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.     

Express saccades in cat: effects of task and target modality

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

Influence of gap and overlap paradigms on saccade latencies and vergence eye movements in seven-year-old children

The latency of eye movements is influenced by the fixation task; when the fixation stimulus is switched off before the target presentation (gap paradigm) the latency becomes short and express movements occur. In contrast, when the fixation stimulus remains on when the target appears (overlap paradigm), eye movement latency is longer. Several previous studies have shown increased rates of express saccades in children; however the presence of an express type of latency for vergence and combined movements in children has never been explored. The present study examines the effects of the gap and the overlap paradigms on horizontal saccades at far (150 cm) and at close (20 cm) viewing distances, on vergence along the median plane, and on saccades combined with convergence or divergence in 15 normal seven-year-old children. The results show that the gap paradigm produced shorter latency for all eye movements than the overlap paradigm, but the difference was only significant for saccades at close viewing distances, for divergence (pure and combined), and for saccades combined with vergence. The gap paradigm produced significantly higher rates of express latencies for saccades at close viewing distances, for divergence, and for saccades combined with divergence; in contrast, the frequencies of express latencies for saccades at far viewing distances and for convergence (pure or combined) were similar in the gap and the overlap paradigms. Interestingly, the rate of anticipatory latencies (<80 ms) was particularly high for divergence in the gap paradigm. Our collective findings suggest that the initiation of saccades at close viewing distances and of divergence is more reflexive, particularly in the gap paradigm. The finding of frequent anticipatory divergence that occurs at similar rates for seven-year-old children (this study) and for adults (Coubard et al., 2004, Exp Brain Res 154:368–381) indicates that predictive initiation of divergence is dominant.     

Gap effects on saccade and vergence latency

To explore visual space, we make saccades, vergence, and, most frequently, combined saccade–vergence eye movements. The initiation of saccades is well studied, while that of vergence is less explored. Saccade latency is influenced by the fixation task: when the target appears simultaneously with the offset of the fixation point, latencies tend to be regular, whereas the introduction of a gap period before target onset causes the emergence of express latencies (80- to 120-ms). This study examines in ten normal adults whether the gap paradigm has a similar effect on the latency of vergence and combined eye movements. The second goal is to identify contextual factors that favor the emergence of short latencies, by comparing a condition in which gap and simultaneous trials were performed in separate blocks (pure blocks) with a condition in which the two types of trials were interleaved randomly (mixed blocks). The results are: (1) the gap paradigm reduced similarly (by approximately –30 ms) the mean latency of saccades, convergence, divergence, and both the saccadic and vergence components of combined eye movements; (2) the gap paradigm was responsible for the emergence of 80– to 120-ms latencies for saccades and divergence (pure or combined), but rarely for convergence; (3) inspection of the latency distributions showed that such short latencies formed a clearly distinct population, different from anticipatory responses or regular latencies, for saccades (pure or combined) but not for pure vergence; importantly, distinct express latencies were found also for the convergence and divergence components of combined eye movements; (4) no difference was found for the group of subjects between pure and mixed blocks, but the latter yielded shorter latencies for some subjects, suggesting an idiosyncratic phenomenon. We suggest that distinct express latencies are specific to saccades and could correspond to a specific mode of saccade initiation. Interestingly, the express mode of triggering can be transferred to the vergence component in the ecological condition in which saccade is combined with vergence.     

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.     

The effect of transcranial magnetic stimulation on the latencies of vertical saccades

In this study, we investigated the effect of transcranial magnetic stimulation (TMS) over the right posterior parietal cortex (PPC) on the latency of two different types of visually-guided vertical saccades: reflexive saccades triggered by the sudden onset of a target, and saccades towards target locations known in advance. For this reason, we used two oculomotor tasks: a gap and a delay task, respectively. Nine normal subjects performed vertical saccades at ±7.5 and ±15°. TMS was applied at 80 and 100 ms after target onset in the gap task, and after fixation offset in the delay task. Without TMS, we confirmed a latency asymmetry in the gap task favouring upward saccades at the lower eccentricity (7.5°), and a latency symmetry in the delay task. TMS increased the latencies of all saccades in the delay task, when delivered at 100 ms. This effect was mostly pronounced for downward saccades at 7.5°. As a result, saccade latencies showed an asymmetry in this condition, similar to the one observed in the gap task without TMS. The gap task with TMS resulted in a variable latency distribution and no significant overall effect on saccade latency. Our results indicate that the right PPC is involved in the initiation of vertical saccades in the delay task, and that this involvement appears to be enhanced for downward saccades. A conclusion for the involvement of this area in the gap task could not be drawn from this study.     

Development of prosaccade and antisaccade task performance in participants aged 6 to 26 years

There are few studies on the development of oculomotor functions during childhood. B. Fischer, M. Biscaldi, and S. Gezeck (1997) reported improvement of antisaccade task performance between ages 6 and 16 years. The present study is a replication and extension of those results. In three age groups (6-7, 10-11, 18-26 years), saccades during pro- and antisaccade tasks with 200-ms gap and overlap and during a fixation task were measured. Adults exhibited faster saccades and less prosaccades during the antisaccade tasks than 10-11-year-old children; these two groups had faster saccades during all tasks and less prosaccades during the anti- and the fixation task than 6-7-year-old subjects. Both children groups made more express saccades than adults. Results suggest different degrees of age-related improvement for different saccadic parameters, the effects being greatest for prosaccade inhibition during the antisaccade task and in line with the assumed protracted development of prefrontal functions.     

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.     

The role of fixation point and subjects' readiness in the occurrence of express saccades as revealed by the self-initiation paradigm

The role of fixation and the subjects' response preparedness in producing express saccades were explored in seven human subjects. The occurrence frequencies of the express saccades were compared in the overlap (continuous presentation of fixation point), gap (fixation point offset 0-400 ms prior to target onset) and no-fixation tasks under the conventional and self-initiation paradigms. In the latter paradigm, the subjects, when ready, touched a sensor in order to ignite the target lamp with a delay time of 0-400 ms (target onset delay time). Therefore, the subjects' response preparedness might be expected to be higher than that in the normal paradigm and equated in each subject at the time when the subjects touched a sensor regardless of the paradigms. Although express saccades were produced neither in the normal overlap nor in the normal no-fixation tasks, they could be produced at the rate of 24 and 48% in the overlap and no-fixation tasks under the self-initiation paradigm, respectively. The highest occurrence frequency of express saccades was obtained when the gap paradigm was combined with the self-initiation paradigm with a delay time of 100 ms (62%). The value was higher by 20% than in the normal gap task. At a target onset delay time of 0 ms under the self-initiation paradigm, the occurrence frequency of express saccades was higher in the overlap task than in the gap task. These results suggest that the subjects' response preparedness has a potentiality to produce express saccades without fixation point offset and that fixation point offset at the same time of the target stimulus onset has an interference, rather than facilitatory, influence on the generation of express saccades.     

Reversible inactivation of macaque dorsomedial frontal cortex: effects on saccades and fixations

 Neural recording and electrical stimulation results suggest that the dorsomedial frontal cortex (DMFC) of macaque is involved in oculomotor behavior. We reversibly inactivated the DMFC using lidocaine and examined how saccadic eye movements and fixations were affected. The inactivation methods and monkeys were the same as those used in a previous study of the frontal eye field (FEF), another frontal oculomotor region. In the first stage of the present study, monkeys performed tasks that required the generation of single saccades and fixations. During 15 DMFC inactivations, we found only mild, infrequent deficits. This contrasts with our prior finding that FEF inactivation causes severe, reliable deficits in performance of these tasks. In the second stage of the study, we investigated whether DMFC inactivation affected behavior when a monkey was required to make more than one saccade and fixation. We used a double-step task: two targets were flashed in rapid succession and the monkey had to make two saccades to foveate the target locations. In each of five experiments, DMFC inactivation caused a moderate, significant deficit. Both ipsi- and contraversive saccades were disrupted. In two experiments, the first saccades were made to the wrong place and had increased latencies. In one experiment, first saccades were unaffected, but second saccades were made to the wrong place and had increased latencies. In the remaining two experiments, specific reasons for the deficit were not detected. Saline infusions into DMFC had no effect. Inactivation of FEF caused a larger double-step deficit than did inactivation of DMFC. The FEF inactivation impaired contraversive first or second saccades of the sequence. In conclusion, our results suggest that the DMFC makes an important contribution to generating sequential saccades and fixations but not single saccades and fixations. Compared with the FEF, the DMFC has a weaker, less directional, more task-dependent oculomotor influence. Received: 12 January 1998 / Accepted: 17 July 1998     

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.     

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.     

Expectations can modulate the frequency and timing of multiple saccades: a TMS study

This study was undertaken to determine if target predictability could modulate saccadic planning and timing at the level of the frontal eye fields (FEF). To this end, healthy participants performed two gap saccade tasks in which the targets were displaced left or right of the midline in either a predictable or a random fashion. Additionally, half of the participants were informed about this manipulation. Single pulse transcranial magnetic stimulation (TMS) was applied to the left FEF before, during, or after the target onset. We examined both the saccade latency and the frequency of multiple saccades (MS) (i.e., saccades that covered <90 % of the distance to the target and were subsequently followed by a corrective saccade). Findings revealed that saccadic reaction times were quickest to the more predictable target side and also confirmed that MS were released more quickly than single saccades. Further, the frequency of MS differed between target locations; higher frequencies of MS were found on the unpredictable side, showing more vulnerability to TMS disruption. In conclusion, we have demonstrated that target predictability modulates saccade planning and that this modulation takes place at least in part in the FEF. The influence of FEF in these processes is observed both in the latencies with which saccades are executed and in the timing and characteristics of the multiple saccades that are observed under different task constraints. Finally, the timing of the FEF contribution also appears to be influenced by the manipulation of target predictability. Each of these observations serves to further clarify the role of the human FEF in saccade planning.     

Differing effects of prosaccades and antisaccades on postural stability

The goal of the study was to examine the effect of different types of eye movements on postural stability. Ten healthy young adults (25 ± 3 years) participated in the study. Postural control was measured by the TechnoConcept© platform and recorded in Standard Romberg and Tandem Romberg conditions while participants performed five oculomotor tasks: two fixation tasks (central fixation cross, without and with distractors), two prosaccade tasks toward peripheral targets displayed 4° to the left or to the right of the fixation cross (reactive saccades induced by a gap 0 ms paradigm and voluntary saccades induced by an overlap 600 ms paradigm) and one antisaccade task (voluntary saccade made in the opposite direction of the visual target). The surface, the length, and the mean speed of the center of pressure were analyzed. We found that saccadic eye movements improved postural stability with respect to the fixation tasks. Furthermore, antisaccades were found to decrease postural stability compared to prosaccades (reactive as well as voluntary saccades). This result is in line with the U-shaped nonlinear model described by Lacour et al. (Neurophysiol Clin 38:411–421, 2008), showing that a secondary task performed during a postural task could increase (prosaccade task) or decrease (antisacade task) postural stability depending on its complexity. We suggest that the different degree of attentional resources needed for performing prosaccade or antisaccade tasks are, most likely, responsible for the different effect on postural control.     

Fixation offset facilitates saccades and manual reaching for single but not multiple target displays

Turning off a fixation point, typically for 200 ms, before the onset of a peripheral target substantially reduces saccadic reaction times. This facilitatory effect generated by an inserted temporal gap between fixation offset and the target appearance is called the “gap” effect [J Opt Soc Am 57:1030–1033, 1967]. We show that the gap reduces the initial latency of both saccades and manual pointing in single and multiple target displays. Yet, in multiple target displays, the gap increased the movement duration because eye or hand movements were frequently misdirected toward distractors so that the trajectory had to be corrected. Thus, in spite of the shortened latency, the total time for trial completion was not shortened in multiple target displays, whereas it was reduced in single target displays. This selective gap effect for a single target was not restricted to goal-directed motor tasks because perceptual discrimination tasks, where no motor response is required, also demonstrated the gap effect only for single target displays. Our results suggest that the gap may facilitate attentional disengagement, but it does not help target selection in motor and perceptual discrimination tasks, where the allocation of attention to the target is required.     

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.     

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     

Muscimol-induced inactivation of monkey frontal eye field: effects on visually and memory-guided saccades.

Muscimol-induced inactivation of the monkey frontal eye field: effects on visually and memory-guided saccades. Although neurophysiological, anatomic, and imaging evidence suggest that the frontal eye field (FEF) participates in the generation of eye movements, chronic lesions of the FEF in both humans and monkeys appear to cause only minor deficits in visually guided saccade generation. Stronger effects are observed when subjects are tested in tasks with more cognitive requirements. We tested oculomotor function after acutely inactivating regions of the FEF to minimize the effects of plasticity and reallocation of function after the loss of the FEF and gain more insight into the FEF contribution to the guidance of eye movements in the intact brain. Inactivation was induced by microinjecting muscimol directly into physiologically defined sites in the FEF of three monkeys. FEF inactivation severely impaired the monkeys' performance of both visually guided and memory-guided saccades. The monkeys initiated fewer saccades to the retinotopic representation of the inactivated FEF site than to any other location in the visual field. The saccades that were initiated had longer latencies, slower velocities, and larger targeting errors than controls. These effects were present both for visually guided and for memory-guided saccades, although the memory-guided saccades were more disrupted. Initially, the effects were restricted spatially, concentrating around the retinotopic representation at the center of the inactivated site, but, during the course of several hours, these effects spread to flanking representations. Predictability of target location and motivation of the monkey also affected saccadic performance. For memory-guided saccades, increases in the time during which the monkey had to remember the spatial location of a target resulted in further decreases in the accuracy of the saccades and in smaller peak velocities, suggesting a progressive loss of the capacity to maintain a representation of target location in relation to the fovea after FEF inactivation. In addition, the monkeys frequently made premature saccades to targets in the hemifield ipsilateral to the injection site when performing the memory task, indicating a deficit in the control of fixation that could be a consequence of an imbalance between ipsilateral and contralateral FEF activity after the injection. There was also a progressive loss of fixation accuracy, and the monkeys tended to restrict spontaneous visual scanning to the ipsilateral hemifield. These results emphasize the strong role of the FEF in the intact monkey in the generation of all voluntary saccadic eye movements, as well as in the control of fixation.     

Fixation and saccade control in an express-saccade maker

In express-saccade makers a large incidence of express saccades (latencies around 100 ms) is paralleled by a reduced ability to suppress saccade generation when required. Such a behavior occurs frequently in dyslexies. We studied the latencies and the metrical properties of saccades in the very rare case of an adult, nondyslexic express-saccade maker (male, age 29 years). The subject produced 65–95% express saccades in the gap (fixation point removed 200 ms before target onset) as well as in the overlap (fixation point not removed) paradigm, which qualified the subject as the most clear case of an express-saccade maker found so far. The number of express saccades increased rather than decreased when fixation foreperiod, gap duration, and target location were randomized from trial to trial as compared to when they remained constant. In the memory-guided saccade and in the antisaccade paradigms in which immediate saccade execution to a visual target had to be suppressed, the subject often reacted to the target with express saccades in an involuntary way. The amplitudes of express saccades were — in some conditions — found to progressively decrease with increasing latency, giving rise to amplitude transition functions. The present findings disprove the notion that express saccades are generated based on the prediction of the time and location of target appearance and support the notion that they are the result of an optomotor reflex. It is argued that the operation of the reflex is gated by a separate fixation system. Express-saccade makers are described as subjects with a dysfunction of the fixation system. Recent neurophysiological findings suggest that the subject studied in the present study has a selective dysfunction of the fixation system at the level of the superior colliculus.