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Expected surprise, defined as the anticipation of uncertainty associated with the occurrence of a future event, plays a major role in gaze shifting and spatial attention. In the present study, we analyzed its impact on oculomotor behavior. We hypothesized that the occurrence of anticipatory saccades could decrease with increasing expected surprise and that its influence on visually-guided responses could be different given the presence of sensory information and perhaps competitive attentional effects. This hypothesis was tested in humans using a saccadic reaction time task in which a cue indicated the future stimulus position. In the ‘no expected surprise’ condition, the visual target could appear only at one previously cued location. In other conditions, more likely future positions were cued with increasing expected surprise. Anticipation was more frequent and pupil size was larger in the ‘no expected surprise’ condition compared with all other conditions, probably due to increased arousal. The latency of visually-guided saccades increased linearly with the logarithm of surprise (following Hick’s law) but their maximum velocity repeated the arousal-related pattern. Therefore, expected surprise affects anticipatory and visually-guided responses differently. Moreover, these observations suggest a causal chain linking surprise, attention and saccades that could be disrupted in attentional or impulse control disorders.

Patient suffering of major depressive disorder (MDD) often complain that subjective time seems to "drag" with respect to physical time. This may point toward a generalized dysfunction of temporal processing in MDD. In the present study, we investigated temporal preparation in MDD. "Temporal preparation" refers to an increased readiness to act before an expected event; consequently, reaction time should be reduced. MDD patients and age-matched controls were required to make a saccadic eye movement between a central and an eccentric visual target after a variable duration preparatory period. We found that MDD patients produced a larger number of premature saccades, saccades initiated prior to the appearance of the expected stimulus. These saccades were not temporally controlled; instead, they seemed to reflect reduced inhibitory control causing oculomotor impulsivity. In contrast, the latency of visually guided saccades was strongly influenced by temporal preparation in controls; significantly less so, in MDD patients. This observed reduced temporal preparation in MDD was associated with a faster decay of short-term temporal memory. Moreover, in patients producing a lot of premature responses, temporal preparation to early imperative stimuli was increased. In conclusion, reduced temporal preparation and short-term temporal memory in the oculomotor domain supports the hypothesis that temporal processing was altered in MDD patients. Moreover, oculomotor impulsivity interacted with temporal preparation. These observed deficits could reflect other underlying aspects of abnormal time experience in MDD.

Causality is a unique feature of human perception. We present here a behavioral investigation of the influence of physical causality during visual pursuit of object collisions. Pursuit and saccadic eye movements of human subjects were recorded during ocular pursuit of two concurrently launched targets, one that moved according to the laws of Newtonian mechanics (the causal target) and the other one that moved in a physically implausible direction (the non-causal target). We found that anticipation of collision evoked early smooth pursuit decelerations. Saccades to non-causal targets were hypermetric and had latencies longer than saccades to causal targets. In conclusion, before and after a collision of two moving objects the oculomotor system implicitly predicts upcoming physically plausible target trajectories.

During visual pursuit of a moving target, expected changes in its trajectory often evoke anticipatory smooth pursuit responses. In the present study, we investigated characteristics of anticipatory smooth pursuit decelerations before a change or the end of a target trajectory. Healthy humans had to pursue with the eyes a target moving along a circular path that predictably or unpredictably reversed direction and then retraced its movement back to the starting position. We found that anticipatory eye decelerations were often evoked in temporal expectation of target reversal and of the end of the trajectory. The latency of anticipatory decelerations initiated before target reversal was variable, had poor temporal accuracy and depended on the history of previous trials. Anticipations of the end of the trajectory were more accurate, more precise and were not influenced by previous trials. In this case, subjects probably based their estimate of the end of the trajectory on the duration just experienced before target motion reversal. These results suggest that anticipatory eye decelerations are based on the characteristics of the current or preceding trials depending on the most reliable information available.

Expectation of upcoming events is an essential cognitive function on which anticipatory actions are based. The neuronal basis of this prospective representation is poorly understood. We trained rhesus monkeys in a smooth-pursuit task in which the direction of upcoming target motion was indicated using a color cue. Under these conditions, directional expectation frequently evoked anticipatory smooth movements. We found that the activity of a population of neurons in the supplementary eye fields encoded the expected future direction of the target. Neuronal activity increased after presentation of the cue, indicating future target motion in the preferred direction. Neuronal activity either remained unaltered or was reduced if the antipreferred direction was cued. In addition, ∼30% of these neurons were more active during trials with anticipatory pursuit in the preferred direction than during trials when monkeys did not anticipate target motion onset. This subset of recorded neurons encoded the direction of the subsequent anticipatory pursuit. We hypothesize that the neural representation of directional expectation could be conceptualized as a competitive interaction between pools of neurons representing likely future events, with the winner of this competition determining the direction of the subsequent anticipatory movement. Similar mechanisms could drive prediction before movement initiation in other motor domains.

RATIONALE: It has been shown that antagonism of the glutamatergic N-methyl-D-aspartate (NMDA) receptor with subanesthetic doses of ketamine perturbs the perception of elapsed time. Anticipatory eye movements are based on an internal representation of elapsed time. Therefore, the occurrence of anticipatory saccades could be a particularly sensitive indicator of abnormal time perception due to NMDA receptors blockade. OBJECTIVES: To determine whether the occurrence of anticipatory saccades could be selectively altered by a subanesthetic dose of ketamine. METHODS: Three Rhesus monkeys were trained in a simple visually-guided saccadic task with a variable delay. Monkeys were rewarded for making a visually-guided saccade at the end of the delay. Premature anticipatory saccades to the future position of the eccentric target initiated before the end of the delay were not rewarded. A sub-anesthetic dose of ketamine (0.25 mg/kg) or a saline solution of the same volume was injected i.m. during the task. RESULTS: We found that the injected dose of ketamine did not induce sedation or abnormal behavior. However, in ~4 minutes, ketamine induced a strong reduction of the occurrence of anticipatory saccades but did not reduce the occurrence of visually-guided saccades. CONCLUSION: This unexpected reduction of anticipatory saccade occurrence could be interpreted as resulting from an altered use of the perception of elapsed time during the delay period induced by NMDA receptors antagonism.

A saccade triggered during sustained smooth pursuit is programmed using retinal information about the relative position and velocity of the target with respect to the eye. Thus the smooth pursuit and saccadic systems are coordinated by using common retinal inputs. Yet, in the absence of retinal information about the relative motion of the eye with respect to the target, the question arises whether the smooth and saccadic systems are still able to be coordinated possibly by using extraretinal information to account for the saccadic and smooth eye movements. To address this question, we flashed a target during smooth anticipatory eye movements in darkness, and the subjects were asked to orient their visual axis to the remembered location of the flash. We observed multiple orientation saccades (typically 2–3) toward the memorized location of the flash. The first orienting saccade was programmed using only the position error at the moment of the flash, and the smooth eye movement was ignored. However, subsequent saccades executed in darkness compensated gradually for the smooth eye displacement (mean compensation ≅ 70%). This behavior revealed a 400-ms delay in the time course of orientation for the compensation of the ongoing smooth eye displacement. We conclude that extraretinal information about the smooth motor command is available to the saccadic system in the absence of visual input. There is a 400-ms delay for smooth movement integration, saccade programming and execution.

The aim of the present study was to find out whether smooth eye movements (SEMs) evoked by superior colliculus (SC) stimulation are, as suggested by Breznen et al. (1996), artefactual eye movements resulting from a non-physiological response of the saccadic generator. This question was reinvestigated in head-restrained cats. Long-lasting SC stimulation was found to evoke, in a comparable proportion, either a single saccade followed by an uninterrupted SEM or a staircase of two or three saccades interleaved with SEMs. These two different patterns of eye movements could be elicited at a near-threshold current and at low stimulation frequencies. In most cases, SEM direction clearly differed from that of the preceding saccade. This difference between SEM and saccade directions varied in a systematic way as a function of the initial saccade direction. As demonstrated by computer simulation, this observation can be explained if the neural circuit controlling SEMs reaches a saturation level earlier than the saccadic burst-generator. Our results in cats were reminiscent of those reported by Breznen et al. (1996) in the monkey only in some instances, when high frequency stimulation (400-600 Hz) was applied. Indeed, in the case of near-threshold stimulation-elicited staircase saccades, increasing the stimulation frequency led to a progressive disappearance of the smaller subsequent saccades that were substituted by uninterrupted SEM-like movements. Altogether, the present results confirm the view that SEMs are genuine eye movements. These results rule out the hypothesis that SEMs result from a saturation of the saccadic generator and strengthen the hypothesis that SEMs and saccades are distinct movements. We suggest that the same collicular efferent cells carry out the motor command to saccadic and SEM circuits and that the position error originating from the SC may be distributed amongst separate downstream motor systems.

During visual tracking of a moving stimulus, primates orient their visual axis by combining two very different types of eye movements, smooth pursuit and saccades. The purpose of this paper was to investigate quantitatively the catch-up saccades occurring during sustained pursuit. We used a ramp-step-ramp paradigm to evoke catch-up saccades during sustained pursuit. In general, catch-up saccades followed the unexpected steps in position and velocity of the target. We observed catch-up saccades in the same direction as the smooth eye movement (forward saccades) as well as in the opposite direction (reverse saccades). We made a comparison of the main sequences of forward saccades, reverse saccades, and control saccades made to stationary targets. They were all three significantly different from each other and were fully compatible with the hypothesis that the smooth pursuit component is added to the saccadic component during catch-up saccades. A multiple linear regression analysis was performed on the saccadic component to find the parameters determining the amplitude of catch-up saccades. We found that both position error and retinal slip are taken into account in catch-up saccade programming to predict the future trajectory of the moving target. We also demonstrated that the saccadic system needs a minimum period of approximately 90 ms for taking into account changes in target trajectory. Finally, we reported a saturation (above 15°/s) in the contribution of retinal slip to the amplitude of catch-up saccades.

When tracking moving visual stimuli, primates orient their visual axis by combining two kinds of eye movements, smooth pursuit and saccades, that have very different dynamics. Yet, the mechanisms that govern the decision to switch from one type of eye movement to the other are still poorly understood, even though they could bring a significant contribution to the understanding of how the CNS combines different kinds of control strategies to achieve a common motor and sensory goal. In this study, we investigated the oculomotor responses to a large range of different combinations of position error and velocity error during visual tracking of moving stimuli in humans. We found that the oculomotor system uses a prediction of the time at which the eye trajectory will cross the target, defined as the “eye crossing time” ( T XE). The eye crossing time, which depends on both position error and velocity error, is the criterion used to switch between smooth and saccadic pursuit, i.e., to trigger catch-up saccades. On average, for T XEbetween 40 and 180 ms, no saccade is triggered and target tracking remains purely smooth. Conversely, when T XEbecomes smaller than 40 ms or larger than 180 ms, a saccade is triggered after a short latency (around 125 ms).

Anticipatory eye movements are often evoked by the temporal expectation of an upcoming event. Temporal expectation is based on implicit timing about when a future event could occur. Implicit timing emerges from observed temporal regularities in a changing stimulus without any voluntary estimate of elapsed time, unlike explicit timing. The neural bases of explicit and implicit timing are likely different. It has been shown that the basal ganglia (BG) play a central role in explicit timing. In order to determine the influence of BG in implicit timing, we investigated the influence of early Parkinson’s disease (PD) and aging on the latency of anticipatory eye movements. We hypothesized that a deficit of implicit timing should yield inadequate temporal expectations, and consequently abnormally timed anticipatory eye movements compared with age-matched controls. To test this hypothesis, we used an oculomotor paradigm where anticipation of a salient target event plays a central role. Participants pursued a visual target that moved along a circular path at a constant velocity. After a randomly short (1200 ms) or long (2400 ms) forward path, the target reversed direction, returned to its starting position and stopped. Target motion reversal caused an abrupt "slip" of the pursued target image on the retina and was a particularly salient event evoking anticipatory eye movements. Anticipatory eye movements were less frequent in PD patients. However, the timing of anticipation of target motion reversal was statistically similar in PD patients and control subjects. Other eye movements showed statistically significant differences between PD and controls, but these differences could be attributed to other factors. We conclude that all anticipatory eye movements are not similarly impaired in PD and that implicit timing of salient events seems largely unaffected by this disease. The results support the hypothesis that implicit and explicit timing are differently affected by BG dysfunction.

When two objects such as billiard balls collide, observers perceive that the action of one caused the motion of the other. We have previously shown (Badler, Lefèvre, & Missal, 2010) that this extends to the oculomotor domain: subjects make more predictive movements in the expected direction of causal motion than in a noncausal direction. However, predictive oculomotor and reactive psychophysical responses have never been directly compared. They should be correlated if they tap into the same mental processes. To test this, we recorded oculomotor responses to launching stimuli, then asked subjects to manually classify those stimuli as causal or noncausal. Overall the psychophysical classifications matched the oculomotor biases, although correlations across subjects were mostly absent. In subsequent experiments, 50% of the trials had a 300-millisecond delay after the collision to impede the perception of causality. Subjects maintained their causal oculomotor bias but used different classification strategies, usually grouping the stimuli either by delay or by direction. In addition, there was no evidence that the two response types were correlated on a trial-by-trial basis. The results suggest divergent processes underlying oculomotor responses to and judgments of causal stimuli.

Recent experimental results show evidence for the corrective role of postsaccadic drifts. This paper addresses the modeling of these slow correcting gaze movements (SCMs). Classical arguments to explain drifts are presented, both in the head fixed condition (pulse-step mismatch) and in the head free condition (vestibulo-ocular reflex (VOR)). The most significant behavioral and electrophysiological experimental data related to SCMs are then briefly reported, with the conclusion that SCMs have a clear corrective role, incompatible with classical explanations of drifts. Based on these experimental data, existing models of the saccadic system are then compared. A theoretical comparison of the classical Robinson model with an alternative model is proposed. Two possible (slow and fast) pathways involved in the control of SCMs are examined, and simulation results are presented. Finally, the discussion addresses the observed species differences in SCMs. The link between natural SCMs and electrical SC stimulations, and the interactions between saccades, VOR, and SCMs are also discussed.

When viewing one object move after being struck by another, humans perceive that the action of the first object “caused” the motion of the second, not that the two events occurred independently. Although established as a perceptual and linguistic concept, it is not yet known whether the notion of causality exists as a fundamental, preattentional “Gestalt” that can influence predictive motor processes. Therefore, eye movements of human observers were measured while viewing a display in which a launcher impacted a tool to trigger the motion of a second “reaction” target. The reaction target could move either in the direction predicted by transfer of momentum after the collision (“causal”) or in a different direction (“noncausal”), with equal probability. Control trials were also performed with identical target motion, either with a 100 ms time delay between the collision and reactive motion, or without the interposed tool. Subjects made significantly more predictive movements (smooth pursuit and saccades) in the causal direction during standard trials, and smooth pursuit latencies were also shorter overall. These trends were reduced or absent in control trials. In addition, pursuit latencies in the noncausal direction were longer during standard trials than during control trials. The results show that causal context has a strong influence on predictive movements.

The ability to predict upcoming events is important to compensate for relatively long sensory-motor delays. When stimuli are temporally regular, their prediction depends on a representation of elapsed time. However, it is well known that the allocation of attention to the timing of an upcoming event alters this representation. The role of attention on the temporal processing component of prediction was investigated in a visual smooth pursuit task that was performed either in isolation or concurrently with a manual response task. Subjects used smooth pursuit eye movements to accurately track a moving target after a constant-duration delay interval. In the manual response task, subjects had to estimate the instant of target motion onset by pressing a button. The onset of anticipatory pursuit eye movements was used to quantify the subject's estimate of elapsed time. We found that onset times were delayed significantly in the presence of the concurrent manual task relative to the pursuit task in isolation. There was also a correlation between the oculomotor and manual response latencies. In the framework of Scalar Timing Theory, the results are consistent with a centralized attentional gating mechanism that allocates clock resources between smooth pursuit preparation and the parallel timing task.

The timing of an upcoming event depends on two factors: its temporal position, proximal or distal with respect to the present moment, and the unavoidable stochastic variability around this temporal position. We searched for a general mechanism that could describe how these two factors influence the anticipation of an upcoming event in an oculomotor task. Monkeys were trained to pursue a moving target with their eyes. During a delay period inserted before target motion onset, anticipatory pursuit responses were frequently observed. We found that anticipatory movements were altered by the temporal position of the target. Increasing the timing uncertainty associated with the stimulus resulted in an increase in the width of the latency distribution of anticipatory pursuit. These results show that monkeys relied on an estimation of the changing probability of target motion onset as time elapsed during the delay to decide when to initiate an anticipatory smooth eye movement.

To organise behaviors, the brain makes predictions from past experience. Addicted subjects, who persist in consuming despite negative consequences, may be unable to learn from past experience. Anticipatory oculomotor movements can be accurately studied in with eye tracking techniques and may serve as an index of procedural learning processes. …

A classic paradigm to study anticipatory pursuit consists in training monkeys to look at a target that appears in the center of a visual display, disappears during a short “gap” period, then reappears and immediately starts to move. To determine the role of prior directional information on anticipatory pursuit eye movements, we trained rhesus monkeys to associate the color of a centrally presented visual cue with the direction of an upcoming target motion. In a first experiment, a gap period occurred randomly in 50% of the trials. Consequently, two possible choices of timing of target motion onset were given to subjects to guide their anticipatory responses. In a second experiment, a gap period occurred during each trial and only a single choice of timing of target motion onset was given to subjects. We found that monkeys used the learned association between the color of the cue and the direction of future target motion to voluntarily initiate anticipatory pursuit movements in the appropriate direction. Anticipatory movements could be classified in two distinct populations: early and late movements. Early movements were most frequent when prior directional information was provided and when two choices of timing of target motion onset were given. The latency of visual pursuit was shortened and its velocity was larger when prior directional information was provided. We conclude that cognitive expectation of future target motion plays a dominant role in determining characteristics of anticipatory pursuit in the monkey.

Anticipatory movements are motor responses occurring before likely sensory events in contrast to reflexive actions. Anticipatory movements are necessary to compensate for delays present in sensory and motor systems. Smooth pursuit eye movements are often used as a paradigmatic example for the study of anticipation. However, the neural control of anticipatory pursuit is unknown. A previous study suggested that the supplementary eye fields (SEFs) could play a role in the guidance of smooth pursuit to predictable target motion. In this study, we favored anticipatory responses in monkeys by making the parameters of target motion highly predictable and electrically stimulated the SEF before and during this behavior. Stimulation sites were restricted to regions of the SEF where saccades could not be evoked at the same low currents. We found that electrical microstimulation in the SEF increased the velocity of anticipatory pursuit movements and decreased their latency. These effects will be referred to as anticipatory pursuit facilitation. The degree of facilitation was the largest if the stimulation train was delivered near the end of the fixation period, before the moment when anticipatory pursuit usually begins. No anticipatory smooth eye movements could be evoked during fixation without an expectation of target motion. These results suggest that the SEF pursuit area might be involved in the process of guiding anticipatory pursuit.

A long-standing belief holds that the saccadic and smooth-pursuit eye movement systems are composed of largely separate premotor circuits, at least in the brainstem. One crucial prediction predicated on this belief is that the tonic discharge of omnipause neurons (OPNs), which are thought to be part of only the saccadic system, should not be modulated during pursuit eye movements. This report shows that the discharge of OPNs, in contradiction, is modulated downward during pursuit movements. In contrast to their behavior during saccades, where they pause completely for the duration of the movement, the downward modulation during pursuit did not totally silence OPNs. The depth of the downward modulation was correlated with the speed of the ongoing pursuit movement. Another type of cell, which we have named saccade/pursuit neurons, was recorded in the paramedian pontine reticular formation near the location of OPNs. This subpopulation of burst cells discharged a cascade of spikes for saccades in a preferred direction. They also displayed a lower-frequency sustained discharge of spikes for the duration of pursuit in the same preferred direction. These data suggest a new type of combined model for the organization of the brainstem saccade/pursuit system. In this new combined model, the OPNs form a common inhibitory mechanism for both types of movements, and the saccade/pursuit neurons participate in the eye-velocity modulation of OPN discharge or membrane polarization during either type of movement.

Briefly flashed visual stimuli presented during smooth object- or self-motion are systematically mislocalized. This phenomenon is called the "flash-lag effect" (Nijhawan, 1994). All previous studies had one common characteristic, the subject's sense of motion. Here we asked whether motion perception is a necessary condition for the flash-lag effect to occur. In our first experiment, we briefly flashed a target during smooth anticipatory eye movements in darkness and subjects had to orient their gaze toward the perceived flash position. Subjects reported to have no sense of eye motion during anticipatory movements. In our second experiment, subjects had to adjust a cursor on the perceived position of the flash. As a result, we show that gaze orientation reflects the actual perceived flash position. Furthermore, a flash-lag effect is present despite the absence of motion perception. Moreover, the time course of gaze orientation shows that the flash-lag effect appeared immediately after the egocentric to allocentric reference frame transformation.

The role of the supplementary eye fields (SEF) during smooth pursuit was investigated with electrical microstimulation. We found that stimulation in the SEF increased the acceleration and velocity of the eyes in the direction of target motion during smooth pursuit initiation but not during sustained pursuit. The increase in eye velocity during initiation will be referred to as pursuit facilitation and was observed at sites where saccades could not be evoked with the same stimulation parameters. On average, electrical stimulation increased eye velocity by ∼20%. At most sites, the threshold for a significant facilitation was 50 μA with a stimulation frequency of 300 Hz. Facilitation of pursuit initiation depended on the timing of stimulation trains. The effect was most pronounced if the stimulation was delivered before smooth pursuit initiation. On average, eye velocity in stimulation trials increased linearly as a function of eye velocity in control trials, and this function had a slope greater than one, suggesting a multiplicative influence of the stimulation. Stimulation during a fixation task did not evoke smooth eye movements. The latency of catch-up saccades was increased during facilitation, but their accuracy was not affected. Saccades toward stationary targets were not affected by the stimulation. The results are further evidence that the SEF plays a role in smooth pursuit in addition to its known role in saccade planning and suggest that this role may be to control the gain of smooth pursuit during initiation. The covariance between pursuit facilitation and the timing of the catch-up saccade as a result of stimulation suggests that these different eye movements systems are coordinated to achieve a common goal.

Differences between gaze shifts evoked by collicular electrical stimulation and those triggered by the presentation of a visual stimulus were studied in head-free cats by increasing the head moment of inertia. This maneuver modified the dynamics of these two types of gaze shifts by slowing down head movements. Such an increase in the head moment of inertia did not affect the metrics of visually evoked gaze saccades because their duration was precisely adjusted to compensate for these changes in movement dynamics. In contrast, the duration of electrically evoked gaze shifts remained constant irrespective of the head moment of inertia, and therefore their amplitude was significantly reduced. These results suggest that visually and electrically evoked gaze saccades are controlled by different mechanisms. Whereas the accuracy of visually evoked saccades is likely to be assured by on-line feedback information, the absence of duration adjustment in electrically evoked gaze shifts suggests that feedback information necessary to maintain their metrics is not accessible or is corrupted during collicular stimulation. This is of great importance when these two types of movements are compared to infer the role of the superior colliculus in the control of orienting gaze shifts.

We tested whether macaque inferior temporal neurons can signal the presence of their preferred shape independently of other shapes simultaneously present, by comparing the responses and selectivity of TE neurons to shapes (figures), either presented in isolation or overlapping another shape (background). The two overlapping shapes differed in color or texture and thus were easily segmentable. We found that the response and selectivity of TE neurons to the figure can be dramatically altered, most commonly reduced, when the figure is overlapping the background. This reduction in response was also present when the monkey was required to actively discriminate the figures from varying backgrounds during recording. The level of suppression depended on the shape of the background and on whether or not figure and background can be discriminated. These results indicate that responses of TE cells are not only determined by the properties of the figures but also are influenced by the properties of stimuli in the background.

Introduction Learning involves setting up predictions in identical situations. The dopaminergic striato-frontocortical loop plays a major role in setting up these predictions. The activity of the loop may be assessed by measuring the anticipation of eye movements, following a target on a screen, using an eye tracker. We have recently observed that deficits in anticipation were correlated with scores of impulsivity on the UPPS scale. Objectives Measure learning abilities in AD subjects, where PET studies have shown a decrease in striatal dopaminergic activity. Aims Test the capacity of AD and control subjects on the anticipatory eye movements. Methods: 24 AD subjects were tested at onset and end of a 3 weeks detox and compared with 24 age and gender matched controls also tested twice, in an occulomotor task. Results Control subjects displayed higher velocity than AD subjects in the period of anticipation of the target on the screen. Furthermore, control subjects displayed improvement in the velocity and amplitude of this anticipatory eye movement, suggesting the existence of learning abilities supported by the striato-frontocortical loop. On the contrary, no improvement occured in AD subjects even after detox, suggesting a blunting of the learning abilities of these subjects. Conclusions The striato-fronto cortical loops that probably plays a role in the learning of the addiction to alcohol in AD subjects is no longer able to support learning processes, which may explain the difficulty of these subjects to unlearn the addiction.

Humans are very sensitive to the presence of other living persons or animals in their surrounding. Human actions can readily be perceived, even in a noisy environment. We recently demonstrated that biological motion, which schematically represents human motion, influences smooth pursuit eye movements during the initiation period (Orban de Xivry, Coppe, Lefèvre, & Missal, 2010). This smooth pursuit response is driven both by a visuomotor pathway, which transforms retinal inputs into motor commands, and by a memory pathway, which is directly related to the predictive properties of smooth pursuit. To date, it is unknown which of these pathways is influenced by biological motion. In the present study, we first use a theoretical model to demonstrate that an influence of biological motion on the visuomotor and memory pathways might both explain its influence on smooth pursuit initiation. In light of this model, we made theoretical predictions of the possible influence of biological motion on smooth pursuit during and after the transient blanking of the stimulus. These qualitative predictions were then compared with recordings of eye movements acquired before, during and after the transient blanking of the stimulus. The absence of difference in smooth pursuit eye movements during blanking of the stimuli and the stronger visually guided smooth pursuit reacceleration after reappearance of the biological motion stimuli in comparison with control stimuli suggests that biological motion influences the visuomotor pathway but not the memory pathway.

The capacity to identify and perceive biologically relevant actions is essential for survival and social skills. We hypothesized that the perception of biological motion could induce a different behavioral response compared with stimuli devoid of biological relevance. To check this hypothesis, we analyzed the eye movement response to a moving walker, or its scrambled version. Subjects were asked to pursue a point-light walker, created using Cutting’s algorithm, or its scrambled version. The walker consisted of 11 dots; a green fixation point on the hip dot and 10 red dots. The control stimulus - scrambled version- was obtained by shuffling the mean vertical position of the dots of the walker -except the hip dot- to disrupt the global form while keeping the same local motion. The point-light walker, or its scrambled version, then appeared and began to move in the randomized heading direction for 800ms before disappearing behind an invisible occluder. The stimulus reappeared 890ms later for 800ms. The type of stimulus -biological motion or scrambled version- its direction and its velocity were selected at random for each trial. We analyzed separately 4 different phases in the responses. In the first one -the reactive smooth pursuit-, we saw that the motor response was stronger with a biological stimulus. The smooth eye velocity was significantly greater for the walker 200ms after pursuit onset. We noticed no difference in eye movement velocity between both stimuli neither during the steady-state phase nor in the occlusion phase. But after reappearance of the stimulus, we saw a stronger smooth pursuit response for the biological motion. The mean acceleration of smooth pursuit eye movements for the first 150ms after stimulus reappearance was significantly higher for biological motion than for the control stimulus. These results show that the biological relevance of an action can influence the behavioral response to the visual stimulus.

The premotor pathways subserving saccades and smooth-pursuit eye movements are usually thought to be different. Indeed, saccade and smooth-pursuit eye movements have different dynamics and functions. In particular, a group of midline cells in the pons called omnipause neurons (OPNs) are considered to be part of the saccadic system only. It has been established that OPNs keep premotor neurons for saccades under constant inhibition during fixation periods. Saccades occur only when the activity of OPNs has completely stopped or paused. Accordingly, electrical stimulation in the region of OPNs inhibits premotor neurons and interrupts saccades. The premotor relay for smooth pursuit is thought to be organized differently and omnipause neurons are not supposed to be involved in smooth-pursuit eye movements. To investigate this supposition, OPNs were recorded during saccades and during smooth pursuit in the monkey (Macaca mulatta). Unexpectedly, we found that neuronal activity of OPNs decreased during smooth pursuit. The resulting activity reduction reached statistical significance in approximately 50% of OPNs recorded during pursuit of a target moving at 40 degrees /s. On average, activity was reduced by 34% but never completely stopped or paused. The onset of activity reduction coincided with the onset of smooth pursuit. The duration of activity reduction was correlated with pursuit duration and its intensity was correlated with eye velocity. Activity reduction was observed even in the absence of catch-up saccades that frequently occur during pursuit. Electrical microstimulation in the OPNs' area induced a strong deceleration of the eye during smooth pursuit. These results suggest that OPNs form an inhibitory mechanism that could control the time course of smooth pursuit. This inhibitory mechanism is part of the fixation system and is probably needed to avoid reflexive eye movements toward targets that are not purposefully selected. This study shows that saccades and smooth pursuit, although they are different kinds of eye movements, are controlled by the same inhibitory system.