Saccadic Compensation for Smooth Eye and Head Movements During Head-Unrestrained Two-Dimensional Tracking

2010 ◽  
Vol 103 (1) ◽  
pp. 543-556 ◽  
Author(s):  
P. M. Daye ◽  
G. Blohm ◽  
P. Lefèvre

Spatial updating is the ability to keep track of the position of world-fixed objects while we move. In the case of vision, this phenomenon is called spatial constancy and has been studied in head-restraint conditions. During head-restrained smooth pursuit, it has been shown that the saccadic system has access to extraretinal information from the pursuit system to update the objects' position in the surrounding environment. However, during head-unrestrained smooth pursuit, the saccadic system needs to keep track of three different motor commands: the ocular smooth pursuit command, the vestibuloocular reflex (VOR), and the head movement command. The question then arises whether saccades compensate for these movements. To address this question, we briefly presented a target during sinusoidal head-unrestrained smooth pursuit in darkness. Subjects were instructed to look at the flash as soon as they saw it. We observed that subjects were able to orient their gaze to the memorized (and spatially updated) position of the flashed target generally using one to three successive saccades. Similar to the behavior in the head-restrained condition, we found that the longer the gaze saccade latency, the better the compensation for intervening smooth gaze displacements; after about 400 ms, 62% of the smooth gaze displacement had been compensated for. This compensation depended on two independent parameters: the latency of the saccade and the eye contribution to the gaze displacement during this latency period. Separating gaze into eye and head contributions, we show that the larger the eye contribution to the gaze displacement, the better the overall compensation. Finally, we found that the compensation was a function of the head oscillation frequency and we suggest that this relationship is linked to the modulation of VOR gain. We conclude that the general mechanisms of compensation for smooth gaze displacements are similar to those observed in the head-restrained condition.

2005 ◽  
Vol 93 (3) ◽  
pp. 1223-1234 ◽  
Author(s):  
Daniel J. Tollin ◽  
Luis C. Populin ◽  
Jordan M. Moore ◽  
Janet L. Ruhland ◽  
Tom C. T. Yin

In oculomotor research, there are two common methods by which the apparent location of visual and/or auditory targets are measured, saccadic eye movements with the head restrained and gaze shifts (combined saccades and head movements) with the head unrestrained. Because cats have a small oculomotor range (approximately ±25°), head movements are necessary when orienting to targets at the extremes of or outside this range. Here we tested the hypothesis that the accuracy of localizing auditory and visual targets using more ethologically natural head-unrestrained gaze shifts would be superior to head-restrained eye saccades. The effect of stimulus duration on localization accuracy was also investigated. Three cats were trained using operant conditioning with their heads initially restrained to indicate the location of auditory and visual targets via eye position. Long-duration visual targets were localized accurately with little error, but the locations of short-duration visual and both long- and short-duration auditory targets were markedly underestimated. With the head unrestrained, localization accuracy improved substantially for all stimuli and all durations. While the improvement for long-duration stimuli with the head unrestrained might be expected given that dynamic sensory cues were available during the gaze shifts and the lack of a memory component, surprisingly, the improvement was greatest for the auditory and visual stimuli with the shortest durations, where the stimuli were extinguished prior to the onset of the eye or head movement. The underestimation of auditory targets with the head restrained is explained in terms of the unnatural sensorimotor conditions that likely result during head restraint.


Author(s):  
Kerstin Rosander

Gaze control involves eyes, head, and body movements and is guided by mainly three types of information: visual, vestibular, and proprioceptive. Appropriate gaze control is a basis for actions such as reaching, grasping, eating, and manipulation, all of which develop during the first year of life. The development of gaze control is about how young infants gain access to these different kinds of information, how they come to use them, and how they come to coordinate head and eyes to accomplish it. This control develops during the first few weeks of life. A major challenge for the gaze controlling system is how gaze is stabilized on a moving target to keep vision clear, including during self-motion or the compensation of other sudden movements. Furthermore, the tracking has to be timed relative to the object motion. This requires prediction, which is a part of smooth pursuit that emerges at around six weeks and is in full function at three months. The smooth eye and head movements must add up in time and space to the object motion. Then the vestibular and visual neural signals must be properly added. Catch-up saccades compensate when the smooth pursuit is insufficient. In other situations, saccades shift the gaze between objects or situations. Moreover, if a moving object temporarily disappears out of view, one or several saccades predictively recapture the object at the reappearance position (four months). The complex and fast development of gaze has inspired the design of robotic vision (iCub) through processes similar to human development, thus increasing the robot’s flexibility and learning abilities


2008 ◽  
Vol 100 (4) ◽  
pp. 1848-1867 ◽  
Author(s):  
Sigrid M. C. I. van Wetter ◽  
A. John van Opstal

Such perisaccadic mislocalization is maximal in the direction of the saccade and varies systematically with the target-saccade onset delay. We have recently shown that under head-fixed conditions perisaccadic errors do not follow the quantitative predictions of current visuomotor models that explain these mislocalizations in terms of spatial updating. These models all assume sluggish eye-movement feedback and therefore predict that errors should vary systematically with the amplitude and kinematics of the intervening saccade. Instead, we reported that errors depend only weakly on the saccade amplitude. An alternative explanation for the data is that around the saccade the perceived target location undergoes a uniform transient shift in the saccade direction, but that the oculomotor feedback is, on average, accurate. This “ visual shift” hypothesis predicts that errors will also remain insensitive to kinematic variability within much larger head-free gaze shifts. Here we test this prediction by presenting a brief visual probe near the onset of gaze saccades between 40 and 70° amplitude. According to models with inaccurate gaze-motor feedback, the expected perisaccadic errors for such gaze shifts should be as large as 30° and depend heavily on the kinematics of the gaze shift. In contrast, we found that the actual peak errors were similar to those reported for much smaller saccadic eye movements, i.e., on average about 10°, and that neither gaze-shift amplitude nor kinematics plays a systematic role. Our data further corroborate the visual origin of perisaccadic mislocalization under open-loop conditions and strengthen the idea that efferent feedback signals in the gaze-control system are fast and accurate.


1995 ◽  
Vol 73 (4) ◽  
pp. 1632-1652 ◽  
Author(s):  
J. O. Phillips ◽  
L. Ling ◽  
A. F. Fuchs ◽  
C. Siebold ◽  
J. J. Plorde

1. We studied horizontal eye and head movements in three monkeys that were trained to direct their gaze (eye position in space) toward jumping targets while their heads were both fixed and free to rotate about a vertical axis. We considered all gaze movements that traveled > or = 80% of the distance to the new visual target. 2. The relative contributions and metrics of eye and head movements to the gaze shift varied considerably from animal to animal and even within animals. Head movements could be initiated early or late and could be large or small. The eye movements of some monkeys showed a consistent decrease in velocity as the head accelerated, whereas others did not. Although all gaze shifts were hypometric, they were more hypometric in some monkeys than in others. Nevertheless, certain features of the gaze shift were identifiable in all monkeys. To identify those we analyzed gaze, eye in head position, and head position, and their velocities at three points in time during the gaze shift: 1) when the eye had completed its initial rotation toward the target, 2) when the initial gaze shift had landed, and 3) when the head movement was finished. 3. For small gaze shifts (< 20 degrees) the initial gaze movement consisted entirely of an eye movement because the head did not move. As gaze shifts became larger, the eye movement contribution saturated at approximately 30 degrees and the head movement contributed increasingly to the initial gaze movement. For the largest gaze shifts, the eye usually began counterrolling or remained stable in the orbit before gaze landed. During the interval between eye and gaze end, the head alone carried gaze to completion. Finally, when the head movement landed, it was almost aimed at the target and the eye had returned to within 10 +/- 7 degrees, mean +/- SD, of straight ahead. Between the end of the gaze shift and the end of the head movement, gaze remained stable in space or a small correction saccade occurred. 4. Gaze movements < 20 degrees landed accurately on target whether the head was fixed or free. For larger target movements, both head-free and head-fixed gaze shifts became increasingly hypometric. Head-free gaze shifts were more accurate, on average, but also more variable. This suggests that gaze is controlled in a different way with the head free. For target amplitudes < 60 degrees, head position was hypometric but the error was rather constant at approximately 10 degrees.(ABSTRACT TRUNCATED AT 400 WORDS)


2018 ◽  
Vol 71 (9) ◽  
pp. 1860-1872 ◽  
Author(s):  
Stephen RH Langton ◽  
Alex H McIntyre ◽  
Peter JB Hancock ◽  
Helmut Leder

Research has established that a perceived eye gaze produces a concomitant shift in a viewer’s spatial attention in the direction of that gaze. The two experiments reported here investigate the extent to which the nature of the eye movement made by the gazer contributes to this orienting effect. On each trial in these experiments, participants were asked to make a speeded response to a target that could appear in a location toward which a centrally presented face had just gazed (a cued target) or in a location that was not the recipient of a gaze (an uncued target). The gaze cues consisted of either fast saccadic eye movements or slower smooth pursuit movements. Cued targets were responded to faster than uncued targets, and this gaze-cued orienting effect was found to be equivalent for each type of gaze shift both when the gazes were un-predictive of target location (Experiment 1) and counterpredictive of target location (Experiment 2). The results offer no support for the hypothesis that motion speed modulates gaze-cued orienting. However, they do suggest that motion of the eyes per se, regardless of the type of movement, may be sufficient to trigger an orienting effect.


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.


2019 ◽  
Author(s):  
Benedikt V. Ehinger ◽  
Katharina Groß ◽  
Inga Ibs ◽  
Peter König

ABSTRACTEye-tracking experiments rely heavily on good data quality of eye-trackers. Unfortunately, it is often that only the spatial accuracy and precision values are available from the manufacturers. These two values alone are not sufficient enough to serve as a benchmark for an eye-tracker: Eye-tracking quality deteriorates during an experimental session due to head movements, changing illumination or calibration decay. Additionally, different experimental paradigms require the analysis of different types of eye movements, for instance smooth pursuit movements, blinks or microsaccades, which themselves cannot readily be evaluated by using spatial accuracy or precision alone. To obtain a more comprehensive description of properties, we developed an extensive eye-tracking test battery. In 10 different tasks, we evaluated eye-tracking related measures such as: the decay of accuracy, fixation durations, pupil dilation, smooth pursuit movement, microsaccade detection, blink detection, or the influence of head motion. For some measures, true theoretical values exist. For others, a relative comparison to a gold standard eye-tracker is needed. Therefore, we collected our gaze data simultaneously from a gold standard remote EyeLink 1000 eye-tracker and compared it with the mobile Pupil Labs glasses.As expected, the average spatial accuracy of 0.57° for the EyeLink 1000 eye-tracker was better than the 0.82° for the Pupil Labs glasses (N=15). Furthermore, we detected less fixations and shorter saccade durations for the Pupil Labs glasses. Similarly, we found fewer microsaccades using the Pupil Labs glasses. The accuracy over time decayed only slightly for the EyeLink 1000, but strongly for the Pupil Labs glasses. Finally we observed that the measured pupil diameters differed between eye-trackers on the individual subject level but not the group level.To conclude, our eye-tracking test battery offers 10 tasks that allow us to benchmark the many parameters of interest in stereotypical eye-tracking situations, or addresses a common source of confounds in measurement errors (e.g. yaw and roll head movements).All recorded eye-tracking data (including Pupil Labs’ eye video files), the stimulus code for the test battery and the modular analysis pipeline are available (https://github.com/behinger/etcomp).BVE, KG, IIandPKconceived the experiment.IIandBVEcreated the experiment and recorded the gaze data.BVEandKGperformed the analysis.BVE, KGandPKreviewed the manuscript critically.


1997 ◽  
Vol 77 (2) ◽  
pp. 654-666 ◽  
Author(s):  
Douglas Tweed

Tweed, Douglas. Three-dimensional model of the human eye-head saccadic system. J. Neurophysiol. 77: 654–666, 1997. Current theories of eye-head gaze shifts deal only with one-dimensional motion, and do not touch on three-dimensional (3-D) issues such as curvature and Donders' laws. I show that recent 3-D data can be explained by a model based on ideas that are well established from one-dimensional studies, with just one new assumption: that the eye is driven toward a 3-D orientation in space that has been chosen so that Listing's law of the eye in head will hold when the eye-head movement is complete. As in previous, one-dimensional models, the eye and head are feedback-guided and the commands specifying desired eye position eye pass through a neural “saturation” so as to stay within the effective oculomotor range. The model correctly predicts the complex, 3-D trajectories of the head, eye in space, and eye in head in a variety of saccade tasks. And when it moves repeatedly to the same target, varying the contributions of eye and head, the model lands in different eye-in-space positions, but these positions differ only in their cyclotorsion about the line of sight, so they all point that line at the target—a behavior also seen in real eye-head saccades. Between movements the model obeys Listing's law of the eye in head and Donders' law of the head on torso, but during certain gaze shifts involving large torsional head movements, it shows marked, 8° deviations from Listing's law. These deviations are the most important untested predictions of the theory. Their experimental refutation would sink the model, whereas confirmation would strongly support its central claim that the eye moves toward a 3-D position in space chosen to obey Listing's law and, therefore, that a Listing operator exists upstream from the eye pulse generator.


1999 ◽  
Vol 81 (6) ◽  
pp. 3105-3109 ◽  
Author(s):  
T. Belton ◽  
R. A. McCrea

Contribution of the cerebellar flocculus to gaze control during active head movements. The flocculus and ventral paraflocculus are adjacent regions of the cerebellar cortex that are essential for controlling smooth pursuit eye movements and for altering the performance of the vestibulo-ocular reflex (VOR). The question addressed in this study is whether these regions of the cerebellum are more globally involved in controlling gaze, regardless of whether eye or active head movements are used to pursue moving visual targets. Single-unit recordings were obtained from Purkinje (Pk) cells in the floccular region of squirrel monkeys that were trained to fixate and pursue small visual targets. Cell firing rate was recorded during smooth pursuit eye movements, cancellation of the VOR, combined eye-head pursuit, and spontaneous gaze shifts in the absence of targets. Pk cells were found to be much less sensitive to gaze velocity during combined eye–head pursuit than during ocular pursuit. They were not sensitive to gaze or head velocity during gaze saccades. Temporary inactivation of the floccular region by muscimol injection compromised ocular pursuit but had little effect on the ability of monkeys to pursue visual targets with head movements or to cancel the VOR during active head movements. Thus the signals produced by Pk cells in the floccular region are necessary for controlling smooth pursuit eye movements but not for coordinating gaze during active head movements. The results imply that individual functional modules in the cerebellar cortex are less involved in the global organization and coordination of movements than with parametric control of movements produced by a specific part of the body.


2000 ◽  
Vol 84 (3) ◽  
pp. 1614-1626 ◽  
Author(s):  
Timothy Belton ◽  
Robert A. McCrea

The contribution of the flocculus region of the cerebellum to horizontal gaze pursuit was studied in squirrel monkeys. When the head was free to move, the monkeys pursued targets with a combination of smooth eye and head movements; with the majority of the gaze velocity produced by smooth tracking head movements. In the accompanying study we reported that the flocculus region was necessary for cancellation of the vestibuloocular reflex (VOR) evoked by passive whole body rotation. The question addressed in this study was whether the flocculus region of the cerebellum also plays a role in canceling the VOR produced by active head movements during gaze pursuit. The firing behavior of 121 Purkinje (Pk) cells that were sensitive to horizontal smooth pursuit eye movements was studied. The sample included 66 eye velocity Pk cells and 55 gaze velocity Pk cells. All of the cells remained sensitive to smooth pursuit eye movements during combined eye and head tracking. Eye velocity Pk cells were insensitive to smooth pursuit head movements. Gaze velocity Pk cells were nearly as sensitive to active smooth pursuit head movements as they were passive whole body rotation; but they were less than half as sensitive (≈43%) to smooth pursuit head movements as they were to smooth pursuit eye movements. Considered as a whole, the Pk cells in the flocculus region of the cerebellar cortex were <20% as sensitive to smooth pursuit head movements as they were to smooth pursuit eye movements, which suggests that this region does not produce signals sufficient to cancel the VOR during smooth head tracking. The comparative effect of injections of muscimol into the flocculus region on smooth pursuit eye and head movements was studied in two monkeys. Muscimol inactivation of the flocculus region profoundly affected smooth pursuit eye movements but had little effect on smooth pursuit head movements or on smooth tracking of visual targets when the head was free to move. We conclude that the signals produced by flocculus region Pk cells are neither necessary nor sufficient to cancel the VOR during gaze pursuit.


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