Oculomotor behavior can be adjusted on the basis of artificial feedback signals indicating externally caused errors.

Whether a saccade is accurate and has reached the target cannot be evaluated during its execution, but relies on post-saccadic feedback. If the eye has missed the target object, a secondary corrective saccade has to be made to align the fovea with the target. If a systematic post-saccadic error occurs, adaptive changes to the oculomotor behavior are made, such as shortening or lengthening the saccade amplitude. Systematic post-saccadic errors are typically attributed internally to erroneous motor commands. The corresponding adaptive changes to the motor command reduce the error and the need for secondary corrective saccades, and, in doing so, restore accuracy and efficiency. However, adaptive changes to the oculomotor behavior also occur if a change in saccade amplitude is beneficial for task performance, or if it is rewarded. Oculomotor learning thus is more complex than reducing a post-saccadic position error. In the current study, we used a novel oculomotor learning paradigm and investigated whether human participants are able to adapt their oculomotor behavior to improve task performance even when they attribute the error externally. The task was to indicate the intended target object among several objects to a simulated human-machine interface by making eye movements. The participants were informed that the system itself could make errors. The decoding process depended on a distorted landing point of the saccade, resulting in decoding errors. Two different types of visual feedback were added to the post-saccadic scene and we compared how participants used the different feedback types to adjust their oculomotor behavior to avoid errors. We found that task performance improved over time, regardless of the type of feedback. Thus, error feedback from the simulated human-machine interface was used for post-saccadic error evaluation. This indicates that 1) artificial visual feedback signals and 2) externally caused errors might drive adaptive changes to oculomotor behavior.

Adaptive changes to saccade amplitude and target localization do not require pre-saccadic target visibility.

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.

Mislocalization after inhibition of saccadic adaptation.

Saccadic eye movements are often imprecise and result in an error between expected and actual retinal target location after the saccade. Repeated experience of this error produces changes in saccade amplitude to reduce the error and concomitant changes in apparent visual location. We investigated the relationship between these two plastic processes in a series of experiments. Following a recent paradigm of inhibition of saccadic adaptation, in which participants are instructed to look at the initial target position and to continue to look at that position even if the target were to move again, our participants nevertheless perceived a visual probe presented near the saccade target to be shifted in direction of the target error. The location percept of the target gradually shifted and diverged over time from the executed saccade. Our findings indicate that changes in perceived location can be the same even when changes in saccade amplitude differ according to instruction and can develop even when the amplitude of the saccades executed during the adaptation procedure does not change. There are two possible explanations for this divergence between the adaptation states of saccade amplitude and perceived location. Either the intrasaccadic target step might trigger updating of the association between pre- and post-saccadic target positions, causing the localization shift, or the saccade motor command adjusts together with the perceived location at a common adaptation site, downstream from which voluntary control is exerted upon the executed eye movement only.

Flexible use of post-saccadic visual feedback in oculomotor learning.

Saccadic eye movements bring objects of interest onto our fovea. These gaze shifts are essential for visual perception of our environment and the interaction with the objects within it. They precede our actions and are thus modulated by current goals. It is assumed that saccadic adaptation, a recalibration process that restores saccade accuracy in case of error, is mainly based on an implicit comparison of expected and actual post-saccadic position of the target on the retina. However, there is increasing evidence that task demands modulate saccade adaptation and that errors in task performance may be sufficient to induce changes to saccade amplitude. We investigated if human participants are able to flexibly use different information sources within the post-saccadic visual feedback in task-dependent fashion. Using intra-saccadic manipulation of the visual input, participants were either presented with congruent post-saccadic information, indicating the saccade target unambiguously, or incongruent post-saccadic information, creating conflict between two possible target objects. Using different task instructions, we found that participants were able to modify their saccade behavior such that they achieved the goal of the task. They succeeded in decreasing saccade gain or maintaining it, depending on what was necessary for the task, irrespective of whether the post-saccadic feedback was congruent or incongruent. It appears that action intentions prime task-relevant feature dimensions and thereby facilitated the selection of the relevant information within the post-saccadic image. Thus, participants use post-saccadic feedback flexibly, depending on their intentions and pending actions.

Volitional control of saccadic adaptation.

Saccadic adaptation is assumed to be driven by an unconscious and automatic mechanism. We wondered if the adaptation process is accessible to volitional control, specifically whether any change in saccade gain can be inhibited. Participants were exposed to post-saccadic error by using the double-step paradigm in which a target is presented in a peripheral location and then stepped during the saccade to another location. In one condition, participants were instructed to follow the target step and look at the final target location. In the other condition they were instructed to inhibit the adjustment of saccade amplitude and look at the initial target location. We conducted two experiments, which differed in the size of the intra-saccadic target step. We found that when told to inhibit amplitude adjustment, gain change was close to zero for outward steps, but some adaptation remained for inward steps. Saccadic latency was not affected by the instruction type for inward steps, but when the target was stepped outward, latencies were longer in the inhibition than in the adaptation condition. The results show that volitional control can be exerted on saccadic adaptation. We suggest that volitional control affects the remapping of the target, thus having a larger impact on outward adaptation.