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Stefan Everling

Publications and source records attributed to Stefan Everling.

17 recordsLinked to original sources
PubMed2006

Monkey dorsolateral prefrontal cortex sends task-selective signals directly to the superior colliculus.

The dorsolateral prefrontal cortex (DLPFC) has been implicated in the ability to perform complex behaviors requiring the implementation of cognitive control. A central supposition of models of prefrontal function is that the DLPFC engages control by selectively modulating the activity of target structures to which it is connected, but no studies in the primate have directly investigated DLPFC output signals. Here, we recorded the activity of DLPFC neurons identified as sending a direct projection to the superior colliculus, a midbrain oculomotor structure, while monkeys performed alternating blocks of trials in which they had to look toward a flashed peripheral stimulus (prosaccades) and trials in which they had to look away from the stimulus in the opposite direction (antisaccades). We report the first direct evidence that the primate DLPFC sends task-selective signals to a target structure. This supports the notion that the DLPFC orchestrates the activity of other brain areas in accordance with task requirements.

Action Potentials↗
PubMed2006

Inhibition and generation of saccades: rapid event-related fMRI of prosaccades, antisaccades, and nogo trials.

Flexible, adaptive behavior often requires the inhibition of automatic responses in favor of voluntary response generation. The antisaccade task requires active inhibition of the automatic saccade to a peripheral stimulus followed by generation of a voluntary antisaccade to the opposite location. Previous studies demonstrated greater functional magnetic resonance imaging (fMRI) activation for antisaccades than prosaccades in cortical saccade areas but did not distinguish the relative contributions of saccadic inhibition and generation. To address this question, we compared prosaccades, antisaccades, and nogo trials in a rapid event-related fMRI experiment with ten human subjects (6 female, 4 male). Trials were compound, containing a colored fixation point to cue trial type, followed by a 200-ms gap and then peripheral stimulus presentation and response. Required responses for prosaccade, antisaccade, and nogo trials, respectively, were to generate a saccade to the stimulus, look away from it, and inhibit the automatic saccade while maintaining central fixation. Frontal and supplementary eye fields, anterior cingulate cortex, intraparietal sulcus, and precuneus, exhibited surprisingly similar activation patterns for prosaccade and nogo responses, suggesting that BOLD signal in cortical saccade regions might predominantly reflect visual detection and attention processes rather than saccade generation or inhibition. These regions displayed greater activation for antisaccade responses versus prosaccade or nogo responses, possibly due to visuospatial remapping or increased attention levels in the antisaccade task. In right superior frontal sulcus, right supramarginal gyrus, and posterior cingulate sulcus, activation was greater for nogo compared to prosaccade responses, suggesting a role in active saccadic inhibition.

Adult↗
PubMed2006

Selective representation of task-relevant objects and locations in the monkey prefrontal cortex.

In the monkey prefrontal cortex (PFC), task context exerts a strong influence on neural activity. We examined different aspects of task context in a temporal search task. On each trial, the monkey (Macaca mulatta) watched a stream of pictures presented to left or right of fixation. The task was to hold fixation until seeing a particular target, and then to make an immediate saccade to it. Sometimes (unilateral task), the attended pictures appeared alone, with a cue at trial onset indicating whether they would be presented to left or right. Sometimes (bilateral task), the attended picture stream (cued side) was accompanied by an irrelevant stream on the opposite side. In two macaques, we recorded responses from a total of 161 cells in the lateral PFC. Many cells (75/161) showed visual responses. Object-selective responses were strongly shaped by task relevance - with stronger responses to targets than to nontargets, failure to discriminate one nontarget from another, and filtering out of information from an irrelevant stimulus stream. Location selectivity occurred rather independently of object selectivity, and independently in visual responses and delay periods between one stimulus and the next. On error trials, PFC activity followed the correct rules of the task, rather than the incorrect overt behaviour. Together, these results suggest a highly programmable system, with responses strongly determined by the rules and requirements of the task performed.

Action Potentials↗
PubMed2006

Neural activity in monkey prefrontal cortex is modulated by task context and behavioral instruction during delayed-match-to-sample and conditional prosaccade-antisaccade tasks.

Complex behavior often requires the formation of associations between environmental stimuli and motor responses appropriate to those stimuli. Moreover, the appropriate response to a given stimulus may vary depending on environmental context. Stimulus-response associations that are adaptive in one situation may not be in another. The prefrontal cortex (PFC) has been shown to be critical for stimulus-response mapping and the implementation of task context. To investigate the neural representation of sensory-motor associations and task context in the PFC, we recorded the activity of prefrontal neurons in two monkeys while they performed two tasks. The first task was a delayed-match-to-sample task in which monkeys were presented with a sample picture and rewarded for making a saccade to the test picture that matched the sample picture following a delay period. The second task was a conditional visuomotor task in which identical sample pictures were presented. In this task, animals were rewarded for performing either prosaccades or antisaccades following the delay period depending on sample picture identity. PFC neurons showed task selectivity, object selectivity, and combinations of task and object selectivity. These modulations of activity took the form of a reduction in stimulus and delay-related activity, and a pro/anti instruction-based grouping of delay activity in the conditional visuomotor task. These data show that activity in PFC neurons is modulated by experimental context, and that this activity represents the formal demands of the task currently being performed.

Action Potentials↗
PubMed2005

Task-dependent effects of social attention on saccadic reaction times.

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

Adult↗
PubMed2005

Neural processes associated with antisaccade task performance investigated with event-related FMRI.

One of the hallmarks of cognitive control is the suppression of prepotent but inappropriate responses. Here we used event-related functional MRI to measure functional brain activation during a stimulus-response incompatibility task. Subjects were instructed before a stimulus appeared either to look at the stimulus (prosaccade) or to look away from the stimulus (antisaccade). Eye movements were recorded so that functional brain activation could be grouped into prosaccades, correct antisaccades, and errors (saccades toward the stimulus on antisaccade trials). Correct antisaccade trials were associated with significantly more activation in frontal and parietal cortical areas compared with prosaccade trials during the late preparatory period before stimulus appearance. Correct antisaccades evoked more activation than errors in the right dorsolateral prefrontal cortex, anterior cingulate cortex (ACC), and presupplementary eye fields during this period. No significant differences were found for any comparisons early in the preparatory period. Our data suggest that the preparation of an antisaccade activates a large frontal and parietal network that may be involved in presetting the oculomotor system for the antisaccade task. These findings indicate that a large network of frontal and posterior areas is modulated during the latter component of the preparatory period on antisaccade compared with prosaccade trials. The results further suggest that the activation level of frontal cortical areas before stimulus presentation is associated with subjects' performance in the antisaccade task. In contrast, we found no areas that were more active for correct antisaccades than prosaccades or for correct antisaccades than error antisaccades during the stimulus-response period. In fact, a number of posterior cortical areas and a few areas in the superior frontal lobe were more active during the stimulus-response period on prosaccade trials than on antisaccade trials. Error antisaccades showed a larger activation in the ACC during the stimulus-response period compared with correct antisaccades.

Adult↗
PubMed2005

Rule-dependent activity for prosaccades and antisaccades in the primate prefrontal cortex.

Everyday life typically requires behavior that involves far more than simple stimulus-response associations. Environmental cues are often ambiguous and require different actions depending on the situation. The prefrontal cortex (PFC) is thought to be crucial for this flexible control of behavior. An important task that probes this ability is the antisaccade task in which subjects have to suppress a glance towards a suddenly presented peripheral stimulus and instead look away from the stimulus to its mirror location. Here we recorded the activity of PFC neurons in monkeys trained to alternate between blocks of prosaccade and antisaccade trials with no external instruction cues. We found that the activity of many neurons was different between the two tasks during the fixation period before the peripheral stimulus was presented. These differences were already present on the first correct trials after a task switch. The activity of these neurons also discriminated between correct responses and errors. We hypothesize that the PFC provides bias signals to saccade-related areas that are necessary to preset the oculomotor system for different tasks.

Action Potentials↗
PubMed2004

Effect of stimulus probability on anti-saccade error rates.

Subjects sometimes fail to suppress a reflexive saccade towards the flashed stimulus in an anti-saccade task. Here, we studied how error rates in the anti-saccade task varied as a function of saccadic probability. Ten subjects performed 200 anti-saccade trials for each of three saccade-direction probability conditions (20%, 50%, and 80%). We found that as the likelihood of a saccade in a given direction increased, the percentage of pro-saccade errors also increased for stimulus presentations in this direction. These results provide support for the hypothesis that errors in the anti-saccade task are the result of an increased level of motor preparation.

Adult↗
PubMed2003

Focused attention modulates visual responses in the primate prefrontal cortex.

Several current models propose an important role of the prefrontal cortex (PFC) in attention. To test the effects of attention in PFC, we recorded from PFC neurons in monkeys performing a task in which they had to attend to one hemifield and wait for a single stimulus that matched a previously presented cue. Neurons exhibited a slight decrease in their initial response and an enhanced activity late in the response to a stimulus at the cued location. The data demonstrate attentional effects on the activity of PFC neurons but they also show that single visual stimuli are initially represented in the activity of PFC neurons even when they are behaviorally irrelevant.

Animals↗
PubMed2003

Effects of ethanol on anti-saccade task performance.

It has been shown that saccade-related neurons in the superior colliculus (SC) display an increased level of prestimulus activity and a higher stimulus-related burst in action potentials preceding direction errors in the anti-saccade task compared with correct anti-saccades. From this, it has been hypothesized that errors occur when the incoming visual signal in the SC passes a threshold and triggers a reflexive saccade. This hypothesis predicts that an attenuated visual signal will reduce the number of direction errors. Since ethanol has been shown to have a suppressive effect on cortical visual event-related potentials (ERPs), the purpose of the present study was to investigate the effects of moderate ethanol consumption on anti-saccade performance. Under both placebo and ethanol conditions, we recorded ERPs and measured eye movements in male subjects during the performance of an anti-saccade task in which the fixation point disappeared 200 ms prior to stimulus presentation. Compared with the placebo condition, we found in the ethanol condition: (1). a decrease in ERP amplitudes during the gap period and after stimulus presentation, (2). an increase in the latencies of anti-saccades, and (3). a decrease in the percentage of direction errors. These data demonstrate the effects of ethanol on anti-saccade task performance and provide further support for the hypothesis that errors in the anti-saccade task are triggered by the incoming visual signal.

Adult↗
PubMed2003

Preparatory set associated with pro-saccades and anti-saccades in humans investigated with event-related FMRI.

Previous studies have shown that the BOLD functional MRI (fMRI) signal is increased in several cortical areas when subjects perform anti-saccades compared with pro-saccades. It remains unknown, however, whether this increase is due to an increased cortical motor signal for anti-saccades or due to differences in preparatory set between pro- and anti-saccade trials. To address this question, we measured event-related fMRI in a paradigm that allowed us to separate instruction-related brain activity from saccade-related brain activity. In this paradigm, the instruction to either generate a pro-saccade or an anti-saccade was conveyed by a switch in the color of the central fixation stimulus and preceded the presentation of a peripheral stimulus by either 6, 10, or 14 s. Cortical areas were functionally mapped using the general linear model comparing standard pro- and anti-saccade blocks with fixation blocks. When the trials were aligned on the onset of the instruction stimulus, bilateral frontal eye fields and right hemisphere dorsolateral prefrontal cortex showed an increased signal during the instruction period on anti-saccade trials as compared with pro-saccade trials. When the trials were aligned on the movement stimulus and the instruction period activity was subtracted, there were no differences between pro- and anti-saccades. This finding suggests that the increased cortical activation found in previous blocked designs originates predominately from differences in preparatory set and not from differences in the motor signal between pro- and anti-saccades.

Adult↗
PubMed2002

Contribution of the primate superior colliculus to inhibition of return.

The phenomenon of inhibition of return (IOR) has generated considerable interest in cognitive neuroscience because of its putative functional role in visual search, that of placing inhibitory tags on objects that have been recently inspected so as to direct further search to novel items. Many behavioral parameters of this phenomenon have been clearly delineated, and based on indirect but converging evidence, the widely held consensus is that the midbrain superior colliculus (SC) is involved in the generation of IOR. We had previously trained monkeys on a saccadic IOR task and showed that they displayed IOR in a manner similar to that observed in humans. Here we recorded the activity of single neurons in the superficial and intermediate layers of the SC while the monkeys performed this IOR task. We found that when the target was presented at a previously cued location, the stimulus-related response was attenuated and the magnitude of this response was correlated with subsequent saccadic reaction times. Surprisingly, this observed attenuation of activity during IOR was not caused by active inhibition of these neurons because (a) they were, in fact, more active following the presentation of the cue in their response field, and (b) when we repeated the same experiment while using the saccadic response time induced by electrical micro-stimulation of the SC to judge the level of excitability of the SC circuitry during the IOR task, we found faster saccades were elicited from the cued location. Our findings demonstrate that the primate SC participates in the expression of IOR; however, the SC is not the site of the inhibition. Instead, the reduced activity in the SC reflects a signal reduction that has taken place upstream.

Action Potentials↗
PubMed2002

Hand-eye coordination for rapid pointing movements. Arm movement direction and distance are specified prior to saccade onset.

Visually guided arm movements such as reaching or pointing are accompanied by saccadic eye movements that typically begin prior to motion of the arm. In the past, some degree of coupling between the oculomotor and limb motor systems has been demonstrated by assessing the relative onset times of eye and arm movement, and by the demonstration of a gap effect for arm movement reaction times. However, measures of limb movement onset time based on kinematics are affected by factors such as the relatively high inertia of the limb and neuromechanical delays. The goal of the present study was thus to assess the relative timing of rapid eye and arm movements made to visual targets by examining electromyographic (EMG) activity of limb muscles in conjunction with eye and arm position measures. The observation of a positive correlation between eye and limb EMG onset latencies, and the presence of a gap effect for limb EMG onset times (a reduction in reaction time when a temporal gap is introduced between the disappearance of a central fixation point and the appearance of a new target) both support the idea that eye and arm movement initiation are linked. However, limb EMG onset in most cases precedes saccade onset, and the magnitude of EMG activity prior to eye movement is correlated with both the direction and amplitude of the upcoming arm movement. This suggests that, for the rapid movements studied here, arm movement direction and distance are specified prior to the onset of saccades.

Arm↗
PubMed2002

Cortical activation associated with midtrial change of instruction in a saccade task.

The appearance of a visual stimulus in the peripheral visual field can elicit different saccade responses depending on prior instruction. This flexibility is commonly attributed to differences in motor set. Little is known about how the brain switches between one saccade response and another. To investigate the neural processes associated with switches between saccade motor sets, we recorded event-related potentials (ERPs) in 13 subjects, in three tasks that required subjects to generate prosaccades to a visual stimulus on 75% of the trials. On 25% of the trials, the color of the fixation point (FP) changed 300 ms prior to stimulus presentation. In the "ANTI" task, the change of the FP was the instruction to generate an antisaccade; in the "NOGO" task, subjects were instructed to maintain fixation; and in the "PRO" task, subjects were instructed to generate a prosaccade. The switch in motor set from prosaccades to antisaccades in the ANTI task and the cancellation of the prosaccade motor set in the NOGO task modulated frontal and frontocentral channels. Futhermore, the ANTI task but not the NOGO task was associated with differences at central and parietal channels compared with the PRO task. We hypothesize that the frontal activation in the ANTI and NOGO task reflects inhibition and task-switching processes, whereas the parietal activation reflects the preparation of this area for the sensorimotor transformation process that is necessary for the generation of an antisaccade.

Adult↗
PubMed2002

Contribution of the primate prefrontal cortex to the gap effect.

The introduction of a brief temporal gap between the disappearance of the initial fixation point and the presentation of a peripheral target leads to a general reduction in saccadic reaction times (SRTs), known as the gap effect. Moreover, extremely short latency express saccades frequently occur in this paradigm. Disorders of the prefrontal cortex (PFC) are often associated with increased numbers of express saccades and an inability to suppress reflexive saccades. To investigate the role of the PFC in the gap effect and in express saccade generation, we trained two rhesus monkeys on a gap saccade task in which the initial fixation point (FP) disappeared 200 ms or 600 ms before a peripheral stimulus appeared either 8 degrees to its left or right side. We recorded from the lateral PFC (areas 8 Ar and 46) in both monkeys the activity of 214 neurons, 84 (39%) of which exhibited task-related activity. These neurons could be further categorized into separate groups based on their discharge behaviour: fixation neurons with a decrease in activity during the gap (27%), FP offset neurons (12%), preparatory neurons with an increase in activity during the gap (30%), visual neurons (6%), post-saccadic neurons (8%), and reward-related neurons (12%). There were no obvious differences in the topography of these groups. Significant differences between express and regular saccade trials were found for fixation-related neurons. These neurons had a lower activity during the gap prior to the generation of contralateral express saccades. We hypothesize that a reduction in the activity of fixation-related neurons in the PFC may contribute to the elevated rate of express saccades in prefrontal disorders.

Animals↗
PubMed2002

Filtering of neural signals by focused attention in the monkey prefrontal cortex.

Prefrontal cortex is thought to be important in attention and awareness. Here we recorded the activity of prefrontal neurons in monkeys carrying out a focused attention task. Having directed attention to one location, monkeys monitored a stream of visual objects, awaiting a predefined target. Although neurons rarely discriminated between one non-target and another, they commonly discriminated between targets and non-targets. From the onset of the visual response, this target/non-target discrimination was effectively eliminated when the same objects appeared at an unattended location in the opposite visual hemifield. The results show that, in prefrontal cortex, filtering of ignored locations is strong, early and spatially global. Such filtering may be important in blindness to unattended signals--a conspicuous aspect of human selective attention.

Animals↗