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S Everling

Publications and source records attributed to S Everling.

17 recordsLinked to original sources
PubMed2003

Comparison of memory- and visually guided saccades using event-related fMRI.

Previous functional imaging studies have shown an increased hemodynamic signal in several cortical areas when subjects perform memory-guided saccades than that when they perform visually guided saccades using blocked trial designs. It is unknown, however, whether this difference results from sensory processes associated with stimulus presentation, from processes occurring during the delay period before saccade generation, or from an increased motor signal for memory-guided saccades. We conducted fMRI using an event-related paradigm that separated stimulus-related, delay-related, and saccade-related activity. Subjects initially fixated a central cross, whose color indicated whether the trial was a memory- or a visually guided trial. A peripheral stimulus was then flashed at one of 4 possible locations. On memory-guided trials, subjects had to remember this location for the subsequent saccade, whereas the stimulus was a distractor on visually guided trials. Fixation cross disappearance after a delay period was the signal either to generate a memory-guided saccade or to look at a visual stimulus that was flashed on visually guided trials. We found slightly greater stimulus-related activation for visually guided trials in 3 right prefrontal regions and right rostral intraparietal sulcus (IPS). Memory-guided trials evoked greater delay-related activity in right posterior inferior frontal gyrus, right medial frontal eye field, bilateral supplementary eye field, right rostral IPS, and right ventral IPS but not in middle frontal gyrus. Right precentral gyrus and right rostral IPS exhibited greater saccade-related activation on memory-guided trials. We conclude that activation differences revealed by previous blocked experiments have different sources in different areas and that cortical saccade regions exhibit delay-related activation differences.

Adult↗
PubMed2001

Prestimulus cortical potentials predict the performance in a saccadic distractor paradigm.

OBJECTIVE: In distractor paradigms, subjects sometimes respond to the wrong stimulus in a reflex-like manner. It is poorly understood why these errors occur. METHODS: To investigate the cortical processes possibly responsible for these errors, we recorded event-related potentials (ERPs) in human subjects performing a distractor saccade task in which subjects had to select a saccade target based on the colour of the initial fixation point and ignore a distractor on the opposite side. The initial fixation point disappeared 200 ms before the presentation of the visual stimuli (gap period). We compared the ERPs between correct trials in which subjects looked to the correct stimulus with error trials in which they looked towards the wrong stimulus. RESULTS: Correct response trials and error response trials showed a negative potential at the end of the gap period with the greatest amplitude over the fronto-central cortex. However, this potential had a lower amplitude in error response trials compared to correct response trials. CONCLUSION: These findings suggest that task-specific preparatory cortical processes in the frontal lobe prior to stimulus presentation have a major influence on the performance in distractor paradigms.

Adult↗
PubMed2000

Neuronal correlates for preparatory set associated with pro-saccades and anti-saccades in the primate frontal eye field.

Diversity in behavioral responses to sensory stimuli has been attributed to variations in preparatory set. Variability in oculomotor responses toward identical visual stimuli has been well documented, but the neuronal processes underlying this variability are poorly understood. Here, we report evidence for set-related activity for saccadic eye movements in single neurons in the frontal eye field (FEF) in monkeys trained on a task in which they either had to look toward a visual stimulus (pro-saccade) or away from the stimulus (anti-saccade) depending on a previous instruction. A portion of FEF neurons were identified as neurons projecting directly to the superior colliculus (SC) with antidromic activation techniques. Saccade-related neurons in the FEF had lower prestimulus and stimulus-related activity on anti-saccade trials compared with pro-saccade trials. The level of prestimulus activity correlated with saccadic reaction times, express saccade occurrence, and errors in the anti-saccade task. In addition, saccade-related activity in the FEF was higher for pro-saccades than for anti-saccades. These results demonstrate that the direct descending pathway from the FEF to the SC carries preparatory set-related activity for pro-saccades and anti-saccades. The results also provide insights into the neuronal basis of variations in saccadic reaction times and in the control of the prepotent response to glance to a flashed stimulus.

Action Potentials↗
PubMed2000

Influence of stimulus eccentricity and direction on characteristics of pro- and antisaccades in non-human primates.

The ability to inhibit reflexes in favor of goal-oriented behaviors is critical for optimal exploration and interaction with our environment. The antisaccade task can be used to investigate the ability of subjects to suppress a reflexive saccade (prosaccade) to a suddenly appearing visual stimulus and instead generate a voluntary saccade (antisaccade) to its mirror location. To understand the neural mechanisms required to perform this task, our lab has developed a non-human primate model. Two monkeys were trained on a task with randomly interleaved pro- and antisaccade trials, with the color of the central fixation point (FP) instructing the monkey to either make a prosaccade (red FP) or an antisaccade (green FP). In half of the trials, the FP disappeared 200 ms before stimulus presentation (gap condition) and in the remaining trials, the FP remained visible (overlap condition) during stimulus presentation. The effect of stimulus eccentricity and direction was examined by presenting the stimulus at one of eight different radial directions (0-360 degrees ) and five eccentricities (2, 4, 8, 10, and 16 degrees ). Antisaccades had longer saccadic reaction times (SRTs), more dysmetria, and lower peak velocities than prosaccades. Direction errors in the antisaccade task were more prevalent in the gap condition. The difference in mean SRT between correct pro- and antisaccades, the anti-effect, was greater in the overlap condition. The difference in mean SRT between the overlap and the gap condition, the gap effect, was larger for antisaccades than for prosaccades. The manipulation of stimulus eccentricity and direction influenced SRT and the proportion of direction errors. These results are comparable to human studies, supporting the use of this animal model for investigating the neural mechanisms subserving the generation of antisaccades.

Animals↗
PubMed2000

On your mark, get set: brainstem circuitry underlying saccadic initiation.

Saccades are rapid eye movements that are used to move the visual axis toward targets of interest in the visual field. The time to initiate a saccade is dependent upon many factors. Here we review some of the recent advances in our understanding of the these processes in primates. Neurons in the superior colliculus and brainstem reticular formation are organised into a network to control saccades. Some neurons are active during visual fixation, while others are active during the preparation and execution of saccades. Several factors can influence the excitability levels of these neurons prior to the appearance of a new saccadic target. These pre-target changes in excitability are correlated to subsequent changes in behavioural performance. Our results show how neuronal signals in the superior colliculus and brainstem reticular formation can be shaped by contextual factors and demonstrate how situational experience can expedite motor behaviour via the advanced preparation of motor programs.

Animals↗
PubMed1999

Role of primate superior colliculus in preparation and execution of anti-saccades and pro-saccades.

We investigated how the brain switches between the preparation of a movement where a stimulus is the target of the movement, and a movement where a stimulus serves as a landmark for an instructed movement elsewhere. Monkeys were trained on a pro-/anti-saccade paradigm in which they either had to generate a pro-saccade toward a visual stimulus or an anti-saccade away from the stimulus to its mirror position, depending on the color of an initial fixation point. Neural activity was recorded in the superior colliculus (SC), a structure that is known to be involved in the generation of fast saccades, to determine whether it was also involved in the generation of anti-saccades. On anti-saccade trials, fixation during the instruction period was associated with an increased activity of collicular fixation-related neurons and a decreased activity of saccade-related neurons. Stimulus-related and saccade-related activity was reduced on anti-saccade trials. Our results demonstrate that the anti-saccade task involves (and may require) the attenuation of preparatory and stimulus-related activity in the SC to avoid unwanted pro-saccades. Because the attenuated pre-saccade activity that we found in the SC may be insufficient by itself to elicit correct anti-saccades, additional movement signals from other brain areas are presumably required.

Animals↗
PubMed1999

Event-related potentials and saccadic reaction times: effects of fixation point offset or change.

Previous studies have shown that saccadic reaction times (SRTs) are reduced if the initial fixation point (FP) disappears 200 ms (gap period) before a peripheral target is presented. This gap saccade task is associated with a negative cortical potential at the end of the gap period. To determine whether the neural processes underlying this potential account for the reduction of SRTs during gap saccade tasks, we recorded event-related potentials (ERPs) in 19 subjects performing a gap saccade task (gap duration 200 ms), a warning saccade task (the color of the FP changed 200 ms prior to target appearance) and an overlap task (the FP remained visible during the trial). SRTs were shortest during the gap task, longest during the overlap task and intermediate during the warning task. The gap and warning tasks were accompanied by the same widespread negative cortical potential with a maximum at the time of stimulus presentation. These findings indicate that the warning effect mediated by the disappearance of the FP during gap saccade tasks is responsible for the gap negativity which was observed by several authors. Our findings of shorter SRTs during the gap task than the warning task, however, suggest that the gap has an additional effect that probably depends on subcortical mechanisms.

Adult↗
PubMed1998

Event-related potentials associated with correct and incorrect responses in a cued antisaccade task.

In an antisaccade task, subjects are instructed to inhibit a reflexive saccade towards a peripheral stimulus flash and to generate a saccade in the opposite direction. It has been shown recently that normal subjects will generate a high number of incorrect prosaccades in an antisaccade task if the fixation point is extinguished 200 ms before the stimulus appears and if a valid cue for the subsequent antisaccade is given during this gap period. In the present study we recorded cerebral event-related potentials from 19 scalp electrodes from normal subjects prior to correct and incorrect responses in a cued antisaccade task to investigate the neural processes associated with correct antisaccades and incorrect prosaccades in this task. Correct antisaccades and incorrect prosaccades were associated with a negative potential with a maximal amplitude around stimulus onset over the dorsomedial frontal cortex. This potential was higher prior to correct antisaccades than prior to incorrect prosaccades. The execution of a correct antisaccade was preceded by a shift of a negative potential from the parietal hemisphere contralateral to the visual stimulus towards the parietal hemisphere ipsilateral to the stimulus. These results support the view that the supplementary eye fields participate in the inhibition of incorrect saccades in a cued antisaccade task and show that the parietal cortex participates in generating a neural representation of the visual stimulus in the hemifield ipsilateral to the stimulus before generating a motor response.

Adult↗
PubMed1998

The antisaccade: a review of basic research and clinical studies.

The ability to suppress reflexive responses in favor of voluntary motor acts is crucial for everyday life. Both abilities can be tested with an oculomotor task, the antisaccade task. This task requires subjects to suppress a reflexive prosaccade to a flashed visual stimulus and instead to generate a voluntary saccade to the opposite side. This article reviews what is currently known about the neural structures and processes which are involved in the performance of this task. Current data show that a variety of brain lesions, neurological diseases and psychiatric disorders result in errors, i.e. prosaccades towards the stimulus, in this task. Brain imaging studies have shown that a widely distributed cortical and subcortical network is active during the generation of antisaccades. These findings are discussed and the potential of the antisaccade task for diagnostic purposes is evaluated.

Animals↗
PubMed1998

Accuracy of visually and memory-guided antisaccades in man.

Primary saccades to remembered targets are generally not precise, but rather undershoot target position. The major source of this saccadic undershoot may be (a) a memory-related process or (b) a poor spatial resolution in those processes which transfer the retinotopic target information into an intermediate memory-linked representation of space. The aim of this study was to investigate whether distortions of eye positions in the antisaccade task, which are characterized by inherent co-ordinate transformation processes, may completely account for the spatial inaccuracies of memory-guided antisaccades. The results show that the spatial inaccuracy of primary and secondary eye movements in the visually guided antisaccade task was comparable to that in the memory-guided antisaccade task. In both conditions, the direction error component was less dysmetric than the amplitude error component. Secondary eye movements were significantly corrective. This increase of eye position accuracy was achieved by reducing the amplitude error only. It is concluded from this study that at least some of the distortion of memory-guided saccades is due to inaccuracies in the sensorimotor co-ordinate transformations.

Adult↗
PubMed1998

Comparison of the discharge characteristics of brain stem omnipause neurons and superior colliculus fixation neurons in monkey: implications for control of fixation and saccade behavior.

Fixation neurons (SCFNs) in the rostral pole of the superior colliculus (SC) and omnipause neurons (OPNs) in the nucleus raphe interpositus (rip) in the pons share similar discharge properties. Both types of neurons discharge tonically during periods of visual fixation and pause for saccadic eye movements, and their activation by electrical stimulation suppresses saccade generation. On the basis of these similarities and the projection from the rostral SC to the rip, it was hypothesized that SCFNs provide a major excitatory input to OPNs. We investigated the role and relationship of SCFNs and OPNs with respect to both fixation behavior and saccade generation by comparing their activity recorded in the same monkeys performing a gap saccade task. In this task, the central fixation point was extinguished 200 ms before the presentation of an eccentric saccadic target, and the discharges of OPNs and SCFNs were contrasted during visual fixation, nonvisual (gap) fixation, and saccade generation. During visual fixation, the mean discharge rate of OPNs was higher and more regular than that of SCFNs. During the gap period, SCFNs decreased their discharge rate before target appearance, whereas no change in discharge rate was observed in OPNs. For both SCFNs and OPNs, the activity level before target appearance was not correlated to saccadic reaction time. In contrast to SCFNs, several OPNs responded with a transient phasic increase in discharge immediately after the target presentation. Before their saccade-related pause, there was a gradual reduction in the activity of SCFNs, whereas OPNs had an abrupt cessation of discharge. SCFNs paused earlier than OPNs, but the OPN pause onset was better synchronized to saccade onset than the SCFN pause onset. OPNs resumed firing after their pause in activity earlier than SCFNs, and the OPN pause end was better synchronized to saccade end than the SCFN pause end. These physiological data reveal differences in the discharge properties of SCFNs and OPNs that are irreconcilable with the hypothesis that the discharge pattern of OPNs reflects simply the excitatory input from SCFNs. It is most likely that additional inputs to OPNs compensate for the reduction in discharge of SCFNs during these periods.

Animals↗
PubMed1998

Reflex suppression in the anti-saccade task is dependent on prestimulus neural processes.

Reflexive responses often must be suppressed to correctly execute a voluntary behavior. It is largely unknown why this control sometimes fails. To examine the neural processes responsible for these failures, we recorded single-neuron activity in the superior colliculus (SC) in behaving monkeys during an anti-saccade task in which they had to suppress a saccade to a visual stimulus that suddenly appeared in the periphery and generate a saccade to the opposite side. We found that the level and distribution of prestimulus activity of buildup neurons in the SC was highly predictive of whether a correct response or an error occurred. A high level of prestimulus activity in buildup neurons at the location in the SC where the visual stimulus was represented was associated with the generation of a reflexive saccade to the stimulus. These findings suggest that the successful suppression of reflexive saccades is dependent on prestimulus neural processes in the SC.

Animals↗
PubMed1997

Influence of pre-target cortical potentials on saccadic reaction times.

Saccades elicited by suddenly appearing targets show a broad distribution of reaction times. This may depend on variations in the subject's state of preparation before target onset. To test this hypothesis, we recorded scalp event-related potentials from eight human subjects to investigate whether differences in saccadic reaction times (SRTs) are related to differences in cortical slow potentials prior to target onset. Compared with trials with medium SRTs (180-230 ms), trials with fast SRTs (130-180 ms) were found to be preceded by a more negative slow potential and trials with slow SRTs (230-280 ms) were found to be preceded by a more positive slow potential. These results support the hypothesis that cortical activation prior to target appearance influences SRTs.

Adult↗
PubMed1997

Cortical potentials preceding pro- and antisaccades in man.

The antisaccade task has been used widely to assess a frontal lobe deficit. In the present study cortical potentials preceding prosaccades and antisaccades were recorded in 7 healthy subjects with 19 scalp electrodes according to the international 10-20 system. The main results were as follows: (1) In both saccade types, a slow presaccadic negative shift was observed at dorso-medial frontal recording sites. The integral over these potentials was significantly greater for antisaccades at the C3, C4 and Cz location than in prosaccades. (2) The integral over presaccadic positivity was significantly lower for antisaccades compared with prosaccades at the Cz location. It is concluded that these results support an important role of the supplementary eye fields in generating antisaccades.

Adult↗
PubMed1996

Hypometric primary saccades of schizophrenics in a delayed-response task.

In this study, the execution of delayed saccades in 15 DSM-III-R-schizophrenic patients and 15 normal subjects was investigated. While looking at a central fixation cross, a peripheral target was randomly presented at 10 degrees eccentricity. Subjects were instructed to saccade to the target when the fixation cross was switched off after 500 ms. Two experiments were conducted: (a) a delayed-saccade task and, (b) a memory-guided saccade task, that is, the peripheral target was switched off together with the fixation cross. In the delayed-saccade task, amplitudes of regular saccades did not differ between schizophrenic patients and normals. In the memory-guided saccade task, schizophrenic subjects showed marked hypometric saccades. Incorrect delayed saccades (while the fixation cross was on) were also hypometric in schizophrenics, but not in normal controls. The final eye position, i.e., the position reached after the execution of correction saccades, however, did not differ between patients and controls. This means that schizophrenics show a deficit in the programming of primary saccades, if the fixation point and the peripheral target are (a) both visually presented or (b) both memorized. The results support the hypothesis that these saccades are the result of an averaging effect between the fixation point and the peripheral target. It is further hypothesized that these deficits might be explained by a lack of prefrontal inhibition of ocular fixation areas.

Adolescent↗
PubMed1996

Cortical potentials during the gap prior to express saccades and fast regular saccades.

When a temporal gap is introduced between the offset of the central fixation point and the appearance of a new target, saccadic reaction time is reduced (gap effect) and a special population of extremely fast saccades occurs (express saccades). It has been hypothesized that the gap triggers a readiness signal, which is responsible for the reduced saccadic reaction times. Here, we recorded event-related potentials during the gap to investigate the central processes associated with the generation of fast regular saccades and express saccades. Prior to the execution of fast regular saccades, subjects produced a slow negative shift, with a maximum at frontal and central channels that started 40 ms after fixation offset. This widespread negativity is similar to a readiness potential. Anticipatory saccades were preceded by an increased frontal and parietal negativity. Prior to express saccades, a frontal negativity was observed, which started 135 ms after the disappearance of the fixation point. It is assumed that the frontal negativity prior to express saccades corresponds to the fixation-disengagement discharge described in the frontal eye field of monkeys. Therefore, we hypothesize that fast regular saccades are the result of an increased readiness signal, while express saccades are the result of specific preparatory processes.

Adult↗