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Maintenance of equilibrium during tracing eye movements.

Horizontal tracing movements of eyes modify the type of vertical posture maintenance decreasing the role of the lower segment in the regulation of the position of the pressure center. The relationship between fluctuations of the pressure center in the frontal and sagittal planes increases. Periodicity of eye movements corresponds to fluctuations of the pressure center and these signals were phase shifted relatively to each other.

Adult↗

Is target movement as well as target displacement a stimulus for saccadic eye movements?

The effects of visual movement on saccadic eye movement have been examined. In a classic apparent-movement demonstration with two successively exposed line-segment targets the quality of the movement is dependent on the relative orientation of the line segments. If saccadic eye movements are elicited between the targets in this situation, the configuration leading to optimal apparent movement also leads to the shortest-latency saccades. When a single line segment is succeeded by two line segments flanking it on opposite sides, and if one of these has the same orientation as the initial one and the other a different orientation, then apparent motion is seen between the two lines with the same orientation. However, the direction of saccades elicited in this configuration is not influenced by the relative orientations of the line segments. The two results together suggest that the effect of visual movement on saccadic eye movement is nonspecific.

Eye Movements↗

Seeking one's heading through eye movements.

A study of eye movements during simulated travel toward a grove of four stationary trees revealed that observers looked most at pairs of trees that converged or decelerated apart. Such pairs specify that one's direction of travel, called heading, is to the outside of the near member of the pair. Observers looked at these trees more than those that accelerated apart; such pairs do not offer trustworthy heading information. Observers also looked at gaps between trees less often when they converged or diverged apart, and heading can never be between such pairs. Heading responses were in accord with eye movements. In general, if observers responded accurately, they had looked at trees that converged or decelerated apart; if they were inaccurate, they had not. Results support the notion that observers seek out their heading through eye movements, saccading to and fixating on the most informative locations in the field of view.

Adult↗

Estimating heading during real and simulated eye movements.

The ability to judge heading during tracking eye movements has recently been examined by several investigators. To assess the use of retinal-image and extra-retinal information in this task, the previous work has compared heading judgments with executed as opposed to simulated eye movements. For eye movement velocities greater than 1 deg/sec, observers seem to require the eye-velocity information provided by extra-retinal signals that accompany tracking eye movements. When those signals are not provided, such as with simulated eye movements, observers perceive their self-motion as curvilinear translation rather than the linear translation plus eye rotation being presented. The interpretation of the previous results is complicated, however, by the fact that the simulated eye movement condition may have created a conflict between two possible estimates of the heading: one based on extra-retinal solutions and the other based on retina-image solutions. In four experiments, we minimized this potential conflict by having observers judge heading in the presence of rotations consisting of mixtures of executed and simulated eye movements. The results showed that the heading is estimated more accurately when rotational flow is created by executed eye movements alone. In addition, the magnitude of errors in heading estimates is essentially proportional to the amount of rotational flow created by a simulated eye rotation (independent of the total magnitude of the rotational flow). The fact that error magnitude is proportional to the amount of simulated rotation suggests that the visual system attributes rotational flow unaccompanied by an eye movement to a displacement of the direction of translation in the direction of the simulated eye rotation.

Cues↗

EMMA--an eye movement measurement and analysis system.

A system has been developed for stimulation, recording and analysis of a wide range of eye movements. Eye movements are stimulated with an LED bar or a video projector under the control of a PC. The eye movements are measured using a scleral reflection technique (IRIS instrument), and sampled and stored on a PC. A range of tests have been developed to measure saccadic and smooth pursuit eye movements. A variety of tools have been developed to assist in the analysis of the data. Several research studies have ably demonstrated the utility and versatility of the system.

Equipment Design↗

The circular eye-tracking test. I. Simultaneous recording of the horizontal and vertical component of eye movement in the eye-tracking test.

Almost all of the conventional clinical reports on the eye-tracking test are made on the horizontal component of the smooth pursuit function of oculomotor system; few are made on the vertical component. One reason is that during the conventional vertical eye-tracking test the artifacts caused by movement of the upper eyelids often make it difficult to analyze the data. To overcome this problem the authors have developed a method to record simultaneously the horizontal and vertical components of eye movement in the eye-tracking test, and moreover, to reduce artifacts as much as possible. Our method, the circular eye-tracking test, is performed by means of an apparatus that provides a visual stimulus drawing a circle clockwise, at a rate of 120 degrees/sec, with a radius of 25.5 cm, and viewed at a distance of 70 cm.

Brain Diseases↗

3D eye movements--basics and clinical applications.

The eye muscles in humans are arranged such that they allow rotations of the globe about any axis in three-dimensional space. Only 3D analysis techniques will suffice to describe the sometimes complex dependencies between the elicited eye movements and the stimulation conditions. With the recent availability of modern 3D eye movement measurement techniques and the further development of appropriate mathematical descriptions, we are now able to study eye movements in all three degrees of freedom. This article describes the basic mathematical tools for 3D eye movement analysis like rotation vectors, reference frames, coordinate systems, and the concept of Listing's law. In a second part some clinical applications are presented. The close coupling between the vestibular and the oculomotor system suggests that by observing spontaneous and elicited eye movements in the case of an acute unilateral vestibular pathology we might be able to find out which parts of the vestibular system (that is, which of the semicircular canals or which of the otoliths) are affected. The rationale of such an analysis is based on the observation that electrical stimulation of single semicircular canal afferents in animals induces eye movements that lie roughly in the plane of the canal. It is, therefore, possible to deduce which canals cause the eye movements observed when only some parts of the vestibular system are defective. Thus, the analysis of 3D movements not only provides an improved understanding of how the brain organizes movement in 3D space, but also has the potential to significantly improve our diagnostic capabilities.

Electronystagmography↗

Speed-accuracy characteristics of saccadic eye movements.

Speed-accuracy trade-off characteristic of horizontal saccadic eye movements were examined in this study. Unlike limb movements, saccadic eye movements are preprogrammed, unidimensional, and do not involve target impact. Hence, they provide an optimal test of the impulse variability account of the speed-accuracy trade-off in rapid movements. Subjects were required to alternately look at two target lights as fast and as accurately as possible for a period of 10 s. Target lights subtended angles of 5, 10, 15, and 20 degrees. By restricting target distances to less than 20 degrees of arc, the speed-accuracy relation was examined for single horizontal saccadic movements of the eye. movement of the dominant eye was tracked with an infra-red eye monitoring device. Fifty saccadic movements of the eye were recorded for each target distance and used to compute the average amplitude, duration, and velocity of eye movements, as well as, movement endpoint variability. An increase in both average velocity and movement endpoint variability with increasing movement amplitude was found. This, together with the unique features of the eye movement system, support the impulse variability account of the speed-accuracy trade-off in rapid movements.

Journal Article↗

Smooth pursuit eye movements and optokinetic nystagmus elicited by intermittently illuminated stationary patterns.

Stationary periodic visual patterns (row of equally spaced dots or black-white stripes) of the period Ps illuminated stroboscopically with a flash frequency fs induce an apparent movement perception (sigma-movement) when slow eye movements are performed across the periodic pattern. The movement appears in the direction of the eye movements when the angular speed VE of the eyes corresponds to the following condition: Ve = k . Ps . fs [deg . s-1] (1) k is a constant and equals 1 (or exceptionally 2 or 3). The sigma-movement induces a sigma-OKN with an average angular speed of its slow phases corresponding to Eq.(1). sigma-OKN can be elicited when identical foveal or identical extrafoveal stimulus patterns are applied from flash to flash. A considerable random variability of the flash sequence does not interrupt the sigma-movement and the sigma-OKN. Both phenomena can also be elicited by a stimulus pattern with its periodicity hidden in spatial noise and this periodic pattern only becomes visible during the eye movements. It is argued that the sigma-phenomena are caused by efference copy signals of the gaze control system, which interact with the afferent signals (displacement of visual stimuli on the retina) at different levels of the afferent visual system. One interaction is supposed at a cortical level where the extrapersonal visual space is represented.

Adult↗

Modifying oculomotor activity in awake subjects increases the amplitude of eye movements during REM sleep.

The eye movements of human subjects were experimentally modified while they were awake to determine the effect of waking experience on electroculographic activity during rapid eye movement (REM) sleep. After normal eye movements were monitored under controlled conditions, subjects spent 5 days wearing goggles that contained minification lenses and that curtailed vision to a 5 degree field. The amplitude and frequency of eye movements decreased when subjects were awake and increased during REM sleep; sleep stage durations and distributions were unaffected. Values returned to normal in the first 24 hours of recovery.

Adult↗

Idiosyncratic characteristics of saccadic eye movements when viewing different visual environments.

Eye position was recorded in different viewing conditions to assess whether the temporal and spatial characteristics of saccadic eye movements in different individuals are idiosyncratic. Our aim was to determine the degree to which oculomotor control is based on endogenous factors. A total of 15 naive subjects viewed five visual environments: (1) The absence of visual stimulation (i.e. a dark room); (2) a repetitive visual environment (i.e. simple textured patterns); (3) a complex natural scene; (4) a visual search task; and (5) reading text. Although differences in visual environment had significant effects on eye movements, idiosyncrasies were also apparent. For example, the mean fixation duration and size of an individual's saccadic eye movements when passively viewing a complex natural scene covaried significantly with those same parameters in the absence of visual stimulation and in a repetitive visual environment. In contrast, an individual's spatio-temporal characteristics of eye movements during active tasks such as reading text or visual search covaried together, but did not correlate with the pattern of eye movements detected when viewing a natural scene, simple patterns or in the dark. These idiosyncratic patterns of eye movements in normal viewing reveal an endogenous influence on oculomotor control. The independent covariance of eye movements during different visual tasks shows that saccadic eye movements during active tasks like reading or visual search differ from those engaged during the passive inspection of visual scenes.

Adult↗

Eye movement patterns in REM sleep.

Eye movements in 6 healthy men and women were studied for recurrent patterns during REM sleep. The REM periods of nocturnal polysomnograms, on 2 consecutive nights, were analyzed in each subject. A discrete scale from 1 to 8 was used to record each eye position. The total number of recorded eye positions for the 2 nights of testing varied from 1314 to 3006. The distributions of eye movement were similar for males and females, for both nights of testing for each subject, among individual REM periods, and between subjects. This was in spite of marked differences in the number and length of REM periods, and in the number of eye movements per minute of REM sleep. In 5 of 6 subjects there was a marked tendency for the eyes to move between the 2 opposite lateral positions. Regardless of the eye position, the opposite movement was generally most likely, with an underlying tendency to return to the most opposite of the two lateral positions. In the remaining subject the opposite movement was also favored, but in this subject eye movements were more likely to be vertical rather than horizontal. Our data suggest that eye movements in REM sleep are organized in complex recurring patterns, with marked similarities between subjects. The significance of these patterns and the significance of deviations from these patterns require further study.

Adult↗

Quantitative measurement of smooth pursuit eye movements.

Smooth pursuit eye movements were quantitatively assessed in 25 normal subjects and 22 patients. A laboratory digital computer was used to compute 200 eye velocity samples per second and to statistically compare these eye velocity measurements for five different object velocities. Of six statistics evaluated, mode eye velocity showed the least variability in normal subjects and was most frequently abnormal in patients. Compared to normal subjects, patients with brain stem degeneration and cerebellar-pontine angle tumors with brain stem compression had significant impairment of smooth pursuit. Patients with peripheral vestibular lesions and C-P angle tumors without brain stem compression did not have impaired smooth pursuit. These preliminary findings suggest that quantitative measurement of pursuit eye velocity can be a sensitive test for brain stem dysfunction.

Adult↗

Short term modification of disparity vergence eye movements.

Dynamics of disparity vergence eye movements can be modified by adaptive stimuli that generate large transient disparities. These modifications were observed for convergence as well as divergence eye movements. After modification, the peak velocities of the step responses for convergence and divergence were substantially higher than in normal baseline responses, a change observed in all four subjects studied. The change in peak velocity of a step response occurred very rapidly after presentation of the adaptive stimuli. Main sequence plots showed that first-order dynamic characteristics increased for post-adaptive responses with respect to normal step responses. Hence, response modification could be quantified as a change in gain accompanied with an increase in the effective response time constant. The adaptive responses to convergent and divergent 'disappearing' step stimuli revealed that the adaptation process modifies the high-velocity component of both disparity convergence and divergence eye movements. Moreover, a gain change in this component alone could account for both the gain and the time constant modifications seen in the overall response. A process of recovery or de-adaptation was also observed for both convergence and divergence eye movements. This observed short-term modification demonstrates a unique control mechanism for vergence eye movements that is effective in either direction.

Adaptation, Ocular↗

Compensatory eye movements produced during fictive swimming of a deafferented, reduced preparation in vitro.

Movement of the head during locomotion must be offset by movement of the eyes in the opposite direction in order to maintain a stable gaze. These compensatory eye movements are classically attributed to ocular reflexes elicited by vestibular stimulation and by slip of the retinal image. In the present study, compensatory eye movements accompanied episodes of fictive swimming expressed in vitro by a reduced, deafferented preparation of the bullfrog larva (tadpole). Thus, there are mechanisms for generation of eye movements, and their coordination with locomotion, that are endogenous to the central nervous system.

Abducens Nerve↗

Vasoactive intestinal polypeptide microinjections into the oral pontine tegmentum enhance rapid eye movement sleep in the rat.

Rapid eye movement sleep can be elicited in the rat by microinjection of the cholinergic agonist carbachol into the oral pontine reticular nucleus. Intracerebroventricular administration, during the light period, of vasoactive intestinal peptide enhances rapid eye movement sleep in several species. Since this peptide is co-localized with acetylcholine in many neurons in the central nervous system, it was assumed that the oral pontine tegmentum could also be one target for vasoactive intestinal peptide to induce rapid eye movement sleep. This hypothesis was tested by recording the sleep-wakefulness cycle in freely-moving rats injected with vasoactive intestinal peptide or its fragments (1-12 and 10-28) directly into the oral pontine reticular nucleus. when administered into the posterior part of this nucleus, vasoactive intestinal peptide at 1 and 10 ng (in 0.1 microliter of saline), but not its fragments, induced a 2-fold enhancement of rapid eye movement sleep during 4 h, at the expense of wakefulness. At the dose of 10 ng, a significant increase in rapid eye movement sleep persisted for up to 8 h. Moreover, when the peptide was injected into the centre of the positive zone, rapid eye movement sleep was enhanced during three to eight consecutive days. These data provide the first evidence that rapid eye movement sleep can be elicited at both short- and long-term by a single intracerebral microinjection of vasoactive intestinal peptide. Peptidergic mechanisms, possibly in association with cholinergic mechanisms, within the caudal part of the oral pontine reticular nucleus may play a critical role in the long-term regulation of rapid eye movement sleep in rats.

Animals↗

Cortico-cortical networks and cortico-subcortical loops for the higher control of eye movements.

There are multiple distinct regions, or eye fields, in the cerebral cortex that contribute directly to the initiation and control of voluntary eye movements. We concentrate on six of these: the frontal eye field, parietal eye field, supplementary eye field, middle superior temporal area, prefrontal eye field, and area 7 m (precuneus in humans). In each of these regions: (1) there is neural activity closely related to eye movements; (2) electrical microstimulation produces or modifies eye movements; (3) surgical lesions or chemical inactivation impairs eye movements; (4) there are direct neural projections to major structures in the brainstem oculomotor system; and (5) increased activity is observed during eye movement tasks in functional magnetic resonance imaging or positron emission tomography experiments in humans. Each of these eye fields is reciprocally connected with the other eye fields, and each receives visual information directly from visual association cortex. Each eye field has distinct subregions that are concerned with either saccadic or pursuit eye movements. The saccadic subregions are preferentially interconnected with other saccade subregions and the pursuit subregions are preferentially interconnected with other pursuit subregions. Current evidence strongly supports the proposal that there are parallel cortico-cortical networks that control purposeful saccadic and pursuit eye movements, and that the activity in those networks is modulated by feedback information, via the thalamus, from the superior colliculus, basal ganglia, and cerebellum.

Animals↗