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Eye movements evoked by electrical stimualtion of the brain in anesthetized fishes.

Several eye movements were evoked by electrical stimulation of the brain in anesthetized sunfish and goldfish. Conjugate lateral rolling movements, similar to eye movements observed when an unoperated fish is rotated about its long axis, were evoked from the acoustico-lateral area of the medulla and the eminentia granularis and an adjacent medial portion of the cerebellum. Bilateral and unilateral backward rotations, similar to the eye movements observed when unoperated fish are rotated forward about the interpupillary axis, were evoked from the medial longitudinal fasciculus and areas related to the oculomotor nerve. Bilateral forward rotations, comparable to the eye movements resulting when unoperated fish are rotated backward about the interpupillary axis, were elicited by stimulation near the trochlear nerve roots in the valvula of the cerebellum; unilateral responses resulted from stimulation near the exiting trochlear nerves. Convergence was elicited by stimulation in the midline near the oculomotor complex and the medial longitudinal fasciculus while unilateral vergence responses were triggered by stimulation in the medial longitudinal fasciculus and areas lateral to the oculomotor nucleus. Conjugate eye movements in the horizontal plane were frequently evoked but were not studied in detail.

Anesthesia↗

Functional and anatomic organization of three-dimensional eye movements in rabbit cerebellar flocculus.

1. The three-dimensional, binocular eye movements evoked by electrical microstimulation of the cerebellar flocculus of alert, pigmented rabbits were recorded using the scleral search coil technique. The components of these eye movements were obtained in reference to an orthogonal coordinate system consisting of a vertical axis and two horizontal axes at 45 degrees and 135 degrees azimuth. The azimuth coordinate was taken to increase to both sides from the 0 degrees reference in the direction of the nose. 2. Eye movements were evoked most readily by stimulation (0.2-ms pulses at 200 Hz for 1 s, intensity < or = 20 microA) at loci in the deep granular layer and the white matter. They consisted of slow (5-20 deg/s) movements. The responses were either binocular, with the eye ipsilateral to the stimulated flocculus usually having the larger amplitude, or were monocular, in which case they were restricted to the ipsilateral eye. 3. The evoked responses were classified according to the combination of the largest measured component of rotation for the two eyes and its sense of rotation (clockwise, CW, or counterclockwise, CCW). Seventy-eight percent of the evoked eye movements could be placed in one of two classes. For one of these classes the largest response component was a short-latency abduction of the ipsilateral eye about its vertical axis (19%), whereas for the other class (59%), the largest response component was a short-latency CCW rotation of the ipsilateral (left) eye about its 135 degrees axis. This response was frequently (50%) accompanied by a smaller short-latency CW rotation of the contralateral (right) eye about its 45 degrees axis. 4. The two main classes of three-dimensional eye movements are associated differentially with anatomically distinguishable compartments that are revealed by acetylcholinesterase histochemistry. Of the five anatomically distinguishable compartments in the floccular white matter, three are predominant. The middle of these three compartments is associated with the vertical axis class of movements, whereas the two adjacent compartments are associated with the 135 degrees class of eye movements. The eye movement relation of the other two, smaller compartments, was not determined. 5. The spatial orientation of the rotation axes of the two main classes of evoked eye movements closely corresponds to that of the preferred axes of the visual climbing fiber input to the flocculus. This suggests that both are organized in a similar coordinate system.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Antisaccade eye movement abnormalities in Tourette syndrome: evidence for cortico-striatal network dysfunction?

Saccadic eye movements are rapid eye movements which act to redirect the eyes from one object of interest to another. Accurately and objectively measurable, their underlying neuroanatomical mechanisms have been extensively studied. The antisaccade task allows the study of the frontocortico-striatal network involved in the voluntary control of saccadic eye movements. In this task, the subject is instructed to inhibit a reflex eye movement towards a peripheral target light and, instead, to generate a movement in the equal and opposite direction. An error occurs when the subject fails to suppress reflexive saccades towards the target. Significantly high error rates and increased latencies in the antisaccade task have been reported in disorders associated with dysfunction of the frontocortico-striatal network. Increased saccadic eye movement latencies and error rates have been reported in Tourette syndrome patients (n = 4) who were receiving antipsychotic medication. To investigate this further, we tested the antisaccade task on six male Tourette syndrome patients. The results were compared with 18 age- and sex-matched mentally and physically healthy, medication/alcohol-free controls. Antisaccade latencies were (mean +/- SD; ms) 751.2+/-186.7 for the Tourette syndrome group and 417+/-75.3 for controls, and error rates were 59+/-14.3 for the Tourette syndrome group and 11.9+/-6.4 for controls, respectively. These significant results may further support dysfunction of the frontocortico-striatal network in Tourette syndrome.

Adolescent↗

Tonic cervical stimulation: does it influence eye position and eye movements in man?

In healthy subjects eye movements were analysed during body rotation, during trunk torsion either with the head passively held stationary in space or with the head voluntarily stabilized in space, and during voluntary head movements. Trapezoidal movements around the vertical axis were performed (+/- 40 degrees, plateau 10 s, duration of ramp 1 or 4 s). Moreover the influence of a tonic head deviation up to 40 degrees on optokinetic nystagmus and on vestibulo-ocular reflex during sinusoidal turning was examined. Eye movements were recorded by DC-electrooculography. Saccadic and slow components of eye movements and the shift of eye position during the plateau of the trapezoidal stimulus were analysed. For all modes of stimulation during the plateau no nystagmus occurred. At the end of the dynamic phase of the stimulus relatively frequent eye deviations--mostly in the direction of the head deviation--were observed, not only after turning the trunk with the head stabilized in space (cervical stimulation) but also after turning head and trunk together. The fact that such eye deviations are thus observed even in the absence of any tonic, especially cervical stimulus, supports the assumption that they cannot be attributed to a tonic stimulus but merely to an effect of the preceding phasic stimulus which outlasts them. Also amplitude and direction of eye shifts during the plateau do not depend on a tonic stimulus, but merely on the eye deviation reached at the end of the dynamic phase of stimulation. Optokinetic nystagmus and vestibulo-ocular reflex are not influenced by an additional tonic cervical stimulus.

Adult↗

The effect of visual field defects on eye movements and practical fitness to drive.

Eye movements of subjects with visual field defects due to ocular pathology were monitored while performing a dot counting task and a visual search task. Subjects with peripheral field defects required more fixations, longer search times, made more errors, and had shorter fixation durations than control subjects. Subjects with central field defects performed less well than control subjects although no specific impairment could be pinpointed. In both groups a monotonous relationship was observed between the visual field impairment and eye movement parameters. The use of eye movement parameters to predict viewing behavior in a complex task (e.g. driving) was limited.

Adult↗

[Disorders of eye movements in point fixation of the dominant eye in unilateral deprivation amblyopia, squint amblyopia and unilateral organ damage].

In four juvenile patients with unilateral deprivation amblyopia three types of involuntary eye movement were identified by simultaneous binocular infrared reflection oculography (IROG). During apparent steady-point fixation of the dominant eye three cases showed sinusoidal pendular movements (type 1) of the amblyopic eye mainly in a horizontal and slightly in a vertical direction, amplitude (A) 1 to 4 degrees, frequency (F) 0.25-0.4 Hz; only one patient had similar movements with smaller deviations even in the fixing eye. In three cases disturbed fixation of the dominant eye was found, a lateral drift (type 2) up to 3 degrees followed by slow correcting movement (F 0.05-0.1 Hz). At the same time there were disjugate movements of the amblyopic eye, the movements of the two eyes resulting in variations in squint angle of up to 6-7 degrees. These movements of the weaker eyes were clinically observed by Ohm (1958) and were termed pendular flutter to distinguish them from nystagmus. This paper presents the first objectively recorded curves of these movements, and differentiates them according to the various types of movement. Type 3 is a fine jerky or pendular nystagmus (A = max. 1 degree, F = 1.0-2.8 Hz), which was also present in three cases. Two patients had a combination of all three types of motility disturbance. Four cases with unilateral organ lesions, either congenital or acquired in early infancy, were observed and compared. All these cases showed excessive pendular flutter (type 1)--horizontal in the case of the fixing eye and horizontal and vertical in the case of the weaker eye.(ABSTRACT TRUNCATED AT 250 WORDS)

Adolescent↗

Vertical object motion during horizontal ocular pursuit: compensation for eye movements increases with presentation duration.

Smooth pursuit eye movements change the retinal image motion of objects in the visual field. To enable an observer to perceive the motion of these objects veridically, the visual system has to compensate for the effects of the eye movements. The occurrence of the Filehne-illusion (illusory motion of a stationary object during smooth pursuit) shows that this compensation is not always perfect. The amplitude of the illusion appears to decrease with increasing presentation durations of the stationary object. In this study we investigated whether presentation duration has the same effect when an observer views a vertically moving object during horizontal pursuit. In this case, the pursuit eye movements cause the perceived motion path to be oblique instead of vertical; this error in perceived motion direction should decrease with higher presentation durations. In Experiment 1, we found that the error in perceived motion direction indeed decreased with increasing presentation duration, especially for higher pursuit velocities. The results of Experiment 2 showed that the error in perceived motion direction did not depend on the moment during pursuit at which the stimulus was presented, suggesting that the degree of compensation for eye movements is constant throughout pursuit. The results suggest that longer presentation durations cause the eye movement signal that is used by the visual system to increase more than the retinal signal.

Adult↗

Respiratory changes associated with rapid eye movements in normo- and hypercapnia during sleep.

Rapid eye movements during rapid-eye-movement (REM) sleep are associated with rapid, shallow breathing. We wanted to know whether this effect persisted during increased respiratory drive by CO2. In eight healthy subjects, we recorded electroencephalographic, electrooculographic, and electromyographic signals, ventilation, and end-tidal PCO2 during the night. Inspiratory PCO2 was changed to increase end-tidal PCO2 by 3 and 6 Torr. During normocapnia, rapid eye movements were associated with a decrease in total breath time by -0.71 +/- 0.19 (SE) s (P < 0.05) because of shortened expiratory time (-0.52 +/- 0.08 s, P < 0.001) and with a reduced tidal volume (-89 +/- 27 ml, P < 0.05) because of decreased rib cage contribution (-75 +/- 18 ml, P < 0.05). Abdominal (-11 +/- 16 ml, P = 0.52) and minute ventilation (-0.09 +/- 0.21 ml/min, P = 0.66) did not change. In hypercapnia, however, rapid eye movements were associated with a further shortening of total breath time. Abdominal breathing was also inhibited (-79 +/- 23 ml, P < 0.05), leading to a stronger inhibition of tidal volume and minute ventilation (-1.84 +/- 0.54 l/min, P < 0.05). We conclude that REM-associated respiratory changes are even more pronounced during hypercapnia because of additional inhibition of abdominal breathing. This may contribute to the reduction of the hypercapnic ventilatory response during REM sleep.

Adult↗

On the origin of eye movement desensitization.

Francine Shapiro, the founder of eye movement desensitization therapy (EMD) claims to have serendipitously discovered this technique by experiencing spontaneous saccadic eye movements in response to disturbing thoughts during a walk in the woods. This explanation is difficult to accept because normal saccadic eye movements appear to be physiologically undetectable and are typically triggered by external stimuli. In the present limited study, all subjects failed to elicit any experience of eye movements, thus calling into question how best to interpret Dr. Shapiro's experience and the origin of EMD.

Arousal↗

Effects of background motion on eye-movement information.

The effect of background stimulus on eye-movement information was investigated by analyzing the underestimation of the target velocity during pursuit eye movement (Aubert-Fleishl paradox). In the experiment, a striped pattern with various brightness contrasts and spatial frequencies was used as a background stimulus, which was moved at various velocities. Analysis showed that the perceived velocity of the pursuit target, which indicated the magnitudes of eye-movement information, decreased when the background stripes moved in the same direction as eye movement at higher velocities and increased when the background moved in the opposite direction. The results suggest that the eye-movement information varied as a linear function of the velocity of the motion of the background retinal image (optic flow). In addition, the effectiveness of optic flow on eye-movement information was determined by the attributes of the background stimulus such as the brightness contrast or the spatial frequency of the striped pattern.

Eye Movements↗

Apparent movement and appearance of periodic stripes during eye movements across a stroboscopically illuminated random dot pattern.

When the eyes follow a small target moving across a stationary random dot pattern illuminated stroboscopically at 3--45 flashes . s-1, the structure of the random dot pattern is seen as moving in the direction of the eye pursuit movements (sigma-movement). At flash frequencies above 9.5 flashes . s-1, periodic stripes oriented perpendicularly to the direction of the pursuit eye movements appear and are also seen as moving in the direction of the eye pursuit movement. The period Ps of the apparent stripes depended on the flash frequency fs and the angular speed of the eyes Ve: Formula: see text. Apparent sigma-movement and apparent stripes were also seen when eye pursuit movements were initiated outside of the random dot pattern and they continued autonomously without a moving target across the pattern. The angular velocity of the sigma-pursuit movement across the random dot pattern depended on the flash frequency and the angular velocity of the initiating target. With the eyes stationary but a moving random dot pattern, the apparent stripes also appeared at flash frequencies fs greater than or equal to 9.5 flashes . s-1. Eq. (1) was valid when Ve was replaced by the target angular velocity Vs (k = 1).

Adult↗

Acoustic-induced eye movements.

This article reports the influence of moving acoustic signals on eye movements (oculomotor functions) using 350 healthy test subjects. An acoustic test setting developed especially for this purpose has made possible the application of moving acoustic signals of varying frequency and form (square stimuli, sinusoidal stimuli, circular movements), as well as the examination of visual-acoustic and vestibular-acoustic interactions. It must be stressed that according to these investigations, both voluntary and involuntary eye movements can be demonstrated in response to moving acoustic signals. Involuntary eye movements, however, under th given experimental conditions, only are observed in about 20% of all test subjects and their frequency of occurrence is highest in young female subjects. They are most pronounced in darkness and when there is no fixation, and they can be coupled to varying degrees with the given acoustic signal. Voluntary and involuntary eye movements do not differ very much from one another. They consist mainly of a series of saccades that form staircase jerks. In both cases, no typical smooth movements or nystagmoid movement forms occur--which are predominant in tracking eye movements of visual stimuli. Occasionally, slight drifting movements can be demonstrated in the intersaccadic intervals. At a larger movement amplitude and increased movement frequency of the sound signal, pure opposite jerks finally appear, which follow up to a movement frequency of 1 Hz. The eye movements are influenced only slightly by a change in the amplitude of the sound movement (+/- 15 degrees to +/- 90 degrees). However, a close phase relationship between eye and sound movement exists. If instead of continuous acoustic signals (music), single burst signals are given at an increasing repetition rate, this results (at low repetition rates of bursts) in orienting reactions upon the individual signals, while at high signal repetition rates (above 0.5 Hz), continuous eye movements in the form of staircase jerks result. In investigating acoustically induced tracking eye movements, many new questions have arisen that previously have been open only to psychophysical studies.

Auditory Pathways↗

The physiologic basis of conjugate eye movements.

We review the physiologic basis for conjugate eye movements and the role of the cerebellum in their regulation. Specifically, the characteristics of the vestibulo-ocular reflex, the optokinetic reflex, pursuit eye movements, and saccadic eye movements are discussed, emphasizing that the ocular motor system operates to minimize the slip of an image on the retina under various behavioral conditions. Such conditions include changes in the position of the head in space and various combinations of eye and head movements that occur while pursuing a target in visual space. The importance of the cerebellum in regulating smooth pursuit movements is emphasized, and the general categories of deficits resulting from abnormalities in this function are reviewed briefly. The abnormalities in conjugate eye movements observed after cerebellar lesions are related to the abnormalities in movements of the extremities observed in the same patients.

Animals↗

Optokinetic eye movements elicited by radial optic flow in the macaque monkey.

We recorded spontaneous eye movements elicited by radial optic flow in three macaque monkeys using the scleral search coil technique. Computer-generated stimuli simulated forward or backward motion of the monkey with respect to a number of small illuminated dots arranged on a virtual ground plane. We wanted to see whether optokinetic eye movements are induced by radial optic flow stimuli that simulate self-movement, quantify their parameters, and consider their effects on the processing of optic flow. A regular pattern of interchanging fast and slow eye movements with a frequency of 2 Hz was observed. When we shifted the horizontal position of the focus of expansion (FOE) during simulated forward motion (expansional optic flow), median horizontal eye position also shifted in the same direction but only by a smaller amount; for simulated backward motion (contractional optic flow), median eye position shifted in the opposite direction. We relate this to a change in Schlagfeld typically observed in optokinetic nystagmus. Direction and speed of slow phase eye movements were compared with the local flow field motion in gaze direction (the foveal flow). Eye movement direction matched well the foveal motion. Small systematic deviations could be attributed to an integration of the global motion pattern. Eye speed on average did not match foveal stimulus speed, as the median gain was only approximately 0.5-0.6. The gain was always lower for expanding than for contracting stimuli. We analyzed the time course of the eye movement immediately after each saccade. We found remarkable differences in the initial development of gain and directional following for expansion and contraction. For expansion, directional following and gain were initially poor and strongly influenced by the ongoing eye movement before the saccade. This was not the case for contraction. These differences also can be linked to properties of the optokinetic system. We conclude that optokinetic eye movements can be elicited by radial optic flow fields simulating self-motion. These eye movements are linked to the parafoveal flow field, i.e., the motion in the direction of gaze. In the retinal projection of the optic flow, such eye movements superimpose retinal slip. This results in complex retinal motion patterns, especially because the gain of the eye movement is small and variable. This observation has special relevance for mechanisms that determine self-motion from retinal flow fields. It is necessary to consider the influence of eye movements in optic flow analysis, but our results suggest that direction and speed of an eye movement should be treated differently.

Animals↗

Compensatory head and eye movements in the frog and their contribution to stabilization of gaze.

Compensatory head movements, recorded in unrestrained frogs, were compared to compensatory eye movements recorded from animals that had their head fixed. Movements were evoked by oscillating the animal in the dark (vestibular stimulation) or in the light in front of an earth-fixed, patterned visual background (combined stimulation) or by rotating vertical black and white bars (optokinetic stimulation) around the stationary animal. Oscillations occurred in the horizontal plane at frequencies between 0.025 and 0.5 Hz. Gain and phase values of head and eye movements, relative to stimulus movements were calculated. Evoked eye movements were limited in amplitude to +/- 3-6 degrees, increasing with the size of the animal. Head movements were limited to +/- 30-40 degrees. Resetting fast-phases of both head and eyes were very rarely observed during sinusoidal stimulation and no eye movements were recorded in the absence of intended head movements. Vestibularly evoked head movements exhibited a frequency-dependent threshold that was not observed for vestibulo-ocular responses. Above threshold, the gain of evoked head responses increased and reached a frequency-dependent plateau at which the system behaved approximately linearly. Within the linear range, gain of vestibularly evoked responses increased with frequency (from 0.04 at 0.025 Hz to 0.75 at 0.5 Hz) and phase lead decreased (from about 80 degrees to 0 degrees). Vestibularly evoked eye movements similarly increased in gain from 0.05 to 0.56 and decreased in phase lead from about 56 degrees to 10 degrees over the same frequency range. Optokinetically evoked head and eye movements had their highest gains (about 0.8 and 0.5) at low constant velocities (less than or equal to 1-4 degrees/S) or frequencies (less than or equal to 0.025 Hz). At higher constant velocities or frequencies the gain dropped. The phase lag increased from close to zero (at 0.025 Hz) to about 60 degrees for the head and to about 20 degrees for the eye movements (at 0.25 Hz). These phase lags are explained by reaction times of the evoked movements of about 600 ms (head) and 200 ms (eyes). Combined stimulation evoked compensatory head movements with gain and phase values that were frequency-independent in the linear range. Head movements compensated for about 80-90% of the imposed gaze shift with a small phase lag (0-10 degrees). Evoked eye movements were found to be large enough in amplitude and fast enough in time to enable a frog to stabilize its gaze exclusively with slow phase compensatory movements for a large variety of frequency and amplitude combinations. The two motor systems controlling movements of the head and the eye are matched in such a way that the non-linearities of the evoked eye movements can compensate for the non-linearities of the evoked head movements.

Animals↗

Breech presentation: is there a difference in eye movement patterns compared with cephalic presentation in the human fetus at term?

OBJECTIVE: Our purpose was to characterize eye movement patterns in the human fetus at term persisting in breech presentation. STUDY DESIGN: Studied were 11 fetuses in breech presentation and 12 in cephalic presentation at 36 to 41 weeks' gestation, of which the presentation remained unchanged from 32 weeks' gestation until delivery. With real-time ultrasonography we analyzed three measurements by means of the Mann-Whitney rank-sum test: (1) duration of eye-movement and no-eye-movement periods, (2) frequency of eye movements, and (3) proportion of horizontal, vertical, and oblique eye movements. RESULTS: There were significant differences in the eye movement directions between breech and cephalic fetuses: horizontal (median 70.3% vs 80.2%), vertical (17.4% vs 11.0%), and oblique (13.1% vs 9.1%), respectively. No differences were noted in the duration of eye-movement and no-eye-movement periods or in the frequency of eye movements. CONCLUSION: Eye movement pattern, in direction, of term fetuses persisting in breech presentation run on a different in-utero developmental course of neural control from those in cephalic fetuses.

Breech Presentation↗

Smooth pursuitlike eye movements evoked by microstimulation in macaque nucleus reticularis tegmenti pontis.

1. Smooth pursuitlike eye movements were evoked with low current microstimulation delivered to rostral portions of the nucleus reticularis tegmenti pontis (rNRTP) in alert macaques. Microstimulation sites were selected by the observation of modulations in single-cell firing rates that were correlated with periodic smoothpursuit eye movements. Current intensities ranged from 10 to 120 microA and were routinely < 40 microA. Microstimulation was delivered either in the dark with no fixation, 100 ms after a fixation target was extinguished, or during maintained fixation of a stationary or moving target. Evoked eye movements also were studied under open-loop conditions with the target image stabilized on the retina. 2. Eye movements evoked in the absence of a target rapidly accelerated to a constant velocity that was maintained for the duration of the microstimulation. Evoked eye speeds ranged from 3.7 to 23 deg/s and averaged 11 deg/s. Evoked eye speed appeared to be linearly related to initial eye position with a sensitivity to initial eye position that averaged 0.23 deg.s-1.deg-1. While some horizontal and oblique smooth eye movements were elicited, microstimulation resulted in upward eye movements in 89% of the sites. 3. Evoked eye speed was found to be dependent on microstimulation pulse frequency and current intensity. Within limits, evoked eye speed increased with increases in stimulation frequency or current intensity. For stimulation frequencies < 300-400 Hz, only smooth pursuit-like eye movements were evoked. At higher stimulation frequencies, accompanying saccades consistently were elicited. 4. Feedback of retinal image motion interacted with the evoked eye movements to decrease eye speed if the visual motion was in the opposite direction as the evoked, pursuit-like eye movements. 5. The results implicate rNRTP as part of the neuronal substrate that controls smooth-pursuit eye movements. NRTP appears to be divided functionally into a rostral, pursuit-related portion and a caudal, saccade-related area. rNRTP is a component of a corticopontocerebellar circuit that presumably involves the pursuit area of the frontal eye field and that parallels the middle and medial superior temporal cerebral cortical/dorsalateral pontine nucleus (MT/MST-DLPN-cerebellum) pathway known to be involved also with regulating smooth-pursuit eye movements.

Animals↗

Eye movements cannot explain vibration-induced visual motion and motion aftereffect.

Eye movements are thought to account for a number of visual motion illusions involving stationary objects presented against a featureless background or apparent motion of the whole visual field. We tested two different versions of the eye movement account: (a) the retinal slip explanation and (b) the nystagmus-suppression explanation, in particular their ability to account for visual motion experienced during vibration of the neck muscles, and for the visual motion aftereffect following vibration. We vibrated the neck (ventral sternocleidomastoid muscles, bilaterally, or right dorsal muscles) and measured eye movements in conjunction with perceived illusory displacement of an LED presented in complete darkness (N=10). To test the retinal-slip explanation, we compared the direction of slow eye movements to the direction of illusory motion of the visual target. To test the suppression explanation, we estimated the direction of suppressed slow-phase eye movements and compared it to the direction of illusory motion. Two main findings show that neither actual nor suppressed eye movements cause the illusory motion and motion aftereffect. Firstly, eye movements do not reverse direction when the illusory motion reverses after vibration stops. Secondly, there are large individual differences with regards to the direction of eye movements in observers who all experience a similar visual illusion. We conclude that, rather than eye movements, a more global spatial constancy mechanism that takes into account head movement is responsible for the illusion. The results also argue against the notion of a single central signal that determines both perceptual experience and oculomotor behaviour.

Adult↗