Adaptation of Saccadic Eye Movements: Transfer and Specificity
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The present study was designed to test whether the adaptation of saccadic eye movements depends only on the eye displacement vector of the trained saccade or also on eye position information. Using the double-step target paradigm in eight human subjects, we first induced in a single session two "opposite directions adaptations" (ODA) of horizontal saccades of the same vector. Each ODA (backward or forward) was linked to one vertical eye position (12.5 degrees up or 25 degrees down) and alternated from trial to trial. The results showed that opposite changes of saccade amplitude can develop simultaneously, indicating that saccadic adaptation depends on orbital eye position. This finding has important functional implications because in everyday life our eyes saccade from constantly changing orbital positions. A comparison of these data to two control conditions in which training trials of a single type (backward or forward) were presented at both 12.5 degrees and -25 degrees eye elevations further indicated that eye position specificity is complete for backward, but not for forward, adaptation. Finally, the control conditions also indicated that the adaptation of a single saccade fully transferred to untrained saccades of the same vector, but initiated from different vertical eye positions. In conclusion, our study indicates that saccadic adaptation mechanisms use vectorial eye displacement signals, but can also take eye position signals into account as a contextual cue when the training involves conflicting saccade amplitude changes.Keywords:
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When the oculomotor system faces a discrepancy between the saccade endpoint and the visual target, it can attribute this discrepancy to a change in the environment or to a visual saccadic error. In the laboratory, a progressive adjustment in saccade amplitude is observed in the presence of a systematic displacement of the target during the saccade. According to literature, saccadic adaptation is disrupted when the target is shortly blanked after the saccade, and disappears around a 600ms-blank. Because an increased blank leads to the perception of target displacements, saccade amplitude modifications are thought to be negligible. We however believe that in the presence of a systematically varying environment, we are still likely to adjust our saccadic amplitudes, but that the nature of this adjustment should differ. We examined, in a double-step adaptation paradigm, the effect of various temporal delays in the presentation of the post-saccadic target on the amount of saccadic adaptation. We hypothesize that an automatic and slow modification of saccadic amplitude (implicit learning) should take place for intrassacadic target displacements, whereas a more explicit learning (voluntary and fast) should dominate when target reappearance is delayed. Preliminary results show that saccades’ amplitude is modified up to a 1200ms delay (gain change: M=-14%, SD=10%), and still visible in large proportion up to 600ms (M=-20%, SD=8.9%). Our visual system seems to “adapt” eye movements even when target displacements can be attributed to a change in the environment and not to saccadic errors anymore. Interestingly, we observed an increase in saccadic endpoint variability (entropy, exploration) and saccades’ latency during adaptation, associated with a smaller amount of saccadic adaptation, when target reappearance was delayed and its displacement perceived. Our results indicate that adaptation profiles likely depend on how participants experienced their saccade landing errors.
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Abstract The accuracy of saccadic eye movements is maintained by saccadic adaptation, a learning mechanism that is proposed to rely on visual prediction error, i.e., a mismatch between the pre-saccadically predicted and post-saccadically experienced position of the saccade target. However, recent research indicates that saccadic adaptation might be driven by postdictive motor error, i.e., a retrospective estimation of the pre-saccadic target position based on the post-saccadic image. We investigated whether oculomotor behavior can be adapted based on post-saccadic target information alone. We measured eye movements and localization judgements as participants aimed saccades at an initially invisible target, which was always shown only after the saccade. Each such trial was followed by either a pre- or a post-saccadic localization trial. The target position was fixed for the first 100 trials of the experiment and, during the following 200 trials, successively shifted inward or outward. Saccade amplitude and the pre- and post-saccadic localization judgements adjusted to the changing target position. Our results suggest that post-saccadic information is sufficient to induce error-reducing adaptive changes in saccade amplitude and target localization, possibly reflecting continuous updating of the estimated pre-saccadic target location driven by postdictive motor error.
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Saccadic eye movement
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Saccadic eye movements profoundly influence the perception of space: stimuli presented briefly around saccadic onset are perceived compressed towards the saccadic target. We studied perisaccadic mislocalization with a double-step saccade paradigm, an important technique in eye-movement research where the second saccade needs to be planned before the first has been executed, and must therefore take into account the displacement caused by the first. In our study the saccades were "memory-guided", with both saccadic targets extinguished before commencing the sequence. We measured perisaccadic localization of a small probe dot briefly flashed at various times during the sequence. At onset of the first saccade, probe dots were mislocalized towards the first and also the second saccade target. However, on onset of the second saccade, there was very little mislocalization. We reasoned that the lack of mislocalization could reflect failure to encode the location of the second saccadic target in an appropriate coordinate space (that takes into account the motion of the first saccade). Perhaps this encoding takes time? To test this idea, we increased the viewing duration of the saccade targets (before commencing the saccade sequence), and observed mislocalization at onset of both the first and second saccade. Our data suggest that construction of the spatiotopic representation requires at least 200 ms, a notion reinforced by a series of experiments on saccadic displacement of motion. We conclude that perisaccadic mislocalization towards the saccade target occurs only after neural map has been constructed in suitable coordinates, and the construction of this map requires time. Meeting abstract presented at VSS 2012
Saccadic eye movement
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Abstract The present study was designed to test whether the adaptation of saccadic eye movements depends only on the eye displacement vector of the trained saccade or also on eye position information. Using the double‐step target paradigm in eight human subjects, we first induced in a single session two “opposite directions adaptations” (ODA) of horizontal saccades of the same vector. Each ODA (backward or forward) was linked to one vertical eye position (12.5° up or 25° down) and alternated from trial to trial. The results showed that opposite changes of saccade amplitude can develop simultaneously, indicating that saccadic adaptation depends on orbital eye position. This finding has important functional implications because in everyday life our eyes saccade from constantly changing orbital positions. A comparison of these data to two control conditions in which training trials of a single type (backward or forward) were presented at both 12.5° and −25° eye elevations further indicated that eye position specificity is complete for backward, but not for forward, adaptation. Finally, the control conditions also indicated that the adaptation of a single saccade fully transferred to untrained saccades of the same vector, but initiated from different vertical eye positions. In conclusion, our study indicates that saccadic adaptation mechanisms use vectorial eye displacement signals, but can also take eye position signals into account as a contextual cue when the training involves conflicting saccade amplitude changes.
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The present study was designed to test whether the adaptation of saccadic eye movements depends only on the eye displacement vector of the trained saccade or also on eye position information. Using the double-step target paradigm in eight human subjects, we first induced in a single session two "opposite directions adaptations" (ODA) of horizontal saccades of the same vector. Each ODA (backward or forward) was linked to one vertical eye position (12.5 degrees up or 25 degrees down) and alternated from trial to trial. The results showed that opposite changes of saccade amplitude can develop simultaneously, indicating that saccadic adaptation depends on orbital eye position. This finding has important functional implications because in everyday life our eyes saccade from constantly changing orbital positions. A comparison of these data to two control conditions in which training trials of a single type (backward or forward) were presented at both 12.5 degrees and -25 degrees eye elevations further indicated that eye position specificity is complete for backward, but not for forward, adaptation. Finally, the control conditions also indicated that the adaptation of a single saccade fully transferred to untrained saccades of the same vector, but initiated from different vertical eye positions. In conclusion, our study indicates that saccadic adaptation mechanisms use vectorial eye displacement signals, but can also take eye position signals into account as a contextual cue when the training involves conflicting saccade amplitude changes.
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This article addresses questions about the preparatory processes that immediately precede saccadic eye movements. Saccade latencies were measured in a task in which subjects were provided partial advance information about the spatial location of a target fixation. In one experiment, subjects were faster in initiating saccades when they knew either the direction or amplitude of the required movement in advance compared to a condition with equal uncertainty about the number of potential saccade targets but without knowledge of the parameters required to execute the movement. These results suggest that the direction and amplitude for an upcoming saccade were calculated separately, and not in a fixed serial order. In another experiment, subjects appear to have programmed the saccades more holistically--with computations of direction and amplitude parameters occurring simultaneously. The implications of these results for models of eye movement preparation are discussed.
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Perception of our visual environment strongly depends on saccadic eye movements, which in turn are calibrated by saccadic adaptation mechanisms elicited by systematic movement errors. Current models of saccadic adaptation assume that visual error signals are acquired only after saccade completion, because the high speed of saccade execution disturbs visual processing (saccadic "suppression" and "mislocalization"). Complementing a previous study from our group, here we report that visual information presented during saccades can drive adaptation mechanisms and we further determine the critical time window of such error processing. In 15 healthy volunteers, shortening adaptation of reactive saccades toward a ±8° visual target was induced by flashing the target for 2 ms less eccentrically than its initial location either near saccade peak velocity ("PV" condition) or peak deceleration ("PD") or saccade termination ("END"). Results showed that, as compared to the "CONTROL" condition (target flashed at its initial location upon saccade termination), saccade amplitude decreased all throughout the "PD" and "END" conditions, reaching significant levels in the second adaptation and post-adaptation blocks. The results of nine other subjects tested in a saccade lengthening adaptation paradigm with the target flashing near peak deceleration ("PD" and "CONTROL" conditions) revealed no significant change of gain, confirming that saccade shortening adaptation is easier to elicit. Also, together with this last result, the stable gain observed in the "CONTROL" conditions of both experiments suggests that mislocalization of the target flash is not responsible for the saccade shortening adaptation demonstrated in the first group. Altogether, these findings reveal that the visual "suppression" and "mislocalization" phenomena related to saccade execution do not prevent brief visual information delivered "in-flight" from being processed to elicit oculomotor adaptation.
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Abstract Visual objects presented around the time of saccadic eye movements are strongly mislocalized towards the saccadic target, a phenomenon known as "saccadic compression." Here we show that perisaccadic compression is modulated by the presence of a visual saccadic target. When subjects saccaded to the center of the screen with no visible target, perisaccadic localization was more veridical than when tested with a target. Presenting a saccadic target sometime before saccade initiation was sufficient to induce mislocalization. When we systematically varied the onset of the saccade target, we found that it had to be presented around 100 ms before saccade execution to cause strong mislocalization: saccadic targets presented after this time caused progressively less mislocalization. When subjects made a saccade to screen center with a reference object placed at various positions, mislocalization was focused towards the position of the reference object. The results suggest that saccadic compression is a signature of a mechanism attempting to match objects seen before the saccade with those seen after.
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Saccades are used by the visual system to explore visual space with the high accuracy of the fovea. The visual error after the saccade is used to adapt the control of subsequent eye movements of the same amplitude and direction in order to keep saccades accurate. Saccadic adaptation is thus specific to saccade amplitude and direction. In the present study we show that saccadic adaptation is also specific to the initial position of the eye in the orbit. This is useful, because saccades are normally accompanied by head movements and the control of combined head and eye movements depends on eye position. Many parts of the saccadic system contain eye position information. Using the intrasaccadic target step paradigm, we adaptively reduced the amplitude of reactive saccades to a suddenly appearing target at a selective position of the eyes in the orbitae and tested the resulting amplitude changes for the same saccade vector at other starting positions. For central adaptation positions the saccade amplitude reduction transferred completely to eccentric starting positions. However, for adaptation at eccentric starting positions, there was a reduced transfer to saccades from central starting positions or from eccentric starting positions in the opposite hemifield. Thus eye position information modifies the transfer of saccadic amplitude changes in the adaptation of reactive saccades. A gain field mechanism may explain the eye position dependence found.
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