David Souto

Effects of attention shifts to stationary objects during steady-state smooth pursuit eye movements

A number of studies have shown that stationary backgrounds compromise smooth pursuit eye movements. It has been suggested that poor attentional selection of the pursuit target was responsible for reductions of pursuit gain. To quantify the detrimental effects of attention, we instructed observers to either pay attention to background objects or to ignore them. The to-be-attended object was indicated by peripheral or central cues. Strong reductions of pursuit gain occurred when the following conditions were met: (a) the subject payed attention to the object (b) a salient event was present, for instance the onset of the target or cue and (c) the attended target produced retinal motion. Removing any of the three conditions resulted in no or far smaller decreases of pursuit gain. Further, decreases in pursuit gain were present with perceptual discrimination and simple manual detection.

Dynamics of attention during the initiation of smooth pursuit eye movements.

Many studies indicate that saccades are necessarily preceded by a shift of attention to the target location. There is no direct evidence for the same coupling during smooth pursuit. If smooth pursuit and attention were coupled, pursuit onset should be delayed whenever attention is focused on a stationary, non-target location. To test this hypothesis, observers were instructed to shift their attention to a peripheral location according to a location cue (Experiments 1 and 2) or a symbolic cue (Experiment 3) around the time of smooth pursuit initiation. Attending to static targets had only negligible effects on smooth pursuit latencies and the early open-loop response but lowered pursuit velocity substantially about the onset of closed-loop pursuit. Around this time, eye velocity reflected the competition between the to-be-tracked and to-be-attended object motion, entailing a reduction of eye velocity by 50% compared to the single task condition. The precise time course of attentional modulation of smooth pursuit initiation was at odds with the idea that an attention shift must precede any voluntary eye movement. Finally, the initial catch-up saccades were strongly delayed with attention diverted from the pursuit target. Implications for models of target selection for pursuit and saccades are discussed.

Involuntary cueing effects on accuracy measures: Stimulus and task dependence

Observers reported the orientation of a tilted grating that was presented together with a vertical distractor grating. In the main experiments, target contrast was low. There was location uncertainty because target location varied randomly and differences between target and distractor were small. In contrast to a previous report (T. Liu, F. Pestilli, & M. Carrasco, 2005), our results showed that non-informative peripheral cues do not improve perceptual performance at the cued location. However, informative peripheral or central cues improved perceptual performance. When we changed the task from an unspeeded perceptual task to a speeded reaction time task, the absence of involuntary cueing effects persisted when a distractor was presented. Without distractors, involuntary cueing effects re-emerged. When target contrast was increased, involuntary cueing effects re-emerged with a distractor but were smaller than without. We suggest that more difficult perceptual tasks reduce or abolish involuntary cueing effects.

Like a rolling stone: Naturalistic visual kinematics facilitate tracking eye movements

Newtonian physics constrains object kinematics in the real world. We asked whether eye movements towards tracked objects depend on their compliance with those constraints. In particular, the force of gravity constrains round objects to roll on the ground with a particular rotational and translational motion. We measured tracking eye movements towards rolling objects. We found that objects with rotational and translational motion that was congruent with an object rolling on the ground elicited faster tracking eye movements during pursuit initiation than incongruent stimuli. Relative to a condition without rotational component, we compared objects with this motion with a condition in which there was no rotational component, we essentially obtained benefits of congruence, and, to a lesser extent, costs from incongruence. Anticipatory pursuit responses showed no congruence effect, suggesting that the effect is based on visually-driven predictions, not on velocity storage. We suggest that the eye movement system incorporates information about object kinematics acquired by a lifetime of experience with visual stimuli obeying the laws of Newtonian physics.

Evidence for an attentional component in saccadic inhibition of return

After presentation of a peripheral cue, facilitation at the cued location is followed by inhibition of return (IOR). It has been recently proposed that IOR may originate at different processing stages for manual and ocular responses, with manual IOR resulting from inhibited attentional orienting, and ocular IOR resulting form inhibited motor preparation. Contrary to this interpretation, we found an effect of target contrast on saccadic IOR. The effect of contrast decreased with increasing reaction times (RTs) for saccades, but not for manual key-press responses. This may have masked the effect of contrast on IOR with saccades in previous studies (Hunt and Kingstone in J Exp Psychol Hum Percept Perform 29:1068–1074, 2003) because only mean RTs were considered. We also found that background luminance strongly influenced the effects of gap and target contrast on IOR.

Saccadic adaptation induced by a perceptual task.

The human motor system and muscles are subject to fluctuations in the short and long term. Motor adaptation is classically thought of as a low-level process that compensates for the error between predicted and executed movements in order to maintain movement accuracy. Contrary to a low-level account, accurate movements might be only a means to support high-level behavioral and perceptual goals. To isolate the influence of high-level goals in adaptation of saccadic eye movements, we manipulated perceptual task requirements in the absence of low-level errors. Observers had to discriminate one character within a peripheral array of characters. Between trials, the location of this character within the array was changed. This manipulation led to an immediate strategic change and a slower, gradual adaptation of saccade amplitude and direction. These changes had a similar magnitude to classical saccade adaptation and transferred at least partially to reactive saccades without a perceptual task. These results suggest that a perceptual task can modify oculomotor commands by generating a top-down error signal in saccade maps just like a bottom-up visual position error. Hence saccade adaptation not only maintains saccadic targeting accuracy, but also optimizes gaze behavior for the behavioral goal, showing that perception shapes even low-level oculomotor mechanisms.

Visual Flicker in the Gamma-Band Range Does Not Draw Attention

External transients, such as a flash or a startling sound, are believed to capture attention. Bauer, Cheadle, Parton, Müller, and Usher reported that attention can also be captured by a stimulus that flickers subliminally at 50 Hz, presumably by entrainment of neurons to the flicker frequency. In their reaction time (RT) task, participants had to locate a subtle change in the spatial frequency content of one of three Gabors (the target). Prior to target onset, presumably subliminal 50-Hz flicker in one of the Gabors served as a spatial cue. Bauer et al. found faster RTs when the cued location was congruent with the target location than when the cue was incongruent with the target location. In their experiments, the cue stopped to flicker at 50 Hz at target onset and was replaced by a stimulus flickering at 100 Hz (i.e., the screen refresh rate). In the present study, we show that the transition from 50 to 100 Hz results in a flash-like impression that can be localized above chance. We suggest that the illusory transition flash interfered with the localization of the subtle target, which contributed to the congruency effect. In support of this view, participants selected the flickering object more often than the non-flickering object when they failed to respond to the target. Further, no cueing effects were observed when the cue continued to flicker until the end of the trial or when the target was a salient change in polarity. In our view, the cueing effect occurs because observers confuse the illusory transition flash with the target when the two are similar. When truly subliminal flicker is used (70-Hz flicker), very small cueing effects persist in the absence of an illusory transition flash but may be accounted for by small effects on reaction time unrelated to attention.

Saliency Changes Appearance

Numerous studies have suggested that the deployment of attention is linked to saliency. In contrast, very little is known about how salient objects are perceived. To probe the perception of salient elements, observers compared two horizontally aligned stimuli in an array of eight elements. One of them was salient because of its orientation or direction of motion. We observed that the perceived luminance contrast or color saturation of the salient element increased: the salient stimulus looked even more salient. We explored the possibility that changes in appearance were caused by attention. We chose an event-related potential indexing attentional selection, the N2pc, to answer this question. The absence of an N2pc to the salient object provides preliminary evidence against involuntary attentional capture by the salient element. We suggest that signals from a master saliency map flow back into individual feature maps. These signals boost the perceived feature contrast of salient objects, even on perceptual dimensions different from the one that initially defined saliency.

Involuntary cueing effects during smooth pursuit: facilitation and inhibition of return in oculocentric coordinates

Peripheral cues induce facilitation with short cue-target intervals and inhibition of return (IOR) with long cue-target intervals. Modulations of facilitation and IOR by continuous displacements of the eye or the cued stimuli are poorly understood. Previously, the retinal coordinates of the cued location were changed by saccadic or smooth pursuit eye movements during the cue-target interval. In contrast, we probed the relevant coordinates for facilitation and IOR by orthogonally varying object motion (stationary, moving) and eye movement (fixation, smooth pursuit). In the pursuit conditions, cue and target were presented during the ongoing eye movement and observers made a saccade to the target. Importantly, we found facilitation and IOR of similar size during smooth pursuit and fixation. The results suggest that involuntary orienting is possible even when attention has to be allocated to the moving target during smooth pursuit. Comparison of conditions with stabilized and moving objects suggest an oculocentric basis for facilitation as well as inhibition. Facilitation and IOR were reduced with objects that moved on the retina both with smooth pursuit and eye fixation.

Perceptual task induces saccadic adaptation by target selection

Adaptation of saccades can be induced by different error signals, such as retinal position errors, prediction errors, or reinforcement learning. Recently, we showed that a shift in the spatial goal of a perceptual task can induce saccadic adaptation, in the absence of a bottom-up position error. Here, we investigated whether this top-down effect is mediated by the visibility of the task-relevant object, by reinforcement due to the feedback about the perceptual judgment or by a target selection mechanism. Participants were asked to discriminate visual stimuli arranged in a vertical compound. To induce adaptation, the discrimination target was presented at eccentric locations in the compound. In the first experiment, we compared adaptation with an easy and difficult discrimination. In the second experiment, we compared adaptation when feedback about the perceptual task was valid and when feedback was provided but was unrelated to performance. In the third experiment, we compared adaptation with instructions to fixate one of the elements in the compound—target selection—to the perceptual task condition—target selection and discrimination. To control for a bottom-up stimulus effect, we ran a fourth experiment in which the only instruction was to look at the compound. The saccade amplitude data were fitted by a two-state model distinguishing between an immediate and a gradual error correction process. We replicated our finding that a perceptual task can drive adaptation of saccades. Adaptation showed no effect of feedback reliability, nor an effect of the perceptual task beyond target selection. Adaptation was induced by a top-down signal since it was absent when there was no target selection instruction and no perceptual task. The immediate error correction was larger for the difficult than for the easy condition, suggesting that task difficulty affects mainly voluntary saccade targeting. In addition, the repetition of experiments one week later increased the magnitude of the gradual error correction. The results dissociate two distinct components of adaptation: an immediate and a gradual error correction. We conclude that perceptual-task induced adaptation is most likely due to top-down target selection within a larger object.