Thérèse Collins

Post-saccadic location judgments reveal remapping of saccade targets to non-foveal locations

The present study addresses the question of how objects are localized across saccades. In a task requiring participants to compare the location of a post-saccadic probe with the pre-saccadic target, we investigated the roles of saccade landing site and post-saccadic probe location. Saccade landing sites vary from trial to trial because of oculomotor error but can also be shifted by saccadic adaptation. Visual targets were extinguished during the saccade and reappeared after a short blank to counteract saccadic suppression of displacement. Performance in localizing targets after unadapted saccades was nearly veridical and independent of actual landing site, showing that trial-to-trial oculomotor error did not contribute to post-saccadic localization. This result suggests that much of the oculomotor error of saccades is included in the efference copy vector and this allows the recovery of a remapped target location that is often not foveal, but stable and accurate across trials. Displacement judgments relative to this remapped location will be independent of trial-to-trial variability in landing site. After adapted saccades, post-saccadic localization shifted in the direction opposite to adaptation but again, trial-by-trial landing site variability did not correlate with performance. This result suggests that the efference copy matches the planned upcoming saccade, be it adapted or not.

Motor space structures perceptual space: Evidence from human saccadic adaptation

Saccadic adaptation is the progressive correction of systematic saccade targeting errors. When a saccade to a particular target is adapted, saccades within a spatial window around the target, the adaptation field, are affected as a function of their distance from the adapted target. Furthermore, previous studies suggest that saccadic adaptation might modify the perceptual localization of objects in space. We investigated the localization of visual probes before and after saccadic adaptation, and examined whether the spatial layout of the observed mislocalizations was structurally similar to the saccadic adaptation field. We adapted a horizontal saccade directed towards a target 12 degrees to the right. Thirty-eight saccades towards the right visual hemifield were then used to measure the adaptation field. The adaptation field was asymmetric: transfer of adaptation to saccades larger than the adapted saccade was greater than transfer to smaller saccades. Subjects judged the localization of 39 visual probes both within and outside the adaptation field. The perceived localization of a probe at a given position was proportional to the amount of transfer from the adapted saccade to the saccade towards that position. This similar effect of saccadic adaptation on both the action and perception representations of space suggests that the system providing saccade metrics also contributes to the metric used for the perception of space.

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

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

Action Goal Selection and Motor Planning Can Be Dissociated by Tool Use.

The preparation of eye or hand movements enhances visual perception at the upcoming movement end position. The spatial location of this influence of action on perception could be determined either by goal selection or by motor planning. We employed a tool use task to dissociate these two alternatives. The instructed goal location was a visual target to which participants pointed with the tip of a triangular hand-held tool. The motor endpoint was defined by the final fingertip position necessary to bring the tool tip onto the goal. We tested perceptual performance at both locations (tool tip endpoint, motor endpoint) with a visual discrimination task. Discrimination performance was enhanced in parallel at both spatial locations, but not at nearby and intermediate locations, suggesting that both action goal selection and motor planning contribute to visual perception. In addition, our results challenge the widely held view that tools extend the body schema and suggest instead that tool use enhances perception at those precise locations which are most relevant during tool action: the body part used to manipulate the tool, and the active tool tip.

Saccadic adaptation shifts the pre-saccadic attention focus

The well-documented phenomenon of the spatial coupling between saccadic programming and the orienting of attention refers to the fact that visual attention is directed toward the location that the eyes are aiming for. However, the question remains open as to whether saccades and attention are two independent processes that can be directed concurrently toward a common goal, or whether their relationship is tighter, with the motor components of the saccade program influencing the selection of the position towards which visual attention is directed. To investigate this issue, an experiment was carried out in which the initial saccade goal was dissociated from the final executed motor vector. This was done by using a saccadic adaptation paradigm and a discrimination task. Results showed that best perceptual performance, which is taken to be an indicator of the locus of visual attention, followed the motor modifications arising from saccadic adaptation. This suggests that visual attention is directed toward the actual saccade landing position and that the perceptual system must have access to information regarding the motor vector before saccade execution.

Decision and metrics of refixations in reading isolated words

Eye movements were recorded during the reading of long words presented in isolation. Overall, the decision to refixate was found to depend on both length and frequency of the word, while refixation amplitude depended only on word length. This finding corroborates the assumption that most refixation saccades are preplanned on the basis of the parafoveal word length. However, cancellation of such a plan is possible and could be linked to the lexical processing during the first fixation into the word. Finally, a small proportion of refixations are corrective saccades, related to an oculomotor error. Theoretical implications for models of eye movement control during reading are discussed.

Extraretinal signal metrics in multiple-saccade sequences

Executing sequences of memory-guided movements requires combining sensory information with information about previously made movements. In the oculomotor system, extraretinal information must be combined with stored visual information about target location. The use of extraretinal signals in oculomotor planning can be probed in the double-step task. Using this task and a multiple-step version, the present study examined whether an extraretinal signal was used on every trial, whether its metrics represented desired or actual eye displacement, and whether it was best characterized as a direct estimate of orbital eye position or a vector representation of eye displacement. The results show that accurate information, including saccadic adaptation, about the first saccade is used to plan the second saccade. Furthermore, with multiple saccades, endpoint variability increases with the number of saccades. Controls ruled out that this was due to the perceptual or memory requirements of storing several target locations. Instead, each memory-guided movement depends on an internal copy of an executed movement, which may present a small discrepancy with the actual movement. Increasing the number of estimates increases the variability because this small discrepancy accumulates over several saccades. Such accumulation is compatible with a corollary discharge signal carrying metric information about saccade vectors.

Visual Versus Motor Vector Inversions in the Antisaccade Task: A Behavioral Investigation With Saccadic Adaptation

In the antisaccade task, subjects must execute an eye movement away from a visual target. Correctly executing an antisaccade requires inhibiting a prosaccade toward the visual target and programming a movement to the opposite side. This movement could be based on the inversion of the visual vector, corresponding to the distance between the fixation point and the visual target, or the motor vector of the unwanted prosaccade. We dissociated the two vectors by means of saccadic adaptation. Adaptation can be observed when systematic targeting errors are caused by the displacement of the visual target during the saccade. Adaptation progressively modifies saccade amplitude (defined by the motor vector) such that it becomes appropriate to the postsaccadic stimulus position and thus different from the visual vector of the target. If antisaccade preparation depended on visual vector inversion, rightward prosaccade adaptation should not transfer to leftward antisaccades (which are based on the same visual vector) but should transfer to rightward antisaccades (which are based on a visual vector inside the adaptation field). If antisaccade preparation depended on motor vector inversion, rightward prosaccade adaptation should transfer to leftward antisaccades (which are based on the same, adapted motor vector) but should not transfer to rightward antisaccades (which are based on a nonadapted motor vector). The results are in line with the first hypothesis, showing that vector inversion precedes saccadic adaptation and suggesting that antisaccade preparation depends on the inversion of the visual target vector.

The use of recurrent signals about adaptation for subsequent saccade programming depends on object structure

Executing sequences of accurate saccadic eye movements supposes the use of signals carrying information about the first saccade for updating the predetermined motor plan of the subsequent saccades. The present study examines the signals used in planning a second saccade when subjects made two successive saccades towards one long or two short peripheral objects displayed before the first saccade execution. Different first eye movement signals could be used: desired eye movement signals, representing the movement necessary for attaining the intended target, or actual eye movement signals, representing the movement actually executed. Experimental dissociation of desired and actual eye movement signals is made possible by adaptive modifications of the first saccade, obtained by transfer of single saccade adaptation, during which the motor vector was progressively modified in response to the systematic intra-saccadic step of a single target. Whether the second saccade used the actual eye movement signal to compensate or not for the adaptive changes in the first saccade depended on which object properties were relevant for saccade planning. Compensation was observed for saccades that aimed for a new object (between-object saccades) because adaptation modifies relative object location. No compensation was observed for saccades that explored an extended object (within-object saccades). Implications for the on-line control of subsequent eye movements are discussed.

Visual target selection and motor planning define attentional enhancement at perceptual processing stages.

Extracting information from the visual field can be achieved by covertly orienting attention to different regions, or by making saccades to bring areas of interest onto the fovea. While much research has shown a link between covert attention and saccade preparation, the nature of that link remains a matter of dispute. Covert presaccadic orienting could result from target selection or from planning a motor act toward an object. We examined the contribution of visual target selection and motor preparation to attentional orienting in humans by dissociating these two habitually aligned processes with saccadic adaptation. Adaptation introduces a discrepancy between the visual target evoking a saccade and the motor metrics of that saccade, which, unbeknownst to the participant, brings the eyes to a different spatial location. We examined attentional orienting by recording event-related potentials (ERPs) to task-irrelevant visual probes flashed during saccade preparation at four equidistant locations including the visual target location and the upcoming motor endpoint. ERPs as early as 130–170 ms post-probe were modulated by attention at both the visual target and motor endpoint locations. These results indicate that both target selection and motor preparation determine the focus of spatial attention, resulting in enhanced processing of stimuli at early visual-perceptual stages.