The Influence of Briefly Presented Randomized Target Motion on the Extraretinal Component of Ocular Pursuit

2008 ◽  
Vol 99 (2) ◽  
pp. 831-842 ◽  
Author(s):  
G. R. Barnes ◽  
C. J. S. Collins

We assessed the ability to extract velocity information from brief exposure of a moving target and sought evidence that this information could be used to modulate the extraretinal component of ocular pursuit. A step-ramp target motion was initially visible for a brief randomized period of 50, 100, 150, or 200 ms, but then extinguished for a randomized period of 400 or 600 ms before reappearing and continuing along its trajectory. Target speed (5–20°/s), direction (left/right), and intertrial interval (2.7–3.7 s) were also randomized. Smooth eye movements were initiated after about 130 ms and comprised an initial visually dependent component, which reached a peak velocity that increased with target velocity and initial exposure duration, followed by a sustained secondary component that actually increased throughout extinction for 50- and 100-ms initial exposures. End-extinction eye velocity, reflecting extraretinal drive, increased with initial exposure from 50 to 100 ms but remained similar for longer exposures; it was significantly scaled to target velocity for 150- and 200-ms exposures. The results suggest that extraretinal drive is based on a sample of target velocity, mostly acquired during the first 150 ms, that is stored and forms a goal for generating appropriately scaled eye movements during absence of visual input. End-extinction eye velocity was significantly higher when target reappearance was expected than when it was not, confirming the importance of expectation in generating sustained smooth movement. However, end-extinction eye displacement remained similar irrespective of expectation, suggesting that the ability to use sampled velocity information to predict future target displacement operates independently of the control of smooth eye movement.

2006 ◽  
Vol 16 (1-2) ◽  
pp. 1-22 ◽  
Author(s):  
Junko Fukushima ◽  
Teppei Akao ◽  
Sergei Kurkin ◽  
Chris R.S. Kaneko ◽  
Kikuro Fukushima

In order to see clearly when a target is moving slowly, primates with high acuity foveae use smooth-pursuit and vergence eye movements. The former rotates both eyes in the same direction to track target motion in frontal planes, while the latter rotates left and right eyes in opposite directions to track target motion in depth. Together, these two systems pursue targets precisely and maintain their images on the foveae of both eyes. During head movements, both systems must interact with the vestibular system to minimize slip of the retinal images. The primate frontal cortex contains two pursuit-related areas; the caudal part of the frontal eye fields (FEF) and supplementary eye fields (SEF). Evoked potential studies have demonstrated vestibular projections to both areas and pursuit neurons in both areas respond to vestibular stimulation. The majority of FEF pursuit neurons code parameters of pursuit such as pursuit and vergence eye velocity, gaze velocity, and retinal image motion for target velocity in frontal and depth planes. Moreover, vestibular inputs contribute to the predictive pursuit responses of FEF neurons. In contrast, the majority of SEF pursuit neurons do not code pursuit metrics and many SEF neurons are reported to be active in more complex tasks. These results suggest that FEF- and SEF-pursuit neurons are involved in different aspects of vestibular-pursuit interactions and that eye velocity coding of SEF pursuit neurons is specialized for the task condition.


2012 ◽  
Vol 5 (3) ◽  
Author(s):  
Graham R. Barnes

When humans pursue sinusoidal target motion they rapidly learn to track with minimal phase error despite inherent visuo-motor processing delays; prior evidence suggests that prediction might even occur within the first cycle. Here, this has been examined by evoking reactive responses to single cycle stimuli having randomised periodicity and peak velocity. Periodicity was varied within three specific ranges with differing average periodicity. Initial responses in the first half-cycle were remarkably similar within periodicity ranges, irrespective of target velocity or frequency, but differed between ranges. In contrast, in the second half-cycle eye velocity closely matched the target in velocity and timing, irrespective of differences in eye velocity in the first half. Abrupt transitions occurred between first and second half-cycles, consistent with the hypothesis that target motion information is sampled and stored within the first half-cycle, irrespective of actual eye velocity evoked, and then released as a predictive estimate in the second half.


1992 ◽  
Vol 67 (3) ◽  
pp. 625-638 ◽  
Author(s):  
D. Goldreich ◽  
R. J. Krauzlis ◽  
S. G. Lisberger

1. Our goal was to discriminate between two classes of models for pursuit eye movements. The monkey's pursuit system and both classes of model exhibit oscillations around target velocity during tracking of ramp target motion. However, the mechanisms that determine the frequency of oscillations differ in the two classes of model. In "internal feedback" models, oscillations are controlled by internal feedback loops, and the frequency of oscillation does not depend strongly on the delay in visual feedback. In "image motion" models, oscillations are controlled by visual feedback, and the frequency of oscillation does depend on the delay in visual feedback. 2. We measured the frequency of oscillation during pursuit of ramp target motion as a function of the total delay for visual feedback. For the shortest feedback delays of approximately 70 ms, the frequency of oscillation was between 6 and 7 Hz. Increases in feedback delay caused decreases in the frequency of oscillation. The effect of increasing feedback delay was similar, whether the increases were produced naturally by dimming and decreasing the size of the tracking target or artificially with the computer. We conclude that the oscillations in eye velocity during pursuit of ramp target motion are controlled by visual inputs, as suggested by the image motion class of models. 3. Previous experiments had suggested that the visuomotor pathways for pursuit are unable to respond well to frequencies as high as the 6-7 Hz at which eye velocity oscillates in monkeys. We therefore tested the response to target vibration at an amplitude of +/- 8 degrees/s and frequencies as high as 15 Hz. For target vibration at 6 Hz, the gain of pursuit, defined as the amplitude of eye velocity divided by the amplitude of target velocity, was as high as 0.65. We conclude that the visuomotor pathways for pursuit are capable of processing image motion at high temporal frequencies. 4. The gain of pursuit was much larger when the target vibrated around a constant speed of 15 degrees/s than when it vibrated around a stationary position. This suggests that the pursuit pathways contain a switch that must be closed to allow the visuomotor pathways for pursuit to operate at their full gain. The switch apparently remains open for target vibration around a stationary position. 5. The responses to target vibration revealed a frequency at which eye velocity lagged target velocity by 180 degrees and at which one monkey showed a local peak in the gain of pursuit.(ABSTRACT TRUNCATED AT 400 WORDS)


1998 ◽  
Vol 79 (4) ◽  
pp. 1918-1930 ◽  
Author(s):  
Stephen G. Lisberger

Lisberger, Stephen G. Postsaccadic enhancement of initiation of smooth pursuit eye movements in monkeys. J. Neurophysiol. 79: 1918–1930, 1998. Step-ramp target motion evokes a characteristic sequence of presaccadic smooth eye movement in the direction of the target ramp, catch-up targets to bring eye position close to the position of the moving target, and postsaccadic eye velocities that nearly match target velocity. I have analyzed this sequence of eye movements in monkeys to reveal a strong postsaccadic enhancement of pursuit eye velocity and to document the conditions that lead to that enhancement. Smooth eye velocity was measured in the last 10 ms before and the first 10 ms after the first saccade evoked by step-ramp target motion. Plots of eye velocity as a function of time after the onset of the target ramp revealed that eye velocity at a given time was much higher if measured after versus before the saccade. Postsaccadic enhancement of pursuit was recorded consistently when the target stepped 3° eccentric on the horizontal axis and moved upward, downward, or away from the position of fixation. To determine whether postsaccadic enhancement of pursuit was invoked by smear of the visual scene during a saccade, I recorded the effect of simulated saccades on the presaccadic eye velocity for step-ramp target motion. The 3° simulated saccade, which consisted of motion of a textured background at 150°/s for 20 ms, failed to cause any enhancement of presaccadic eye velocity. By using a strategically selected set of oblique target steps with horizontal ramp target motion, I found clear enhancement for saccades in all directions, even those that were orthogonal to target motion. When the size of the target step was varied by up to 15° along the horizontal meridian, postsaccadic eye velocity did not depend strongly either on the initial target position or on whether the target moved toward or away from the position of fixation. In contrast, earlier studies and data in this paper show that presaccadic eye velocity is much stronger when the target is close to the center of the visual field and when the target moves toward versus away from the position of fixation. I suggest that postsaccadic enhancement of pursuit reflects activation, by saccades, of a switch that regulates the strength of transmission through the visual-motor pathways for pursuit. Targets can cause strong visual motion signals but still evoke low presaccadic eye velocities if they are ineffective at activating the pursuit system.


1994 ◽  
Vol 72 (6) ◽  
pp. 2714-2728 ◽  
Author(s):  
A. F. Fuchs ◽  
F. R. Robinson ◽  
A. Straube

1. We recorded single-unit activity from neurons of an output of the cerebellum, the fastigial nucleus, in two rhesus macaques while the monkeys tracked small moving targets with their eyes. Many neurons in the caudal part of the fastigial nucleus exhibited a modulation in their discharge rates when smooth-pursuit eye movements were elicited by either sinusoidal or step-ramp motions of a small target. 2. The pursuit direction that elicited the most vigorous modulation in unit firing to sinusoidal target motion could be horizontal, vertical, or oblique. Most often, the preferred direction was in the contralateral and/or downward direction (50 of 69 neurons) or in the ipsilateral and/or upward direction (13 of 69). 3. For units whose preferred smooth-pursuit directions were either contralateral/downward or ipsilateral/upward during sinusoidal pursuit, peak firing as measured by the phase shift of periodic modulation at 0.5-0.8 Hz occurred near the time of peak velocity. The discharge of 80% of the neurons with contralateral/downward preferred directions preceded eye velocity by an average of -27 degrees; thus these neurons discharged maximally during eye acceleration. In contrast, neurons with ipsilateral/upward preferred directions lagged peak velocity by an average of +10.5 degrees and therefore discharged during eye deceleration. 4. The average eye velocity sensitivity for sinusoidal pursuit between 0.5 and 0.8 Hz was 0.83 +/- 0.57 (SD) spikes/s per degrees/s. We also tested 36 units during pursuit at a variety of frequencies in their preferred directions and found that firing rates increased monotonically with peak eye velocity. However, the firing rate saturated at velocities ranging from 20 to 60 degrees/s for different units. 5. When a monkey tracked a step-ramp target motion, three discharge patterns emerged in the 27 units tested. Just over half of the units discharged a burst of spikes that preceded (average lead of 27.4 +/- 17 ms) and lasted throughout the initial third of the eye acceleration; the burst was followed by a subsequent steady firing that continued after the eye had accelerated to its steady velocity. Fewer neurons discharged a burst that began late in the acceleration and was followed by steady firing. Occasional neurons showed only a gradual increase in firing rate during acceleration followed by a steady discharge. 6. Thirty of the 31 fastigial smooth-pursuit units tested also were modulated during sinusoidal yaw and/or pitch oscillations while the animals fixated a spot that rotated with them.(ABSTRACT TRUNCATED AT 400 WORDS)


2010 ◽  
Vol 104 (5) ◽  
pp. 2850-2862 ◽  
Author(s):  
Yan Yang ◽  
Stephen G. Lisberger

We commonly think of motor learning as a gradual process that makes small, adaptive steps in a consistent direction. We now report evidence that learning in pursuit eye movements could start with large, transient short-term alterations that stoke a more gradual long-term process. Monkeys tracked a target that started moving horizontally or vertically. After 250 ms of motion had produced a preinstruction eye velocity close to target velocity, an orthogonal component of target motion created an instructive change in target direction that was randomly in one of the two directions along the orthogonal axis. The preinstruction eye velocity in each trial expressed single-trial learning as a bias toward the direction of the instruction in the prior trial. The single-trial learning was forgotten within 4 to 10 s. Two observations implied that single-trial learning was not simply cognitive anticipation. First, the magnitude of the trial-over-trial change in eye velocity depended on the ongoing eye velocity at the time of the instruction in the prior trial. Single-trial learning was negligible if the prior trial had provided a well-timed cue without evoking any preinstruction eye velocity. Second, regular alternation of the direction of the instructive target motion caused reactive rather than anticipatory trial-over-trial changes in eye velocity. Humans showed very different responses that appeared to be based on cognitive anticipation rather than learning. We suggest that single-trial learning results from a low-level learning mechanism and may be a necessary prerequisite for longer-term modifications that are more permanent.


1990 ◽  
Vol 63 (4) ◽  
pp. 676-688 ◽  
Author(s):  
S. G. Lisberger

1. Monkeys normally use a combination of smooth head and eye movements to keep the eyes pointed at a slowly moving object. The visual inputs from target motion evoke smooth pursuit eye movements, whereas the vestibular inputs from head motion evoke a vestibuloocular reflex (VOR). Our study asks how the eye movements of pursuit and the VOR interact. Is there a linear addition of independent commands for pursuit and the VOR? Or does the interaction of visual and vestibular stimuli cause momentary, "parametric" modulation of transmission through VOR pathways? 2. We probed for the state of the VOR and pursuit by presenting transient perturbations of target and/or head motion under different steady-state tracking conditions. Tracking conditions included fixation at straight-ahead gaze, in which both the head and the target were stationary; "times-zero (X0) tracking," in which the target and head moved in the same direction at the same speed; and "times-two (X2) tracking," in which the target and head moved in opposite directions at the same speed. 3. Comparison of the eye velocities evoked by changes in the direction of X0 versus X2 tracking revealed two components of the tracking response. The earliest component, which we attribute to the VOR, had a latency of 14 ms and a trajectory that did not depend on initial tracking conditions. The later component had a latency of 70 ms or less and a trajectory that did depend on tracking conditions. 4. To probe the latency of pursuit eye movements, we imposed perturbations of target velocity imposed during X0 and X2 tracking. The resulting changes in eye velocity had latencies of at least 100 ms. We conclude that the effects of initial tracking conditions on eye velocity at latencies of less than 70 ms cannot be caused by visual feedback through the smooth-pursuit system. Instead, there must be another mechanism for short-latency control over the VOR; we call this component of the response "short-latency tracking." 5. Perturbations of head velocity or head and target velocity during X0 and X2 tracking showed that short-latency tracking depended only on the tracking conditions at the time the perturbation was imposed. The VOR appeared to be suppressed when the initial conditions were X0 tracking. 6. The magnitude of short-latency tracking depended on the speed of initial head and target movement. During X0 tracking at 15 deg/s, short-latency tracking was modest. When the initial speed of head and target motion was 60 deg/s, the amplitude of short-latency tracking was quite large and its latency became as short as 36 ms.(ABSTRACT TRUNCATED AT 400 WORDS)


2008 ◽  
Vol 100 (3) ◽  
pp. 1544-1556 ◽  
Author(s):  
Carlos R. Cassanello ◽  
Abhay T. Nihalani ◽  
Vincent P. Ferrera

Due to delays in visuomotor processing, eye movements directed toward moving targets must integrate both target position and velocity to be accurate. It is unknown where and how target velocity information is incorporated into the planning of rapid (saccadic) eye movements. We recorded the activity of neurons in frontal eye fields (FEFs) while monkeys made saccades to stationary and moving targets. A substantial fraction of FEF neurons was found to encode not only the initial position of a moving target, but the metrics (amplitude and direction) of the saccade needed to intercept the target. Many neurons also encoded target velocity in a nearly linear manner. The quasi-linear dependence of firing rate on target velocity means that the neuronal response can be directly read out to compute the future position of a target moving with constant velocity. This is demonstrated using a quantitative model in which saccade amplitude is encoded in the population response of neurons tuned to retinal target position and modulated by target velocity.


2006 ◽  
Vol 95 (2) ◽  
pp. 593-601 ◽  
Author(s):  
Matthew T. Avila ◽  
L. Elliot Hong ◽  
Amanda Moates ◽  
Kathleen A. Turano ◽  
Gunvant K. Thaker

Schizophrenia patients exhibit several smooth pursuit abnormalities including poor pursuit initiation. Velocity discrimination is also impaired and is correlated with pursuit initiation performance—suggesting that pursuit deficits are related to impairments in processing velocity information. Studies suggest that pursuit initiation is influenced by prior target motion information and/or expectations and that this is likely caused by expectation-based changes in the perceptual inputs to the pursuit system. We examined whether poor pursuit initiation in schizophrenia results from inaccurate encoding of immediate velocity signals, or whether these deficits reflect a failure to use prior target motion information to “optimize” the response. Twenty-eight patients and 24 controls performed an adapted version of a “remembered pursuit task.” Trials consisted of a series of target motions, the first of which occurred unexpectedly, followed by four to seven identical targets each preceded by an auditory cue and a “catch target” in which a cue was given followed by target extinction. Initiation eye velocity in response to unexpected, first targets was similar in the patient and control groups. In contrast, patients showed lower eye velocity in response to repeated, cued targets compared with controls. Patients also showed reduced eye velocity in response to catch targets. Reduction in pursuit latency across repeated targets was less robust in patients. Results suggest that processing of immediate velocity information is unaffected in schizophrenia and that pursuit initiation deficits reflect an inability to accurately generate, store, and/or access “remembered” velocity signals.


2003 ◽  
Vol 90 (4) ◽  
pp. 2205-2218 ◽  
Author(s):  
Mark M. Churchland ◽  
I-Han Chou ◽  
Stephen G. Lisberger

We recorded the smooth-pursuit eye movements of monkeys in response to targets that were extinguished (blinked) for 200 ms in mid-trajectory. Eye velocity declined considerably during the target blinks, even when the blinks were completely predictable in time and space. Eye velocity declined whether blinks were presented during steady-state pursuit of a constant-velocity target, during initiation of pursuit before target velocity was reached, or during eye accelerations induced by a change in target velocity. When a physical occluder covered the trajectory of the target during blinks, creating the impression that the target moved behind it, the decline in eye velocity was reduced or abolished. If the target was occluded once the eye had reached target velocity, pursuit was only slightly poorer than normal, uninterrupted pursuit. In contrast, if the target was occluded during the initiation of pursuit, while the eye was accelerating toward target velocity, pursuit during occlusion was very different from normal pursuit. Eye velocity remained relatively stable during target occlusion, showing much less acceleration than normal pursuit and much less of a decline than was produced by a target blink. Anticipatory or predictive eye acceleration was typically observed just prior to the reappearance of the target. Computer simulations show that these results are best understood by assuming that a mechanism of eye-velocity memory remains engaged during target occlusion but is disengaged during target blinks.


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