling pursuit eye movements.     

Characteristics of “anti” saccades in man

Summary Four subjects — all made large numbers of Express saccades in the normal gap task — were instructed to make saccades in the direction opposite to the side where a visual stimulus appeared (anti task). Gap and overlap trials were used. Saccadic reaction time (SRT), velocity and amplitude of the corresponding eye movements were analysed and compared to those of saccades made in the normal task. The velocity of anti saccades was found to be slightly (up to 15%) but significantly slower in two subjects. The distributions of SRTs in normal gap tasks show a small group of anticipatory saccades (with SRT below 80 ms and slower velocities) followed by a group of saccades with fast reaction times between 80 ms and 120 ms (Express saccades) followed by another large group ranging up to 180 ms (regular saccades). In the gap anti task there are anticipatory saccades and saccades with SRTs above 100 ms; Express saccades are missing. The distribution of SRTs obtained in the overlap anti task was unimodal with a mean value of 231 ms as compared to 216 ms in the normal task. The introduction of the gap therefore clearly decreases the reaction times of the anti saccades. Control experiments show that the delay of anti saccades is not due to an interhemispheric transfer time but must be attributed to the saccade generating system taking more time to program a saccade to a position where no visual stimulus appears. These data are discussed as providing further evidence for the existence of a reflex-like pathway connecting the retina to the oculomotor nuclei mediating the Express saccade.     

Disconnection of parietal and occipital access to the saccadic oculomotor system

Summary The experiment explored the networks through which signals arising from visual areas of cortex control saccadic eye movements. Electrical stimulation of the inferior parietal and the occipital cortex (here termed the posterior eye fields) normally evokes saccadic eye movements. We replicated previous reports that these evoked eye movements ceased after large tectal ablations. This initial finding suggested that the posterior eye fields depended on a single route of access to the saccade generator, one descending through the superior colliculus (SC). On closer examination, the critical lesion appeared to be one which removed the SC and cut efferents from the frontal eye field (FEE) coursing nearby. Subsequently we confirmed that eye movements evoked from the posterior eye fields ceased after cooling the SC, or cutting its efferents- but only when one of these procedures was combined with FEF ablation. Thus, visual signals from the occipital and inferior parietal cortex have more than one, but perhaps only two routes of access to the oculomotor system. One passes through the superior colliculus, the other through the frontal eye field. Ancillary experiments revealed that inferior parietal and FEF ablations, alone or combined, do not disrupt saccades evoked from the occipital lobe. Striate and prestriate areas can therefore use their own direct input to the SC or to the basal ganglia to drive saccadic eye movements.Abbreviations for Nuclei C interstitial, of Cajal - CL central lateral - CM centromedianum - D Darkschewitsch - HL lateral habenula - IC inferior colliculus - LD lateral dorsal - LP lateral posterior nucl - MD medial dorsal - MG medial geniculate - MR midbrain reticular formation - NPA nuc. of the pretectal area - NPC nuc. of the posterior commissure - PC posterior commissure - PF parafascicular - PI inferior pulvinar - PL lateral pulvinar - PM medial pulvinar - PO oral pulvinar - R red - SG suprageniculate - SL sublentiform - SN substantia nigra - VPL ventral posterolateral - VPM ventral posteromedial - ZI zona incerta - III oculomotor - IV trochlear Supported by grants NIH EY02941 and EY04005, NSF BNS8603915     

Smooth eye movements evoked by electrical stimulation of the cat's superior colliculus

Head-fixed gaze shifts were evoked by electrical stimulation of the deeper layers of the cat superior colliculus (SC). After a short latency, saccades were triggered with kinematics similar to those of visually guided saccades. When electrical stimulation was maintained for more than 150–200 ms, postsaccadic smooth eye movements (SEMs) were observed. These movements were characterized by a period of approximately constant velocity following the evoked saccade. Depending on electrode position, a single saccade followed by a slow displacement or a staircase of saccades interspersed by SEMs were evoked. Mean velocity decreased with increasing deviation of the eye in the orbit in the direction of the movement. In the situation where a single evoked saccade was followed by a smooth movement, the duration of the latter depended on the duration of the stimulation train. In the situation where evoked saccades converged towards a restricted region of the visual field (goal-directed or craniocentric saccades), the SEMs were directed towards the centre of this region and their mean velocity decreased as the eye approached the goal. The direction of induced SEMs depended on the site of stimulation, as is the case for saccadic eye movements, and was not modified by stimulation parameters (place code). On the other hand, mean velocity of the movements depended on the site of stimulation and on the frequency and intensity of the current (rate code), as reported for saccades in the cat. The kinematics of these postsaccadic SEMs are similar to the kinematics of slow, postsaccadic correction observed during visually triggered gaze shifts of the alert cat. These results support the hypothesis that the SC is not exclusively implicated in the control of fast refixation of gaze but also in controlling postsaccadic conjugate slow eye movements in the cat.     

A frontal cortical potential associated with saccades in humans

Summary We describe a frontal EEG potential which begins 25–35 ms before intentional saccadic eye movement. It consists of a 15–20 volt monophasic positive waveform with peak during or just after movement, and returns to EEG baseline 150–200 ms after its onset. The waveform is largest at a midline position just anterior to F_Z (10–20 system), is independent of visual input such as fixation guides, and is not related to saccade direction or amplitude. The potential is difficult to observe in some subjects and is independent of the pre-saccadic spike potential. It may be related to the discharge of single cortical neurons that signal the initiation of saccadic movements, but not their exact metrics; a possible generator is the supplementary eye fields of the dorsomedial prefrontal cortex.     

The gap effect and inhibition of return: interactive effects on eye movement latencies

Three experiments are reported with two types of manipulations that are known to affect the latency with which subjects can initiate saccadic eye movements. The first manipulation involves the temporal relation between the offset of a visual fixation point and the onset of a peripheral target (the gap effect). The second manipulation involves the prior allocation and removal of visual attention (inhibition of return). In two experiments, the gap effect was smaller for saccades to previously attended locations than to previously unattended locations. The results suggest an important link between the two phenomena and provide new insights into the brain mechanisms underlying visual attention and eye movements.     

Saccade and blinking evoked by microstimulation of the posterior parietal association cortex of the monkey

Summary Electrical stimulation with microelectrodes of the posterior parietal association cortex in alert behaving monkeys elicited saccadic eye movements and blinking. The sites in which saccades were elicited by electrical stimulation were concentrated in the anteromedial part of area 7a, especially in the posterior bank of the intraparietal sulcus, in a region which sends efferent projections to the frontal eye field and the superior colliculus, but they were also found in the posterolateral part of area 7a. Compared with the frontal eye fields and the superior colliculus, the threshold current for eliciting saccades was relatively high, on the average 86 A. Moreover, the elicitation of saccade was inconsistent even with suprathreshold stimulation and suppressed during visual fixation. Latencies of the saccades were relatively long, on the average 50 ms; they were longer in the posterolateral part than in the anteromedial part. Direction and amplitude of evoked saccades depended on the site of stimulation, but was independent of eye position in most cases. However, goal-directed saccades which depended on initial eye position were elicited in three penetrations in the posterolateral part of area 7a. Blinking was elicited mainly in the lateral part of area 7a. The threshold of blinking was 70 A and the latency was 50 ms on the average. In contrast to saccades, blinking was elicited constantly with each stimulus even during attentive fixation. We occasionally recorded single unit activity at the site of stimulation with the same electrodes. More than half of the units recorded at the site of blinking responded to approaching visual stimulus. These results suggest that area 7a participates indirectly in the control of saccades by way of its connection to the frontal eye fields or the superior colliculus, and it may also play an important role in blinking in response to a visual threat.Prof. J. Hyvärinen died on February 26, 1983     

The ability to produce express saccades as a function of gap interval among schizophrenia patients

The ability to produce express sacccades is associated with adequate functioning of saccadic burst cells in the superior colliculus. Saccadic burst cells appear to be under the inhibitory control of both the collicular and the dorsolateral frontal fixation systems. Twenty schizophrenia patients and 20 nonpsychiatric subjects were presented a saccade task that included five different gap intervals (0, 100, 200, 300, and 400 ms) between fixation point offset and peripheral target onset (at ±4°). All subjects generated the highest frequency of express saccades in trials with a gap interval of 200 ms. Schizophrenia patients had an increased frequency of express saccades across gap intervals, especially for targets presented in the right visual field. The groups did not differ in the percentages of anticipatory saccades or saccadic amplitudes. These results suggest that schizophrenia patients' saccadic burst cells in the superior colliculus are functioning adequately, but may be consistent with dys-function of dorsolateral frontal cortex and/or its interconnecting subcortical circuitry.     

Goal-directed vestibulo-ocular function in man: gaze stabilization by slow-phase and saccadic eye movements

Summary Vestibular function was examined during passive head movements having profiles that approximated the low-to-intermediate range of natural self-generated movements (10–220°/s peak velocity, about 0.5 s duration). A seated subject looked at a point target on the wall, the lights were extinguished and the chair was briefly turned while the subject tried to look at the just-viewed point. The chair was stopped, the lights were turned on again and the target was re-fixated, if necessary. Ocular stabilization was characterized (1) by net stabilization that was due to the combined effects of both slow-phase and rapid (saccadic or quick-phase) eye movements, (2) by cumulative-slow-phase stabilization that was due to slow-phase eye movements, and (3) by cumulative-saccadic stabilization that was due to effects of all rapid eye movements. It was found that both slow-phase and saccadic eye movements tended to keep the eyes on the actual unseen target. During repeatedly applied head movements, net and cumulative-slow-phase stabilization tended to be almost perfect. However, the average magnitude of the error in net stabilization (i.e., deviation from perfection) was always less than the corresponding error in slow-phase stabilization. This occurred because in a given turn, saccadic movements tended to supplement deficient slow-phase movements and to decrement excessive slow-phases. In 4 of 5 subjects, cumulative-saccadic stabilization tended to equal the error in cumulative-slow-phase stabilization. All results were unaffected by head velocities up to ±220°/s. It was concluded that these saccades tended to stabilize gaze (eye + head) in space during head movements in total darkness.     

Eye position signals in human saccadic processing

Summary 1. We studied saccades to briefly flashed targets in 8 human subjects. The target flash occurred (i) during smooth pursuit (ramp-flash), (ii) just before a saccade to another target (step-flash), or (iii) during steady fixation (flash only). All lights were extinguished after the target flash so that smooth pursuit or saccadic eye movements occurred during the interval of complete darkness between the target flash and the saccade to it. We compared these saccades to those made without intervening eye movement (flash only), and quantified the extent to which the saccadic system compensated for the change in eye position that occurred during the dark interval. 2. Saccades to control flashes were reasonably accurate (mean gain 0.87) and consistent. Compensation for the intervening eye movement in the ramp-flash and step-flash paradigms was highly variable from trial to trial. On average, subjects compensated for 27% of the intervening pursuit eye movement on ramp-flash trials and for 58% of intervening saccadic movement on step-flash trials. 3. Multiple regression analysis showed that the variability did not depend on factors such as variations in underlying saccadic gain, response latency, timing of stimuli or size of the required response. We conclude that this variability is intrinsic to saccadic responses that require the use of an eye position signal. 4. These results show that an eye position signal is available to the saccadic system but that this signal has low fidelity. The high variability and low fidelity of the eye position signal suggest that the saccadic system does not normally operate in spatial coordinates, which require the use of an accurate eye position signal, but rather in retinal coordinates.     

Dorsal premotor areas of nonhuman primate: functional flexibility in time domain

A voluntary motor act requires recognition of the informational content of an instruction. An instruction may contain spatial and temporal information. The recently proved role of the monkey frontal cortex in time computation, as well as in motor preparation and motor learning, suggested that we investigate the relationship between premotor neuron discharges and the temporal feature of the visual instructions. To this purpose, we manipulated the duration of an instructional cue in a visuomotor task while recording unit activity. We found two types of premotor neurons characterised by a discharge varying in relation to the duration of the cue: (1) motor-linked neurons, with a specific premotor activity constantly bounded to the motor act; (2) short-term encoders neurons, with a premotor activity depending on the cue duration. The cue duration was the critical factor in determining the behaviour of the short-term encoders cells: when the cue ranged from 0.5 s to 1 s, they presented a preparatory activity; when the cue was longer, up to 2 s, they lost their preparatory activity; when the cue was blinked the cells anticipated their discharge. The activity changed in few trials. These data confirm and highlight the role of frontal cortex in encoding specific cues with a temporal flexibility, which may be the expression of temporal learning and represent an extended aspect of cortical plasticity in time domain.     

Reversible inactivation of macaque frontal eye field

 The macaque frontal eye field (FEF) is involved in the generation of saccadic eye movements and fixations. To better understand the role of the FEF, we reversibly inactivated a portion of it while a monkey made saccades and fixations in response to visual stimuli. Lidocaine was infused into a FEF and neural inactivation was monitored with a nearby microelectrode. We used two saccadic tasks. In the delay task, a target was presented and then extinguished, but the monkey was not allowed to make a saccade to its location until a cue to move was given. In the step task, the monkey was allowed to look at a target as soon as it appeared. During FEF inactivation, monkeys were severely impaired at making saccades to locations of extinguished contralateral targets in the delay task. They were similarly impaired at making saccades to locations of contralateral targets in the step task if the target was flashed for ≤100 ms, such that it was gone before the saccade was initiated. Deficits included increases in saccadic latency, increases in saccadic error, and increases in the frequency of trials in which a saccade was not made. We varied the initial fixation location and found that the impairment specifically affected contraversive saccades rather than affecting all saccades made into head-centered contralateral space. Monkeys were impaired only slightly at making saccades to contralateral targets in the step task if the target duration was 1000 ms, such that the target was present during the saccade: latency increased, but increases in saccadic error were mild and increases in the frequency of trials in which a saccade was not made were insignificant. During FEF inactivation there usually was a direct correlation between the latency and the error of saccades made in response to contralateral targets. In the delay task, FEF inactivation increased the frequency of making premature saccades to ipsilateral targets. FEF inactivation had inconsistent and mild effects on saccadic peak velocity. FEF inactivation caused impairments in the ability to fixate lights steadily in contralateral space. FEF inactivation always caused an ipsiversive deviation of the eyes in darkness. In summary, our results suggest that the FEF plays major roles in (1) generating contraversive saccades to locations of extinguished or flashed targets, (2) maintaining contralateral fixations, and (3) suppressing inappropriate ipsiversive saccades. Received: 2 February 1996 / Accepted: 26 February 1997     

Ear and eye representation in the frontal cortex,area 8b,of the macaque monkey: an electrophysiological study

We evoked both ear and eye movements in area 8b, the rostral area of frontal cortex, in two monkeys. In some sites it was possible to evoke only ear movements or only eye movements; in other locations we evoked both ear and eye movements by varying the intensity of electrical stimulation. The electrically evoked ear movements were forward, or backward, or oblique (upward-forward; upward-backward). In two penetrations the ear movements were bilateral, in the other penetrations they were contralateral. Ipsilateral ear movements were not observed. The evoked eye movements were mainly fixed-vector saccades, contralateral and with an upward orientation of about 45°. If we considered only the sites where the threshold was equal to or lower than 50 A, the stimulation of this area evoked mainly ear movements. In addition we recorded the electrical activity of 195 neurons. Of these neurons: 74% (145/195) discharged before ear movements (ear cells); 20% (40/195) discharged before ear and eye movements (ear-eye cells); 5% (10/195) discharged only before eye movements (eye cells). Ninety-one percent (132/145) of ear cells presented a preferred direction; 90% (36/40) of ear-eye cells presented a preferred direction for ear movements, and 15% (6/40) presented a preferred direction for eye movements. Eighty-five percent (34/40) of cells did not present a preferred direction for visually guided saccades and were active when the monkey made saccades toward the unlit targets (checking saccades). Our results show that a field of area 8b is related to ear movements and to eye-ear movements. The findings that it is possible to obtain both ear and eye movements with low-intensity currents and that there are cells firing for the two types of movements suggest that area 8b may be involved in the orientation and coordination of both ear and eye. This area might be considered a rostral extension of supplementary eye field (SEF) or a different region. However, based on its distinct functional characteristics and connectivity, it is probably better regarded as a separate field. Regardless, the combination of 8b and SEF may constitute a cortical center for orienting processes.     

Adjustment of saccade characteristics during head movements

Summary Saccade characteristics have been studied during coordinated eyehead movements in monkeys. Amplitude, duration, and peak velocity of saccades with head turning were compared with saccades executed while the head was artificially restrained. The results indicate that the saccade characteristics are modulated as a function of head movement, hence the gaze movement (eye+head) exactly matches saccades with head fixed. Saccade modulation is achieved by way of negative vestibulo-ocular feedback. The neck proprioceptors, because of their longer latency, are effective only if the head starts moving prior to the onset of saccade. It is concluded that saccades made with head turning are not ballistic movements because their trajectory is not entirely predetermined by a central command.     

Effects of frontal eye field stimulation upon activities of the lateral geniculate body of the cat

Summary Effects of electrical stimulation of the frontal eye field (FEF) upon activities of the lateral geniculate body (LG) were studied in encéphale isolé cats. In some experiments the effects were examined by recording field responses of the dorsal nucleus of LG (LGd) and the visual cortex (VC) to electrical stimulation of the optic chiasm (OX). Conditioning repetitive stimulation of FEF exerted no significant effects on the r₁ wave of LGd responses but had a facilitatory effect on the r₂ wave. FEF-induced facilitation of VC responses was prominent in the late postsynaptic components. These effects had latencies of 50–100 msec and durations of 200–500 msec. Transection of the midbrain showed that most of the FEF-effect was not mediated via the brainstem reticular formation.Extracellular unitary recordings were made from 125 neurons, of which 91 were LGd neurons, 23 neurons of the caudal part of the thalamic reticular nucleus (TRc) and 11 neurons of the ventral nucleus of LG (LGv). In 30 of 87 LGd relay neurons FEF stimuli increased response probabilities to OX stimuli and their spontaneous discharges. These FEF-facilitated LGd neurons were distinguished from the non-affected ones in that the former had longer OX-latencies than the latter. The FEF-facilitated neurons probably correspond to X neurons of LGd.In 17 TRc neurons the effects were inhibitory. Their time courses were similar to those of the facilitation in the LGd relay neurons. Seven LGv neurons received facilitatory effects from FEF. Among them 5 neurons showed short-latency (6.7–17 msec) responses to FEF single shocks.The FEF sites inducing conjugate lateral eye movements exerted stronger facilitatory effects than those inducing upward or centering eye movements did.It is suggested that the effects may subserve to cancel the inhibitory convergence onto X-cells just after saccadic eye movements so as to improve visual information transmission through LGd during the eye fixation.     

Cognitive spatial-motor processes

Summary We studied the activity of 123 cells in the arm area of the motor cortex of three rhesus monkeys while the animals performed a 2-dimensional (2-D) step-tracking task with or without a delay interposed between a directional cue and a movement triggering signal. Movements of equal amplitude were made in eight directions on a planar working surface, from a central point to targets located equidistantly on a circle. The appearance of the target served as the cue, and its dimming, after a variable period of time (0.5–3.2 s), as the go stimulus to trigger the movement to the target; in a separate task, the target light appeared dim and the monkey moved its hand towards it without waiting. Population histograms were constructed for each direction after the spike trains of single trials were aligned to the onset of the cue. A significant increase (3–4×) in the population activity was observed 80–120 ms following the cue onset; since the minimum delay was 500 ms and the average reaction time approximately 300 ms, this increase in population activity occurred at least 680–720 ms before the onset of movement. A directional analysis (Georgopoulos et al. 1983, 1984) of the changes in population activity revealed that the population vector during the delay period pointed in the direction of movement that was to be made later.     

Effect of posture on arterial baroreflex control of heart rate in humans

Summary Altered baroreflex function may contribute to the cardiovascular changes associated with weightlessness. Since central blood volume (CBV) increases during simulated weightlessness, we have examined the possibility that acute changes in CBV may modify baroreceptor function. We used graded head-up tilt (HUT) and head-down tilt (HDT) to induce changes in CBV, and neck suction to stimulte carotid baroreceptors, in 6 subjects. The increase in pulse interval induced by a negative pressure of 8.2 kPa (62 mm Hg) imposed for 10 s while supine was compared with the increase while tilted for 8 min at ± 15, ± 30 and ± 45. During HDT at 15 the pulse interval over the first 5 cardiac cycles following suction onset was 51 ± (SEM) 18 ms longer (p<0.05), at 30 it was 61±20 ms longer (p<0.05), and at 45 it was 74±35 ms longer (p<0.01), compared with supine. During HUT at 15 the pulse interval was 25±9 ms shorter (p<0.05) than when supine, but was not significantly different at 30 and 45. These responses occurred independently of changes in brachial blood pressure. Attenuation was also observed after 5 min (56±17 ms; <0.05), and after 40 min (25±9 ms; p<0.05) of 60 HUT compared with supine. We conclude that posture does modify arterial baroreflex control of heart rate. If this occurs primarily as a result of a change in CBV, then the acute effect of weightlessness may be an accentuation, not an attenuation, of baroreflex function.M. H. Harrison was a National Research Council postdoctoral research fellow on leave from the Ministry of Defence, UK     

Motion processing for saccadic eye movements in humans

Summary 1. We studied the latencies and amplitudes of saccades to moving targets in normal human subjects. Targets underwent ramp or step-ramp motions. The goal was to determine how the saccadic system uses information about target velocity. 2. For simple ramp motion saccadic latency decreased as target speed increased. A threshold distance model, which assumes that the target has to move a minimum distance before saccadic processing starts, provided a good fit to the responses of all four subjects and explains discrepancies between previously published findings. 3. A double step experiment showed that target position may have some effect on saccadic amplitude when sampled 70 ms before saccade onset, but it must be sampled at least 140 ms before onset for an accurate saccade to occur. 4. Saccades to simple ramp targets approximated the target position 55 ms before saccade onset. Based on our double step results, this is more compensation than possible by a simple position estimate and implies extrapolation of target motion by the saccadic system. The lack of complete compensation may be due to an underestimate of the target speed and/or of the saccadic latency. 5. A delayed-saccade paradigm resulted in saccades with a longer, constant latency and allowed longer viewing of target motion. These saccades accounted for all but 20 ms of target motion, suggesting that with more processing time of target motion a better extrapolation may be generated. 6. In a step-ramp paradigm the target stepped in one direction, then moved smoothly in the opposite direction. Saccades in this paradigm could be made in either the direction of the step or in the direction of target motion: the direction and latency were determined solely by the time at which the target crossed the fixation point. This time must be calculated from target speed and position, implying that the saccadic system must use speed information to adjust latency or to cancel unnecessary saccades.     

Responses of cat vestibular neurons to stimulation of the frontal cortex

Summary To study the neural basis for the regulation of vestibulocollic reflexes during voluntary head movements, the effects of stimulation of the precruciate cortex near the presylvian sulcus (neck area of the motor cortex) and the frontal eye fields (FEF) on vestibular neurons were studied in cerebellectomized cats anesthetized with chloralose. Neurons were recorded in the medial and descending vestibular nuclei and antidromically identified from C₁. Stimulation of the FEF and precruciate cortex fired 29 and 13% of neurons that did not exhibit spontaneous activity. About 80% of spontaneously discharging neurons were influenced by stimulation of either of the two. Stimulation of the precruciate cortex or FEF suppressed or facilitated labyrinthine evoked monosynaptic activation of vestibulospinal neurons, suggesting that the frontal cortical neurons have the properties to regulate the vestibulocollic reflexes.     

Human express saccades: extremely short reaction times of goal directed eye movements

Summary Human subjects were asked to execute a saccade from a central fixation point to a peripheral target at the time of its onset. When the fixation point is turned off some time ( 200 ms) before target onset, such that there is a gap where subjects see nothing, the distribution of their saccadic reaction times is bimodal with one narrow peak around 100 ms (express saccades) and another peak around 150 ms (regular saccades) measured from the onset of the target. Express saccades have been described earlier for the monkey.