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Extraretinal information, such as corollary discharge (CD), is hypothesized to help compensate for saccade-induced visual input disruptions. However, support for this hypothesis is largely for one-dimensional transsaccadic visual changes, with little comprehensive information on the spatial characteristics. Here we systematically mapped the two-dimensional extent of this compensation by quantifying the insensitivity to different displacement metrics. Human subjects made saccades to targets positioned at different amplitudes (4° or 8°) and directions (rightward, oblique, or upward). After the saccade the initial target disappeared and, after a blank period, reappeared at a shifted location-- a collinear, diagonal, or orthogonal displacement. Subjects reported the perceived shift direction, and we determined the displacement detection based on the perceptual judgments. The two-dimensional insensitivity fields resulting from the perceptual thresholds had spatial features similar to the saccadic eye movement variability: 1) scaled with movement amplitude, 2) oriented (less sensitive to the change) along the saccade vector, and 3) approximately constant in shape when normalized by movement amplitude. In addition, comparing the postsaccadic perceptual estimate of the presaccadic target location to that based solely on the postsaccade visual error showed that overall the perceptual estimate was approximately 50% more accurate and 35% less variable than estimates based solely on this visual information. However, this relationship was not uniform: The benefit of extraretinal information was observed largely for displacements with a component parallel to the saccade vector. These results suggest a graded use of extraretinal information when forming the postsaccadic perceptual evaluation of transsaccadic environmental changes. [ABSTRACT FROM AUTHOR]

Saccadic eye movements direct the high-resolution foveae of our retinas toward objects of interest. With each saccade, the image jumps on the retina, causing a discontinuity in visual input. Our visual perception, however, remains stable. Philosophers and scientists over centuries have proposed that visual stability depends upon an internal neuronal signal that is a copy of the neuronal signal driving the eye movement, now referred to as a corollary discharge (CD) or efference copy. In the old world monkey, such a CD circuit for saccades has been identified extending from superior colliculus through MD thalamus to frontal cortex, but there is little evidence that this circuit actually contributes to visual perception. We tested the influence of this CD circuit on visual perception by first training macaque monkeys to report their perceived eye direction, and then reversibly inactivating the CD as it passes through the thalamus.Wefound that the monkey's perception changed; during CD inactivation, there was a difference between where the monkey perceived its eyes to be directed and where they were actually directed. Perception and saccade were decoupled. We established that the perceived eye direction at the end of the saccade was not derived from proprioceptive input from eye muscles, and was not altered by contextual visual information. We conclude that the CD provides internal information contributing to the brain's creation of perceived visual stability. More specifically, the CD might provide the internal saccade vector used to unite separate retinal images into a stable visual scene. [ABSTRACT FROM AUTHOR]

Corollary discharge (CD) is hypothesized to provide the movement information (direction and amplitude) required to compensate for the saccadeinduced disruptions to visual input. Here, we investigated to what extent these conveyed metrics influence perceptual stability in human subjects with a target-displacement detection task. Subjects made saccades to targets located at different amplitudes (4°, 6°, or 8°) and directions (horizontal or vertical). During the saccade, the target disappeared and then reappeared at a shifted location either in the same direction or opposite to the movement vector. Subjects reported the target displacement direction, and from these reports we determined the perceptual threshold for shift detection and estimate of target location. Our results indicate that the thresholds for all amplitudes and directions generally scaled with saccade amplitude. Additionally, subjects on average produced hypometric saccades with an estimated CD gain <1. Finally, we examined the contribution of different error signals to perceptual performance, the saccade error (movement-to-movement variability in saccade amplitude) and visual error (distance between the fovea and the shifted target location). Perceptual judgment was not influenced by the fluctuations in movement amplitude, and performance was largely the same across movement directions for different magnitudes of visual error. Importantly, subjects reported the correct direction of target displacement above chance level for very small visual errors (<0.75°), even when these errors were opposite the target-shift direction. Collectively, these results suggest that the CD-based compensatory mechanisms for visual disruptions are highly accurate and comparable for saccades with different metrics. [ABSTRACT FROM AUTHOR]

Before each saccade, neurons in frontal eye field anticipate the impending eye movement by showing sensitivity to stimuli appearing where the neuron's receptive field will be at the end of the saccade, referred to as the future field (FF) of the neuron. We explored the time course of this anticipatory activity in monkeys by briefly flashing stimuli in the FF at different times before saccades. Different neurons showed substantial variation in FF time course, but two salient observations emerged. First, when we compared the time span of stimulus probes before the saccade to the time span of FF activity, we found a striking temporal compression of FF activity, similar to compression seen for perisaccadic stimuli in human psychophysics. Second, neurons with distinct FF activity also showed suppression at the time of the saccade. The increase in FF activity and the decrease with suppression were temporally independent, making the patterns of activity difficult to separate. We resolved this by constructing a simple model with values for the start, peak, and duration of FF activity and suppression for each neuron. The model revealed the different time courses of FF sensitivity and suppression, suggesting that information about the impending saccade triggering suppression reaches the frontal eye field through a different pathway, or a different mechanism, than that triggering FF activity. Recognition of the variations in the time course of anticipatory FF activity provides critical information on its function and its relation to human visual perception at the time of the saccade. [ABSTRACT FROM AUTHOR]

Despite saccades changing the image on the retina several times per second, we still perceive a stable visual world. A possible mechanism underlying this stability is that an internal retinotopic map is updated with each saccade, with the location of objects being compared before and after the saccade. Psychophysical experiments have shown that humans derive such location information from a corollary discharge (CD) accompanying saccades. Such a CD has been identified in the monkey brain in a circuit extending from superior colliculus to frontal cortex. There is a missing piece, however. Perceptual localization is established only in humans and the CD circuit only in monkeys. We therefore extended measurement of perceptual localization to the monkey by adapting the target displacement detection task developed in humans. During saccades to targets, the target disappeared and then reappeared, sometimes at a different location. The monkeys reported the displacement direction. Detections of displacement were similar in monkeys and humans, but enhanced detection of displacement from blanking the target at the end of the saccade was observed only in humans, not in monkeys. Saccade amplitude varied across trials, but the monkey's estimates of target location did not follow that variation, indicating that eye location depended on an internal CD rather than external visual information. We conclude that monkeys use a CD to determine their new eye location after each saccade, just as humans do. [ABSTRACT FROM AUTHOR]

Ballistic movements like saccades require the brain to generate motor commands without the benefit of sensory feedback. Despite this, saccades are remarkably accurate. Theory suggests that this accuracy arises because the brain relies on an internal forward model that monitors the motor commands, predicts their sensory consequences, and corrects eye trajectory midflight. If control of saccades relies on a forward model, then the forward model should adapt whenever its predictions fail to match sensory feedback at the end of the movement. Using optimal feedback control theory, we predicted how this adaptation should alter saccade trajectories. We trained subjects on a paradigm in which the horizontal target jumped vertically during the saccade. With training, the final position of the saccade moved toward the second target. However, saccades became increasingly curved, i.e., suboptimal, as oculomotor commands were corrected on-line to steer the eye toward the second target. The adaptive response had two components: (1) the motor commands that initiated the saccades changed slowly, aiming the saccade closer to the jumped target. The adaptation of these earliest motor commands displayed little forgetting during the rest periods. (2) Late in saccade trajectory, another adaptive response steered it still closer to the jumped target, producing curvature. Adaptation of these late motor commands showed near-complete forgetting during the rest periods. The two components adapted at different timescales, with the late-acting component displaying much faster rates. It appears that in controlling saccades, the brain relies on an internal feedback that has the characteristics of a fast-adapting forward model. [ABSTRACT FROM AUTHOR]

The ability to make predictive saccadic eye movements is dependent on neural signals that anticipate the onset of a visual target. We used a novel paradigm—based on the saccade-countermanding task—as a tool to investigate rhythm saccade pacing and to provide information on the mechanisms of predictive timing. In particular, we examined the ability of normal subjects to stop a sequence of periodically paced eye movements when cued by a stop signal that was presented at different times with respect to the last target of the sequence (stop signal delay, SSD). The timing of the stop signal affected the ability to stop the saccadic sequence (make a saccade to a central target rather than to the peripheral alternating targets) in different ways, depending on the preceding tracking behavior. For the same SSD, subjects cancelled fewer trials during predictive tracking (promoted by tracking targets alternating at a fast pacing rate, 1.0 Hz) than during reactive tracking (tracking alternating targets at a low pacing rate, 0.2 Hz). In addition, on non-canceled trials, there was an increase in the delay of the corrective saccade to the central target with increasing SSD for pacing at 0.2 Hz, but the timing of the corrective saccade remained near constant for 1.0 Hz pacing. In examining the timing between movements, we estimate that the repetitive GO process that drives the saccades during predictive tracking begins earlier and has a shorter duration than the repetitive GO process during reactive tracking. These behavioral results provide further insight into the initiation process of predictive responses. In particular, the reduced reaction time and the corresponding short duration of the predictive process may result from a faster accumulation of neuronal discharge to a relatively fixed threshold. [ABSTRACT FROM AUTHOR]

Abstract: Previously we have shown that repetitive predictive saccades to alternating visual targets are mediated by an internal clock. That is, when subjects track a periodic visual stimulus alternating at a high rate (a small inter-stimulus interval, ISI, of 500 or 833ms), they use an internal estimate of stimulus timing to pre-program the eye movement timing. Auditory pacing tones at the same rate also generate predictive saccades. It is natural to ask if an identical internal clock is used to generate the predictive saccades in each case. We hypothesized that if subjects can use auditory information to establish an internal estimate of stimulus timing—as we demonstrated can be done with visual targets—then the distributions of predictive inter-saccade intervals should demonstrate the well-known “Scalar Property” for either Auditory Cued or Visual Cued stimuli: inter-saccade interval histograms should be almost identical when each is divided by its mean. However, when making reactive saccades to a pacing stimulus (at a low rate), there should be a difference in the timing statistics between Auditory and Visual pacing, due to differences in sensory processing. We report here that the variances of inter-saccade intervals at three predictive pacing rates (ISIs of 500, 833, and 1000ms) are equivalent, whereas the variance for Auditory Cued Pacing was greater than that for Visual Cued Pacing during reactive saccades at two reactive pacing rates (ISIs of 1667 and 2500ms). When the inter-saccade interval histograms at the predictive pacing rates were normalized, the distributions were nearly identical for both Visual and Auditory Cued Pacing, which means that the Scalar Property holds for predictive saccades from either pacing stimulus. These results suggest that (1) an internal timing reference (clock) can be established by either auditory or visual information and (2) during predictive tracking the variability in saccade timing is due to the variability in the internal timing representation, while during reactive tracking the variability in saccade timing depends on the sensory modality used to trigger the saccades. [Copyright &y& Elsevier]

Previously we demonstrated the presence of a behavioral phase transition between reactive and predictive eye tracking of alternating targets. Prior studies of repetitive movements have proposed that an “internal clock” is the neural mechanism by which interval timing is achieved. In the present report we tested whether predictive oculomotor (saccade) tracking is based on an internal time reference (clock) by examining the effect of transient perturbations to the periodic pacing stimulus. These perturbations consisted of altering the timing of the stimulus (abruptly increasing or decreasing the inter-stimulus interval) or extinguishing the targets altogether. Although reactive tracking (at low pacing rates) was greatly affected by these timing perturbations, once predictive tracking was established subjects continued to time their eye movement responses at the pre-existing rate despite the perturbation. As expected from certain clock models, inter-stimulus intervals for predictive tracking followed Weber’s law and the scalar property (timing variability increases in proportion to interval duration), but this was not true for reactive tracking. In addition, the perturbation results show that subjects can establish an internal representation of target pacing (the internal clock) in as little as two eye-movement intervals, which suggests that this mechanism is relevant for real-world situations. These findings are consistent with the presence of an internal clock for the generation of these predictive movements, and demonstrate that the neural mechanism responsible for this behavior is temporally accurate and flexible. [ABSTRACT FROM AUTHOR]

Data from previous human and primate studies on saccadic and smooth pursuit eye movements suggest that there are shared internal inputs (for example, perception, attention, expectation, and memory) for the initiation of the two types of movements. Additional reports examining the effect of preparation time on movement responses have shown that when ample time is allowed subjects usually generate long-latency “reactive” responses. When the time allowed to prepare a movement is short, however, subjects respond with reduced latency and often anticipate the stimulus (“predictive” response). Based on these findings, we believe that the shared internal inputs at early stages of movement preparation may result in saccade and pursuit eye movements demonstrating the same dependence on preparation time despite acting through different neural pathways further downstream. Previously we demonstrated a behavioral “phase transition” when normal subjects tracked alternating targets with saccades. When preparation time was long (low-frequency pacing) subjects made reactive saccades (latency ~180 ms). As preparation time monotonically decreased (pacing frequency increased), there was an abrupt transition to a predictive response (latency <100 ms). In the present study we show that a similar transition exists in smooth pursuit tracking and that the point of transition between the two behaviors is the same for both systems. In other words, the same behavior (reactive versus predictive) is selected when pursuit and saccade tracking are tested under the same time constraints. This provides further evidence that the two types of movements are different motor outcomes of a common decision process. [ABSTRACT FROM AUTHOR]

The cerebellum plays an important role in predicting and producing regularly timed motor responses. In the present report, we examined the performance of SCA6 cerebellar patients and normal controls in a simple saccade tracking task. Our results imply that the cerebellum plays a role in maintaining predictive motor behavior once it has been established. [ABSTRACT FROM AUTHOR]

The double-step paradigm investigates the characteristics of planning and execution when the motor system must rapidly adjust for a new goal location. Studies have provided detailed temporal information based on the duration available for the motor system to prepare a new movement trajectory (here referred to as re-preparation time). However, previous work has largely examined single displacement sizes, limiting the spatiotemporal understanding of movement planning and execution. The lack of a description of this behavioral timecourse across increasing displacement sizes is true for saccades, rapid eye movements that redirect the fovea. Furthermore, during the double-step paradigm, the primary saccade often fails to accurately foveate the final target location and a secondary saccade brings the target onto the fovea. However, it is also unknown how this compensation is concurrently modified with the exposure duration and displacement of the movement goal. Here, we examined the amount of time required to change the initial saccade direction to a new target location for relatively small (20°, 30°, and 40°) and large (60° and 90°) target spatial separations. Interestingly, we found a clear relationship between the saccade direction and the amount of time allowed to redirect the movement; across separations, intermediate saccades occurred when approximately 60-140 ms was available to readjust the movement plan. Additionally, there was a consistent relationship between the timing of the secondary saccade and the re-preparation time across jump sizes, suggesting that concurrent movement correction planning was dependent on the amount of exposure to the final movement goal. [ABSTRACT FROM AUTHOR]