Overlap Paradigm Research Articles

The visual strategy specific to humans among hominids: A study using the gap–overlap paradigm

Although an extensive body of literature exists on the cognitive underpinnings of gaze movements in macaques and humans, few studies have investigated this topic from a broader evolutionary perspective. This study used the gap–overlap paradigm to examine the timing of the gaze movements by four hominid species: humans, chimpanzees, gorillas, and orangutans. The saccade latency involved in shifting the gaze from central to peripheral stimuli was measured and compared under two conditions, gap and overlap. The central stimulus disappeared shortly before the onset of the peripheral stimulus under the gap condition, but it remained under the overlap condition. Although all species demonstrated similar saccade latencies under the gap condition, the species clearly differed from one another under the overlap condition, which may suggest their similar perceptual and motor mechanism of making a saccade on the one hand and their differential strategies for coping with the competition between two activities involving fixation and initiation of a saccade (i.e. central vs. peripheral visual stimuli) on the other hand. In particular, humans showed longer saccade latency under the overlap condition compared to the other great apes, which may reflect this species’ unique means of visual processing.

Saccade behaviour in migraine patients

Voxel-based morphometry studies in migraine patients showed significant grey matter volume reduction in regions involved in the control of saccadic eye movements. We hypothesized that these changes would be reflected in dysfunctional saccadic behaviour. Saccades were recorded by infrared oculography using three different paradigms (pro-saccade with gap, pro-saccade overlap and anti-saccade with gap). We compared the results for migraine patients (n = 80) with those for controls (n = 87). No significant differences were found between migraine patients with (n = 46) and without (n = 34) aura. Migraine patients showed a saccadic behaviour that differed from controls in three respects. In migraine patients, the latencies in the pro-saccade with gap paradigm were borderline significantly longer. Moreover, in both the pro-saccade with gap and the pro-saccade overlap paradigm we observed a larger intra-individual variation of the latency in migraine patients. However, the biggest difference was that the patients who received migraine prophylactic therapy made significantly more anti-saccade errors in the anti-saccade with gap paradigm, suggesting that inhibitory saccade control is impaired in migraine patients depending on the severity of the migraine. We suggest a deficient inhibitory control, reflecting an executive dysfunction in the dorsolateral prefrontal cortex or a dysfunction in the cingulate cortex, is present in migraine patients.

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.

The pupil reflects motor preparation for saccades – even before the eye starts to move

The eye produces saccadic eye movements whose reaction times are perhaps the shortest in humans. Saccade latencies reflect ongoing cortical processing and, generally, shorter latencies are supposed to reflect advanced motor preparation. The dilation of the eye’s pupil is reported to reflect cortical processing as well. Eight participants made saccades in a gap and overlap paradigm (in pure and mixed blocks), which we used in order to produce a variety of different saccade latencies. Saccades and pupil size were measured with the EyeLink II. The pattern in pupil dilation resembled that of a gap effect: for gap blocks, pupil dilations were larger compared to overlap blocks; mixing gap and overlap trials reduced the pupil dilation for gap trials thereby inducing a switching cost. Furthermore, saccade latencies across all tasks predicted the magnitude of pupil dilations post hoc: the longer the saccade latency the smaller the pupil dilation before the eye actually began to move. In accordance with observations for manual responses, we conclude that pupil dilations prior to saccade execution reflect advanced motor preparations and therefore provide valid indicator qualities for ongoing cortical processes.

Ocular Motor Fixation Deficits in Friedreich Ataxia

Friedreich ataxia (FRDA) is the most common genetic cause of ataxia with a prevalence of approximately 1 in 29,000. Ocular motor abnormalities are common in FRDA and include fixation instability, saccadic dysmetria, and vestibular dysfunction. It has not yet been determined whether aspects of spatial attention, which are closely coupled to eye movements, are similarly compromised in FRDA. This study examined attentional engagement and disengagement of eye movements in FRDA using a gap overlap task. Thirteen individuals with genetically confirmed FRDA and 12 age-matched unaffected controls participated in the experiment. The gap overlap paradigm was used to examine the effect of early (gap condition), simultaneous (null condition), or late (overlap condition) removal of a central fixation on saccadic latency to a peripheral target stimulus. Although the FRDA group showed a larger gap effect (i.e., difference in saccadic latencies between the overlap and gap condition), these participants demonstrated a greater difference in latencies in the overlap relative to the null condition, suggestive of deficits within the disengagement process of attentional orienting. We propose a role for the cerebellum in these deficits in the disengagement of spatial attention based on evidence of cerebellar connectivity with regions involved in exogenous shifts of attention. The significant correlations between saccadic latency and disease severity as measured by the Friedreich Ataxia Rating Scale further support the proposal that saccadic latency might be useful as a surrogate marker of disease severity and progression in future clinical trials in FRDA.

Improving the saccade peak velocity measurement for detecting fatigue

The aim of the study was to compare saccadic peak velocity (SPV) values measured with video based Fitness Impairment Tester (FIT) and electro-oculography (EOG) during prolonged wakefulness. We tested different numbers of saccades and two saccade paradigms to improve the EOG measurements for detecting fatigue. The SPVs were measured from 11 fast patrol boat navigators with FIT and EOG every sixth hour until 54 h. Subjective sleepiness was assessed with the Karolinska Sleepiness Scale. EOG was measured using an overlap and a gap paradigm and the data was divided into sequential five 20-saccade blocks and cumulative blocks of 20, 40, 60, 80, and 100 saccades. Compared to the gap paradigm, the overlap paradigm produced a higher number of analyzable saccades for a given measurement time. The shorter measurements (20–40 saccades) appeared to be more sensitive for fatigue, whereas the longer measurements (60–100 saccades) were more sensitive to time spent on the task. Thus, the optimal number of saccades varies also depending on the research question. The EOG method was more sensitive to fatigue than FIT. The FIT values measured after 30 and 36 h of wakefulness did not differ significantly from the baseline values, while subjective sleepiness and the EOG values showed that the participants were significantly less alert at these time points. The EOG measurements can be improved for detecting fatigue by using the overlap saccade paradigm. The SPV values measured with the EOG method appear to be somewhat more sensitive in detecting fatigue than the FIT method.

Effects of aging on regular and express latencies of vergence

Vergence eye movements are frequent in every day life and important for depth perception. Yet, studies of vergence in elderly are rare. We examined convergence and divergence between targets placed along median line at 20, 40 or 150cm. Thirteen elderly (70±11years) and ten young (25±3years) adults participated in the study. The gap paradigm (i.e., the fixed stimulus is extinguished prior to target onset) and the overlap paradigm (the fixed stimulus remains illuminated after target onset) were used to elicit reflexive or voluntary eye movements. Latency of convergence and divergence increased with age in both gap and overlap conditions. Both young subjects and elderly showed shorter latency of vergence under the gap condition than under the overlap condition. In the overlap condition, for elderly only, convergence resulted in longer latencies than divergence. In the gap condition express convergence was rare, while express latencies (80–120ms) were frequent for divergence starting from a near fixation point (at 20cm). The rates of express divergence were similar for young (23%) and elderly (21%). These results were in line with prior studies of saccades and indicated different aging effects on regular and express latencies.

Rare express saccades in elderly fallers

To examine horizontal saccades in elderly subjects with falling history; prior extensive screening was done to recruit subjects with falling history in the absence of pathology. Twelve elderly with falling history were tested. Two testing conditions were used: the gap (fixation target extinguishes prior to target onset) and the overlap (fixation stays on after target onset) paradigms. Each condition was run at three viewing distances--20 cm, 40 cm, and 150 cm, corresponding to convergence angle at 17.1 degrees, 8.6 degrees, and 2.3 degrees, respectively. Eye movements were recorded with the photoelectric IRIS (Skalar medical). (i) like in healthy elderly subjects, elderly with falling history produce shorter latencies in the gap paradigm than in the overlap paradigm; (ii) their latencies are shorter at near distances (20 and 40 cm) relative to 150 cm for both paradigms; (iii) the novel result is that they fail to produce express latencies even in the conditions (near viewing distance and the gap task) known to promote high rates of express in adults (25%) or in healthy elderly (20%). Seven from the 10 healthy elderly produced express saccades at rates >12%, while 9 of the 12 older subjects with falling history showed no express saccades at all; the remaining 3 subjects showed low rates <12%. The quasi paucity of express saccades could be due to the disequilibrium of complex cortical/subcortical networks needed for making express saccades. The results support models suggesting specific network for express saccades; missing of such optomotor reflex may go along with missing other reflexes as well increasing the chances of falling.

Repetitive TMS over the human oculomotor cortex: Comparison of 1-Hz and theta burst stimulation

The aim of the study was to compare the effect duration of two different protocols of repetitive transcranial magnetic stimulation (rTMS) on saccade triggering. In four experiments, two regions (right frontal eye field (FEF) and vertex) were stimulated using a 1-Hz and a theta burst protocol (three 30 Hz pulses repeated at intervals of 100 ms). The same number of TMS pulses (600 pulses) was applied with stimulation strength of 80% of the resting motor threshold for hand muscles. Following stimulation the subjects repeatedly performed an oculomotor task using a modified overlap paradigm, and saccade latencies were measured over a period of 60 min. The results show that both 1-Hz and theta burst stimulation had inhibitory effects on saccade triggering when applied over the FEF, but not over the vertex. One-hertz rTMS significantly increased saccade latencies over a period of about 8 min. After theta burst rTMS, this effect lasted up to 30 min. Furthermore, the decay of rTMS effects was protocol-specific: After 1-Hz stimulation, saccade latencies returned to a baseline level much faster than after theta burst stimulation. We speculate that these time course differences represent distinct physiological mechanisms of how TMS interacts with brain function.

Eye movement signals influence perception: Evidence from the adaptation of reactive and volitional saccades

Information about upcoming saccadic eye movements is used to orient visuo-spatial attention across the visual field. Different eye movement signals (intended or actual) could be used according to the intentionality of the saccade in preparation (Reactive or Volitional), and can be dissociated by saccadic adaptation. Gap 0 and overlap paradigms were contrasted to elicit the two saccade populations with different latencies and an asymmetric transfer of saccadic adaptation. Preparation of both saccade types caused a concomitant shift in the attentional focus (indexed by relative perceptual performance) to the actual, not intended, eye position. The attentional shift emerged progressively, earlier for V-saccades but reaching a maximal level around saccade onset for both saccade types. These results suggest that information about actual eye movements mediates the pre-saccadic shift of attention.

Effects of uncertainty and target displacement on the latency of express saccades in man

Saccadic eye movements generated in response to a gap paradigm in which the fixation light spot was extinguished 200 ms prior to presentation of the target light spot showed appreciably shorter latencies than for the overlap paradigm in which the target light spot was presented 200 ms prior to extinction of the fixation light spot. When there was unpredictability in the direction of target presentation, i.e., to the left or right of the fixation light spot, the gap paradigm evoked mainly fast regular saccades of peak latency of 155 ms with relatively few express saccades which were defined as having latencies of less than 120 ms. By contrast, when the target always appeared to the right, a substantial population of express saccades with peak latency 95 ms was now generated. There was also a change in the relationship between saccadic latency and target angular displacement which covered the range 5–35 °. With the overlap paradigm and unpredictability of target direction, the latencies of the slow regular saccades increased markedly with target angular displacement. This was not the case with the same target direction when the latency of slow regular, fast regular, and express saccades remained constant with increasing target angular displacement. This indicates for targets appearing in the same hemifield that the ocular motor system operates with shortest latency irrespective of target angular displacement.

One-Hertz transcranial magnetic stimulation over the frontal eye field induces lasting inhibition of saccade triggering

The aim of the study was to examine the effect of low-frequency repetitive transcranial magnetic stimulation on saccade triggering. In five participants, a train of 600 pulses with a frequency of 1 Hz was applied over the right frontal eye field and--as control condition--over the vertex. After repetitive transcranial magnetic stimulation application, oculomotor performance was evaluated with an overlap paradigm. The results show that the repetitive transcranial magnetic stimulation effect was specific for frontal eye field stimulation. Saccade latencies were found to be increased bilaterally for several minutes after the stimulation, and the time course of recovery was different for the ipsilateral and contralateral sides. The results are discussed in the light of possible local and remote repetitive transcranial magnetic stimulation effects on the oculomotor network.

Total sleep deprivation effect on disengagement of spatial attention as assessed by saccadic eye movements

Objective Previous research has shown that total sleep deprivation (TSD) of short duration (one night) affects performance to some cognitive tasks subserved by a fronto-parietal network. The aim of our study was to assess the effects of TSD on visuo-spatial attention, which is a cognitive task involving this network. Specifically, the disengagement of spatial attention was investigated with gap and overlap paradigms of saccadic eye movements. Methods Ten healthy young male subjects performed the two tasks the morning after a normal night and after a TSD night. The study was conducted using a balanced, crossover design. Results TSD significantly increased the gap effect (difference of latency between overlap and gap). Conclusions This result can be interpreted as an impaired disengagement of attention after TSD. As the peak velocity, which is an indicator of alertness, was not altered by TSD, the impairment in the disengagement of spatial attention does not result from a decrease in alertness. Significance This study shows that saccadic eye movements enable studying alertness and disengagement of spatial attention simultaneously. The idea that specific brain areas are affected by TSD is confirmed by our results.

Latency of Prosaccades and Antisaccades in Professional Shooters

This study evaluated hypothesis that the faster saccadic reaction time in professional clay-target shooters found in a previous study was because of a superiority of athletes arising at the attention level or at level of saccadic motor preparation. Ten shooters with at least 6 yr of shooting training in Olympic shotgun disciplines and 10 control subjects participated in the experiments. In the first experiment, prosaccades were studied by comparing the saccadic latencies obtained from the overlap and gap paradigms. In the overlap paradigm, a target was presented randomly at one of four cardinal positions with the fixation point presented throughout the trial duration. In the gap paradigm, the fixation point was removed at the time of target presentation. In the second experiment, subjects were instructed to saccade as quickly as possible in the direction opposite to that of the target location (antisaccades). Shooters had shorter saccadic latency than controls, both with gap and overlap conditions in the first experiment and in the antisaccade condition of the second experiment. This result indicates that athletes' advantage in saccadic reaction times cannot be attributed to improvement of the attentional mechanism of disengagement. Present results support the hypothesis that shooters develop shorter motor preparation to saccades.

Visual processing and infant ocular latencies in the overlap paradigm.

Young infants have repeatedly been shown to be slower than older infants to shift fixation from a midline stimulus to a peripheral stimulus. This is generally thought to reflect maturation of the neural substrates that mediate the disengagement of attention, but this developmental difference may also be attributable to young infants' slower processing of the midline stimulus. This possibility was tested with 3- and 7-month-old infants in 2 experiments in which the degree of familiarity of the midline stimulus was manipulated across repeated trials. The results of these experiments demonstrated that the processing of midline content does affect infants' ocular latencies to a peripheral stimulus but that developmental differences in such processing do not account for developmental differences in disengagement seen across the 1st year.

Prolongation of latency of horizontal saccades in elderly is distance and task specific

The present study examined horizontal saccades in healthy subjects: 9 adults (20–32 years) and 10 aged subjects (63–83 years), under gap (fixation target extinguishes prior to target onset) and overlap (fixation stays on after target onset). The gap paradigm is known to promote fast initiation of saccades while the overlap paradigm promotes voluntary saccades with longer latency. In real life we perform saccades at various distances. In this study each paradigm was run at three viewing distances—20, 40 and 150 cm, corresponding to a convergence angle of 17.1°, 8.6° and 2.3°, respectively. Eye movements were recorded with the Chronos video eye tracker or with the photoelectric IRIS. The main findings are: (i) increase in latency of saccades with age, with distance and with the overlap condition; (ii) evidence for interaction between these factors, indicating the following anomaly: in the gap condition and at near, aged subjects show short latencies similar to those of young adults; (iii) express type of latencies (between 80 and 120 ms) occur most frequently at near in the gap condition and at similar rates in young (25%) and aged subjects (20%). The specificity of close distance combined with the gap for triggering short latency saccades could be related to both attention and oculomotor fixation disengagement. The strength of coupling between fixation—eye movement control and visual attention control varies for different locations in space, and its decline with aging can be also different.

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.

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.

Inhibition of return, gap effect and saccadic reaction time to a visual target.

Simple manual reaction time (MRT) to a visual target (S2) is shortened when a non-informative cue (S1) is flashed at the S2 location shortly before the onset of S2 (early facilitation). Afterwards, MRT to S2 appearing at the S1 location is lengthened (inhibition of return - IOR). Similar results have been obtained for saccadic reaction time (SRT). Moreover, when there is a temporal gap between offset of the fixation point (FP) and onset of a target (gap paradigm), SRT is shorter than SRT in an overlap paradigm (FP remains on). In the present study, we determined SRT to S2 (10 degrees) after presenting S1 at the same eccentricity (10 degrees) or at a parafoveal position (2 degrees) in the same or in the opposite hemifield. In addition, we employed both gap and overlap paradigms. Twelve subjects were asked not to respond to S1 (2 degrees or 10 degrees) to the right or to the left of FP, but to respond by making a saccadic movement in response to S2. We obtained the following results: 1) a 40-ms gap effect, 2) an interaction between gap effect and IOR, 3) a 39-ms delay (IOR) when S2 appeared at the cued (S1) position, and 4) a smaller (17 ms) but significant inhibition when S1 occurred at 2 degrees in the ipsilateral hemifield. Thus, a parafoveal (2 degrees) S1 elicits an inhibition of SRT towards ipsilateral peripheral targets. Since an inhibition of the ipsilateral hemifield by a 1 degree eccentric cue has been reported to occur when manual responses are employed, we suggest that the postulated functional link between covert and overt orienting of attention is also valid for parafoveal cues.

Idiosyncratic left–right asymmetries of saccadic latencies: examination in a gap paradigm

Subjects were given reflexive- and voluntary-saccade tasks using five different gap intervals (0–500 ms) between the fixation point offset and the target onset and an overlap paradigm (i.e., the fixation point remained on during the target presentation). In the first experiment, targets were monocularly presented, and the latencies of reflexive saccades to a peripheral target were compared between the left and right visual fields in which targets were presented. The data averaged over subjects did not show a significant difference of saccade latencies between the two visual fields. However, individual subjects showed strong left/right asymmetries of saccade latencies: six out of the 12 subjects that participated made a saccade more rapidly to the right than to the left, and two other subjects showed the reverse result. In these cases, the left/right asymmetry was observed on both gap and overlap trials. The saccade latencies were not affected by the hemiretina to which a target was projected. The second experiment was conducted to identify conditions under which the left/right asymmetry can be reproduced. For this purpose, five subjects were given both reflexive-saccade and voluntary-saccade tasks. In the latter task, a cue stimulus for generating saccades was given at the central fixation point. Regardless of whether saccades were made reflexively or voluntarily, and whether the targets were viewed monocularly or binocularly, each subject showed the same pattern of left/right asymmetries of saccade latencies. The results were interpreted as showing that a visuo-spatial attentional bias specific to individual subjects is involved in generating idiosyncratic left/right asymmetries of saccade latencies.