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Eye movement disorders after frontal eye field lesions in humans.

Eye movements were recorded electro-oculographically in three patients with a small ischemic lesion affecting the left frontal eye field (FEF) and in 12 control subjects. Reflexive visually guided saccades (gap and overlap tasks), antisaccades, predictive saccades, memory-guided saccades, smooth pursuit and optokinetic nystagmus (OKN) were studied in the three patients. Staircase saccades and double step saccades were also studied in one of the three patients. For both leftward and rightward saccades, latency in the overlap task (but not in the gap task) and that of correct antisaccades and of memory-guided saccades was significantly increased, compared with the results of controls. There was a significant decrease in the amplitude gain of all rightward saccades programmed using retinotopic coordinates (gap and overlap tasks, predictive and memory-guided saccades), whereas the amplitude gain of corresponding leftward saccades was preserved. Such an asymmetry between leftward and rightward saccades was significant. In the staircase paradigm as well as for the first saccade in the double step paradigm (with the use of retinotopic coordinates in both cases), the amplitude gain of rightward saccades was also significantly lower than that of leftward saccades. Moreover, in the double step paradigm, the amplitude gain of the first rightward saccade was significantly lower than that of the second rightward saccade (programmed using extraretinal signals), which was preserved. The percentage of errors in the antisaccade task did not differ significantly from that of normal subjects. In the predictive saccade paradigm, the percentage of predictive rightward saccades was significantly decreased. The left smooth pursuit gain for all tested velocities, the right smooth pursuit gain for higher velocities, and the left OKN gain were significantly decreased. The results show, for the first time in humans, that the FEF plays an important role in (1) the disengagement from central fixation, (2) the control of contralateral saccades programmed using retinotopic coordinates, (3) saccade prediction and (4) the control of smooth pursuit and OKN, mainly ipsilaterally. In contrast, the left FEF did not appear to be crucial for the control of the only type of saccades programmed using extraretinal signals studied here.

Brain↗

Smooth-pursuit eye movement representation in the primate frontal eye field.

Physiological and behavioral data reported here show an involvement of the primate frontal eye field (FEF) cortex in smooth-pursuit eye movements, in addition to its well-established role in saccadic eye movements. Microstimulation just ventral to the small saccade representation of the FEF elicits eye movements that, in contrast to elicited saccades, have low velocities, continue smoothly without interruption during prolonged stimulation, and are usually directed ipsilaterally to the stimulated hemisphere. Neurons in this region respond in association with smooth-pursuit eye movements and visual motion. Tracking deficits following experimental lesions of the FEF depend critically upon the status of this ventral region: superficial lesions sparing it leave smooth-pursuit eye movements intact, whereas lesions removing it produce substantial deficits in the anticipatory initiation, motion-induced acceleration, asymptotic velocity, and predictive continuation of ipsilateral smooth pursuit.

Animals↗

Effects of linear acceleration on fish behavior and eye movements.

Behavioral responses and eye movements of fish during linear acceleration were reviewed. It is known that displacement of otoliths in the inner ear leads to body movements and/or eye movements. On the ground, the utriculus of the vestibular system is stimulated by otolith displacement caused by gravitational and inertial forces during horizontal acceleration of whole body. When the acceleration is imposed on the fish's longitudinal axis, the fish showed nose-down and nose-up posture for tailward and noseward displacement of otolith respectively. These responses were understood that the fish aligned his longitudinal body axis in a plane perpendicular to the direction of resultant force vector acting on the otoliths. When the acceleration was sideward, the fish rolled around his longitudinal body axis so that his back was tilted against the direction in which the inertial force acted on the otoliths. Linear acceleration applied to fish's longitudinal body axis evoked torsional eye movement. Direction of torsion coincided with the direction of acceleration, which compensate the change of resultant force vector produced by linear acceleration and gravity. Torsional movement of left and right eye coordinated with each other. In normal fish, both sinusoidal and rectangular acceleration of 0.1G could evoke clear eye torsion. Though the amplitude of response increased with increasing magnitude of acceleration up to 0.5 G, the torsion angle did not fully compensate the angle calculated from gravity and linear acceleration. Removal of the otolith on one side reduced the response amplitude of both eyes. The torsion angle evoked by rectangular acceleration was smaller than that evoked by sinusoidal acceleration in both normal and unilaterally labyrinthectomized fish. These results suggest that eye torsion of fish include both static and dynamic components.

Acceleration↗

Brain pathology, eye movement disorders, and basic science of ocular movement.

Diagnosis of eye movement disorders is an important step in evaluation of patients with developmental or acquired brain lesions. A better understanding of the various pathologies requires both a detailed clinical examination and the analysis of eye movement recordings (nystagmus of various types, saccade, and pursuit anomalies). Basic science studies help to interpret these eye movement disorders.

Brain↗

Distinguishing schizophrenic patients from healthy controls by quantitative measurement of eye movement parameters.

BACKGROUND: Eye tracking dysfunction is a putative trait marker for susceptibility to schizophrenia; however, it cannot be recommended as an additional tool for the diagnosis of schizophrenia, due to low sensitivity and specificity. METHODS: To assess the diagnostic potentials of combinations of eye movement paradigms, four smooth pursuit experiments (1: constant velocity of 15 degrees/sec; 2 and 3: combination with either visual or auditory distractors; 4: constant velocity of 30 degrees/sec) and two saccadic eye movement experiments (1: reflexive saccades; 2: voluntary saccades) were conducted. Fourteen patients with residual schizophrenia and 17 healthy controls were studied. Two sets of discriminant analyses (each with the resubstitution and with the "leaving one out" method) were calculated. RESULTS: In the first set, all 10 characteristic variables were included, whereas for the second set, the three most powerful parameters were selected (two from smooth pursuit tasks and one from a voluntary saccade experiment). This procedure provided the best classification results, regarding concordance between clinical diagnoses and eye movement dysfunction (kappa = .67-.80). CONCLUSIONS: Schizophrenic patients of the residual subtype can be differentiated from healthy individuals with considerable criterion validity on the basis of paradigms from two different ocular motor systems.

Adult↗

Ongoing eye movements constrain visual perception.

Eye movements markedly change the pattern of retinal stimulation. To maintain stable vision, the brain possesses a variety of mechanisms that compensate for the retinal consequences of eye movements. However, eye movements may also be important for resolving the ambiguities often posed by visual inputs, because motor commands contain additional spatial information that is necessarily absent from retinal signals. To test this possibility, we used a perceptually ambiguous stimulus composed of four line segments, consistent with a shape whose vertices were occluded. In a passive condition, subjects fixated a spot while the shape translated along a certain trajectory. In several active conditions, the spot, occluder and shape translated such that when subjects tracked the spot, they experienced the same retinal stimulus as during fixation. We found that eye movements significantly promoted perceptual coherence compared to fixation. These results indicate that eye movement information constrains the perceptual interpretation of visual inputs.

Algorithms↗

Electrical activity of superior colliculus associated with eye movements in alert cats.

Eye movement potentials (EMPs) were recorded from the superior colliculus (SC) of awake cats. When cats moved the eyes in front of a patterned or diffuse field, the EMP appeared as a sequence of the three components; EMP1, EMP2 and EMP3. The EMP1-EMP2-complex occurred while the eyes were moving and it had a wave form similar to that of the flash-evoked response. The EMP3 was seen 30-50 msec after the end of eye movement. In dark only the EMP3 was observed; its delay after the end of eye movement was as large as 70 msec. By placing electrodes at various depths of SC, the EMP was found to originate in the superficial layers. In immobilized, encéphale isolé cats, placed in dark or made blind by enucleation, the followings were observed; 1) Passive movements of the eye did not give rise to any significant electrical activity of SC. 2) Spontaneous mass activity of the oculomotor nucleus, that would result in eye movements if the cats were not paralysed, was followed by sharp wave activity of SC and this was the same after the visual cortex had been ablated bilaterally. From these findings it is concluded for the EMPs of awake cats that the EMP1-EMP2-complex is a response to retinal impulses caused by movements of images, whereas the EMP3 is due to an extraretinal factor such as the corollary mechanism. Besides the EMPs as described above, some activity was found to occur about 150 msec before the onset of eye movement. This EMP, called early EMP, seemed to appear when it was followed by eye movements with relatively high speeds.

Action Potentials↗

Eye movements induced by electrical stimulation of the frontal eye fields of marmosets and squirrel monkeys.

Two or three sites in the region of the frontal lobe of marmoset and squirrel monkey are definable as frontal eye fields (FEF) on the basis of electrical stimulation which results in slow and/or saccadic eye movements. In the squirrel monkey these are located on the banks of the arcuate and principal sulci; in the marmoset at estimated analogous locations. Eye ball deviations result from stimulation of these sites with a return to initial position at the end of stimulation. The induced eye movements included large, slow eye movements of varying speeds, micro- and macro-saccades, pendular eye movements, and saccadic stairways. The direction of these eye movements depended on the site stimulated. Both smooth slow and saccadic eye movements were obtained even with stimulation of the same site, depending on the stimulus intensity and the depth of anesthesia. A precise relationship between the site stimulated and the type of eye movement elicited could not be established in these species. Effective stimulus was 300 Hz frequency, 0.2-0.5 ms pulse duration, 0,05-0.05 mA current, and trains of 4-160 pulses. Characteristically, sites for induction of saccadic eye movements were frontal to sites for induction of slow eye movements.

Animals↗

Cytomorphometric changes in rat brain neurons after rapid eye movement sleep deprivation.

Rapid eye movement sleep plays a vital role in the survival of animals. Its deprivation causes alterations in brain functions and behaviors including activities of important enzymes, neurotransmitter levels, impairment of neural excitability and memory consolidation. However, there was a lack of knowledge regarding the effects of rapid eye movement sleep deprivation on neuronal morphology that may get affected much earlier than any permanent damage to the neurons. In the present study, some of these issues have been addressed by studying the effects of rapid eye movement sleep deprivation on various morphological parameters viz. neuronal perimeter, area and shape of neurons located in brain areas known to regulate rapid eye movement sleep and as a control in other brain areas which do not regulate rapid eye movement sleep. The results showed that rapid eye movement sleep deprivation differentially affected neurons depending on their physiological correlates of rapid eye movement sleep and neurotransmitter content. The effects could be reversed if the animals were allowed to recover from rapid eye movement sleep loss or by applying alpha1-adrenergic antagonist, prazosin. The findings in rats support reported data and help explaining previous observations.

Adrenergic alpha-1 Receptor Antagonists↗

Eyelid movements measured by Nightcap predict slow eye movements during quiet wakefulness in humans.

A precipitous decline in eyelid movements (ELMs) has been shown to be a highly reliable indicator of sleep onset. While ELMs correlate well with eye movements during waking and rapid eye movement (REM) sleep, the eye sensor remains silent during the period of slow eye movements (SEMs) typical of sleep onset. If the ELM density (e.g. ELMs per minute) dropped simultaneously with the appearance of SEMs prior to sleep onset, it could be a promising tool for identifying decreases in alertness prior to overt sleep onset. The present study was designed to determine whether the presence of SEMs in the transitional period preceding stage 1 sleep is reflected in decreases in ELM density. ELM densities were computed for 2.5-s epochs with and without SEMs, as well as for 15-s epochs. Decreases in ELM density not only were an excellent correlate of the appearance of SEMs during wakefulness with closed eyes, but also a good predictor of their occurrence (c. 82% accuracy) at a time resolution of 2.5 s. Based on these results, we conclude that ELM density reliably predicts moderate changes in the level of alertness during quiet wakefulness.

Adult↗

Real time binocular detection of horizontal vertical and torsional eye movements by an infra red video-eye tracker.

So far it has only been possible to detect horizontal and vertical eye movements during real time; torsional eye movements have to be calculated from video-tape that have been processed several times, and each investigation takes considerable time. We developed a real time binocular infra red video eye tracker for routine clinical examination which allows real time graphical visualization of horizontal, vertical and torsional eye movements with a bandwidth of 12 Hz (25 Hz refresh frequency). Eye movements in three dimensions can now be obtained as easy as has been done before with electronystagmography for the horizontal component. In contrast to previous systems described, we analyze only those parts of the video-frame that are essential for detection of all three eye movements. This results in a huge data-reduction accompanied by a speed-up of analysis. The black pupil centre is detected by an iterative process of analyzing several horizontal and vertical lines of the even video field. The algorithm searches for regions of minimal intensity. Horizontal and vertical eye rotation is then calculated and transformed into Fick's coordinates. Based upon the centre of the pupil four concentric segments are defined in the iris to detect torsion and to enable correction for false rotation. Torsion is detected by cross-correlation of each segment and estimated from the outcome of the correlation coefficients. All transformations are established by predefined stored transformation matrices thus making real time calculations for transformation unnecessary. After corrections for eccentricity and assuming that fixation of the subjects tested was perfect, accuracy appears to be within 0.3 degrees-1.0 degrees for all three dimensions. Linearity after all corrections is within 4-9% within a visual field of 60 x 40 degrees (horizontal x vertical).

Computer Graphics↗

Slow and saccadic eye movements evoked by microstimulation in the supplementary eye field of the cebus monkey.

1. Intracortical microstimulation was used to map the supplementary eye field (SEF) in eight hemispheres of five Cebus apella monkeys. Monkeys were immobilized during experiments with Telazol (tiletamine HCl and zolazepam HCl), a dissociative anesthetic agent that was demonstrated to have no significant effect on microstimulation-induced eye movement parameters compared with similar experiments in alert, behaviorally trained monkeys. The functional subregions were defined with the use of low-threshold current (< or = 50 microA). Electrically elicited eye movements were videotaped and quantified. Both slow and saccadic eye movements were reliably evoked at low threshold by microstimulation in each of eight hemispheres studied. The two types of eye movements were clearly distinguished by their significantly different duration and velocity (P < 0.0001) and their different responses to long stimulus trains. The results strongly support the proposal that the SEF produces not only saccadic eye movements as previously reported but also slow (pursuit) eye movements.

Anesthetics↗

Oculomotor function in Wernicke-Korsakoff's syndrome: II. Smooth pursuit eye movements.

Smooth pursuit eye movements were studied in three patients with alcoholic Korsakoff's syndrome, one with Wernicke's encephalopathy, and an age-matched control. Horizontal smooth pursuit eye movements were abnormal in all patients: peak eye velocity and the ability to sustain smooth eye velocity were reduced. Also, smooth pursuit gain began to decrease at relatively low target velocities (i.e., 8-10 degrees). These data demonstrate a severe disturbance in smooth pursuit function long after the clinically apparent oculomotor abnormalities have passed.

Aging↗

Computer simulation of overshoot in saccadic eye movements.

The human horizontal eye movement system produces quick, precise, conjugate eye movements called saccades. These are important in normal vision. For example, reading tasks exclusively utilize saccadic eye movements. The majority of saccades have dynamic overshoot. The amplitude of this overshoot is independent of saccadic amplitude, and is such that it places the image of the stimulus within the retinal region of maximum acuity within a minimum of time. A computer based model of the saccadic mechanisms was used to study the origin of this overshoot. It was discussed that dynamic overshoot cannot be attributed to biomechanism properites of the eye movement mechanism, but must instead be explained by variations in the controlling nervous activity. The form of this neural controller signal is very similar to that required for a time optimal response of an inertial system.

Computers↗

Perceived visual motion as effective stimulus to pursuit eye movement system.

Human eye tracking of a foveal afterimage during angular head oscillation in the dark produced smooth eye movements exceeding those for normal vestibular nystagmus, and a reduction in the frequency of the fast phase component of nystagmus eye movements. These results may support a closed loop extension of the corollary discharge theory, with oculomotor commands based on perceived object velocity.

Attention↗

The effect of stimulus predictability and age on human tracking eye movements.

Human tracking eye movements were recorded in healthy volunteers of varying age. The target motion was either predictable (pure sinusoids or sinusoidal frequency sweep) or unpredictable (pseudorandomized). Gain and phase values for the tracking and the smooth pursuit eye movement signal were calculated as a function of target frequency from 0.2 to 2 Hz. Max. target velocity was 20 or 40 deg/sec. In the low frequency area the smooth pursuit gain for predictable stimulation was higher than the gain elicited by the pseudorandomized stimulation. In the high-frequency area, gain values did not differ significantly. For the predictable stimulation, the phase of the smooth component was always a lag, increasing with increasing frequency. At low frequencies of the pseudorandomized signal a phase lead was observed. At higher frequencies the lead turned into a lag that was greater than for the predictable stimulation. The young and old groups showed reduced smooth pursuit gain values, compared with the main group specially when the target waveform was pseudorandomized. The reduced smooth pursuit ability was mostly compensated for by an increased amount of saccades.

Adult↗

Covert attention and eye movements during reading.

Eye movements were monitored during the reading of spatially transformed text in order to examine covert attentional processes in reading. In some conditions, the sequence of letters within a word was congruent with (i.e. in the same direction as) the sequence of words in the sentence; in other conditions the direction of letters within words and the direction of words in the sentence were incongruent. In addition, the window of visible text was varied so that in some conditions only the fixated word (and all preceding words) were visible, whereas in other conditions the fixated word and the succeeding word were both visible. Readers were able to extract more parafoveal information from text when the words themselves were normal than when the letters within the words were transformed. However, with practice, readers were able to use some parafoveal information even when the words were transformed. The most important finding was that the congruity of the word and letter order had no reliable effect on the ability to extract parafoveal information and influenced reading performance only when the words themselves were normal. We conclude that covert attention in reading is not a letter-by-letter scan that sweeps across the page, but either an asymmetric spotlight held constant on each fixation or a shifting of an attentional spotlight extending across multiletter units (possibly words) with the direction of shifts of attention closely coupled to the direction of eye movements.

Attention↗