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Progressive cortical synchronization of ponto-geniculo-occipital potentials during rapid eye movement sleep.

Phasic events, termed ponto-geniculo-occipital potentials, appear in the brainstem, thalamus and cerebral cortex during rapid eye movement sleep. In the cat, the species of choice for ponto-geniculo-occipital studies, these field potentials are usually recorded from the lateral geniculate thalamic nucleus and visual cortex. However, the fact that brainstem cholinergic neurons play a crucial role in the transfer of ponto-geniculo-occipital potentials to the thalamus, coupled with the evidence that mesopontine tegmental neurons project to virtually all thalamic nuclei, together explain why ponto-geniculo-occipital potentials are recorded over widespread territories, beyond the visual thalamocortical system. Here we demonstrate, by means of multi-site unit and field potential recordings from sensory, motor and association cortical areas in behaving cats, that: (i) ponto-geniculo-occipital potentials appear synchronously over the neocortex; and (ii) that their cortical synchronization develops progressively from the period preceding rapid eye movement sleep by 30-90 s (pre-rapid eye movement), to reach the highest degree of intracortical coherence during later epochs of rapid eye movement sleep. We propose that the widespread coherence of cortical ponto-geniculo-occipital potentials underlies the synchronization of fast oscillations (30-40 Hz) during rapid eye movement sleep over many, functionally distinct cortical territories implicated in dreaming, as brainstem-induced ponto-geniculo-occipital-like potentials are consistently followed by such fast oscillations.

Animals↗

Discharge properties of neurons in the monkey thalamus tested with angular acceleration, eye movement and visual stimuli.

Monkeys were trained to make visually evoked eye movements while undergoing simultaneous head rotation. Single units were recorded in the pregeniculate nucleus (PGN). PGN neurons discharged during each saccade, but there was no change in activity with horizontal head acceleration or with various combinations of head and smooth pursuit eye movements as previously described in the cat. Therefore, the anatomical homology between LGNv and PGN does not appear to have a neurophysiological basis. Neurons in the oral part of VPL or occasionally in VPI discharged as a function of head velocity but not with saccades, smooth pursuit or fixation eye movements, nor after brief light flashes or during smooth pursuit across structured backgrounds. This suggests that VPLo and VPI are only vestibular relay nuclei and not concerned with vestibular/visual or vestibular/oculomotor interactions.

Animals↗

Visual motion due to eye movements helps guide the hand.

Movement of the body, head, or eyes with respect to the world creates one of the most common yet complex situations in which the visuomotor system must localize objects. In this situation, vestibular, proprioceptive, and extra-retinal information contribute to accurate visuomotor control. The utility of retinal motion information, on the other hand, is questionable, since a single pattern of retinal motion can be produced by any number of head or eye movements. Here we investigated whether retinal motion during a smooth pursuit eye movement contributes to visuomotor control. When subjects pursued a moving object with their eyes and reached to the remembered location of a separate stationary target, the presence of a moving background significantly altered the endpoints of their reaching movements. A background that moved with the pursuit, creating a retinally stationary image (no retinal slip), caused the endpoints of the reaching movements to deviate in the direction of pursuit, overshooting the target. A physically stationary background pattern, however, producing retinal image motion opposite to the direction of pursuit, caused reaching movements to become more accurate. The results indicate that background retinal motion is used by the visuomotor system in the control of visually guided action.

Brain↗

Cortical wave amplitude and eye movement direction are correlated in REM sleep but not in waking.

Eye movement related potentials were recorded bilaterally from the lateral geniculate body and visual cortex in five cats during REM sleep and waking. During REM sleep polygraphic recordings showed that these potentials have significantly greater amplitudes (primary waves) in the visual cortex and lateral geniculate body ipsilateral to the direction of the corresponding rapid eye movements. This amplitude difference was seen as a mean increase of 38.2% over the contralateral side and was found in 94.0% of wave pairs. The correlation between horizontal eye movement direction and amplitude of the waves is specific to REM sleep as there were no measurable differences in the waking state. These results are discussed with respect to possible mechanisms and functions of this lateralized wave system and its state dependency.

Animals↗

Surgeon offsets and dynamic eye movements in laser refractive surgery.

PURPOSE: To determine the amount of static and dynamic pupil decentrations that occur during laser refractive surgery. SETTING: The Center of Visual Science and the Department of Ophthalmology, University of Rochester, Rochester, New York, USA. METHODS: The surgeon's accuracy in aligning the pupil center with the laser center axis was measured when engaging the eye-tracker in 17 eyes receiving conventional laser in situ keratomileusis (LASIK) procedures (Technolas 217z; Bausch & Lomb). Eye movements were measured subsequently during the treatment in 10 eyes using a pupil camera operating at 50 Hz. Temporal power spectra were calculated from the eye movement measurements. RESULTS: The mean pupil misalignment by the surgeon at the beginning of the procedure was 206.1 microm +/- 80.99 (SD) (with respect to the laser center). The laser center was typically misaligned below (inferiorly) and to the left (nasally and temporally in left and right eyes, respectively) of the pupil [corrected] center. Small amounts of cyclotorsion were observed during the ablation (<2 degrees). The mean magnitude of dynamic pupil decentration from the laser center during treatment was 227.0 +/- 44.07 microm. The mean standard deviation of eye movements was 65.7 +/- 25.64 microm. Temporal power spectra calculated from the horizontal and vertical changes in eye position during the ablation were similar. Ninety-five percent of the total power of the eye movements was contained in temporal frequencies up to 1 Hz, on average, in both directions. CONCLUSIONS: Most eye movements during LASIK are slow drifts in fixation. An eye-tracker with a 1.4 Hz closed-loop bandwidth could compensate for most eye movements in conventional or customized ablations.

Calibration↗

Digital measurement of torsional eye movement due to postural change and its effect on reading performance.

PURPOSE: Posture induced torsional eye movements have rarely been investigated. The current study made use of digital imaging technology to measure the cycloduction resulting from postural change. The effect of cycloduction on reading performance was also investigated. METHODS: Thirty subjects were recruited and pictures of the right eye were captured using a digital camera at three postures, i.e. sitting, 90 degrees tilted to the right and 90 degrees tilted to the left. With the identification of a conjunctival landmark, torsional eye movement was measured. The subjects were then required to read a custom designed near chart while in those three postures, the reading card being rotated 90 degrees clockwise or 90 degrees anti-clockwise, to match with the tilting of the head. The reproducibility of the torsional eye movement and reading performance measure was determined in 12 of those 30 subjects. RESULTS: Incycloduction was induced when tilting to the right and excycloduction when tilting to the left. This method was found to be reproducible with the 95% confidence limits of 0.80 degrees between visits. The mean incycloduction induced was 6.50 degrees (SD 1.51 degrees ) and 6.41 degrees (SD 1.46 degrees ) for excycloduction. No significant difference in amount was demonstrated (paired t-test: t = 0.624, P = 0.538). No significant difference was found in the reading scores at various postures (Repeated measures ANOVA: df = 2, F = 1.881, P = 0.162). CONCLUSIONS: The results presented here demonstrate that we have developed an objective and instantaneous method with good precision, which could be applied in other studies that require the measurement of torsional eye movements. The induced cycloduction did not affect the reading performance.

Adolescent↗

An algorithm separating saccadic from nonsaccadic eye movements automatically by use of the acceleration signal.

An algorithm is described to discriminate automatically between saccades and slow eye movements. Sampled data of the eye position have been used to calculate the momentary acceleration of the eye. The higher acceleration values of the saccadic eye movements as opposed to the slow compensatory or pursuit eye movements served to differentiate between the two. The method is demonstrated by search-coil data in squirrel monkeys.

Algorithms↗

Development of voluntary control of saccadic eye movements. I. Age-related changes in normal children.

To investigate the development of the voluntary control of saccadic eye movement, we examined eye movements in 99 normal children (4-13 years of age). Subjects were asked to fixate a central light for 3-5 s. A target was then presented, either to its right or left. In visually guided saccades, the mean latencies of the child group were longer than those of the adults, and decreased with age until the age of 12 where they reached adult levels. On the other hand, their peak saccadic velocities were not different from those of the adults. In the antisaccade task, they showed a higher rate of directional errors, indicating an inability to suppress reflexive saccades to the target. Mean latencies of correct antisaccades were significantly longer in the children than in the adults. Error rates and antisaccadic latency tended to decrease with age. We also examined the effects of an auditory warning signal during the fixation period and compared the results with those without. The warning stimulus was less effective in children than in adults in both tasks. Similar peak saccadic velocities between children and adults suggest the earlier development of the saccadic burst generator in the brainstem. In contrast, the delayed development of latency of saccades and antisaccades and the error rates of antisaccades suggest delayed maturation of the cerebral cortex, particularly the frontal association areas that are involved in both eye movement control and attentional processes.

Acoustic Stimulation↗

The diagnostic value of eye movement recordings in patients with Huntington's disease and their offspring.

Eye movements were studied with EOG in a large number of Huntington's disease (HD) patients (N = 48) and their offspring (subjects at risk, N = 97). The following oculomotor paradigms were investigated: horizontal and vertical saccades of different sizes (10-80 degrees), smooth pursuit eye movements, optokinetic and vestibular nystagmus. It was found that 75% of the HD patients had pathologically slow saccades. This number rose to 87% if other oculomotor deficits were included. Thus a small, but definite number of HD patients had normal eye movements. This included patients with disease durations of more than 14 years. A comparison with the results of clinical and evoked potential studies on these patients showed that they were clinically only mildly affected but did not form a different group of HD patients. Subjects at risk had statistically (P less than 0.01) slower horizontal saccades (average 427 deg/sec) than the controls (455 deg/sec). For 11% of the subjects at risk horizontal saccade velocity was pathological and altogether 22% had some oculomotor deficits. Subjects with pathological slow horizontal saccades also had a high incidence (64%) of pathological evoked potentials. Thus, subjects with pathological slow horizontal saccades can be considered to have a high risk of developing the clinical symptoms of HD.

Adolescent↗

Effects of early-onset artificial strabismus on pursuit eye movements and on neuronal responses in area MT of macaque monkeys.

In humans, esotropia of early onset is associated with a profound asymmetry in smooth pursuit eye movements. When viewing is monocular, targets are tracked well only when they are moving nasally with respect to the viewing eye. To determine whether this pursuit abnormality reflects an anomaly in cortical visual motion processing, we recorded eye movements and cortical neural responses in nonamblyopic monkeys made strabismic by surgery at the age of 10-60 d. Eye movement recordings revealed the same asymmetry in the monkeys' pursuit eye movements as in humans with early-onset esotropia. With monocular viewing, pursuit was much stronger for nasalward motion than for temporalward motion, especially for targets presented in the nasal visual field. However, for targets presented during ongoing pursuit, temporalward and nasalward image motion was equally effective in modulating eye movement. Single-unit recordings made from the same monkeys, under anesthesia, revealed that MT neurons were rarely driven binocularly, but otherwise had normal response properties. Most were directionally selective, and their direction preferences were uniformly distributed. Our neurophysiological and oculomotor measurements both suggest that the pursuit defect in these monkeys is not due to altered cortical visual motion processing. Rather, the asymmetry in pursuit may be a consequence of imbalances in the two eyes' inputs to the "downstream" areas responsible for the initiation of pursuit.

Animals↗

Hourly fetal urinary production rate and blood velocity waveforms in the fetal renal artery relative to fetal heart rate pattern and fetal eye movements in normal pregnancies at 30-32 weeks of gestation.

In 14 normal pregnancies at 30-32 weeks of gestation, bloodflow velocity waveforms in the fetal renal artery and fetal micturition were studied in relation to fetal heart rate pattern (FHRP) and fetal eye movements. PI in the fetal renal artery was significantly lower in the presence of FHRP-B and eye movements separately and in combination compared with FHRP-A and absent eye movements separately and in combination. Fetal urinary output was higher in the absence of fetal eye movements. There was an inverse relationship between renal artery PI and fetal heart rate. However, since mean fetal heart rate in the various state variables was not essentially different, no standardization for heart rate was necessary. No correlation existed between renal artery PI and fetal urinary production rate. It can be concluded that in the normal fetus at 30-32 weeks of gestation, renal artery PI demonstrates a significant reduction in the presence of FHRP-B and fetal eye movements separately and in combination, suggesting reduced downstream impedance at renal level. However, the decrease in fetal micturition in the presence of fetal eye movements, suggests that reduced urinary output is determined by changes in fetal central nervous system activity rather than by changes in fetal renal perfusion.

Adult↗

Eye movements and industrial inspection.

A brief survey is given of the eye movement parameters which contribute to visual search strategies. The literature on eye movements and industrial inspection is reviewed and some recent studies carried out at the University of Birmingham into the eye movements underlying four inspection tasks are described. These tasks included the inspection of tin cans, electrical edge connectors, men's briefs and woven fabric.

Journal Article↗

Interaction of visual and non-visual signals in the initiation of smooth pursuit eye movements in primates.

The initiation of smooth pursuit eye movements (PEM) by visual and non-visual signals was analysed in humans and monkeys. While PEM latency ranged around 150 ms when a purely visual target was provided, it often dropped to about 0 ms, or even became negative, when target movement was coupled to the subject's arm; this suggests that signals about the intention to move the arm can be evaluated for PEM control. Eye movements always started in the visually correct direction, independent of the sign of coupling between arm and target; from this we conclude that intentional signals are not mere triggers, but also convey directional information. Short-latency PEM trials were intermixed with those characterized by normal latencies, which often resulted in bimodal latency distributions; this suggests that visual and intentional signals compete for the control of PEM.

Animals↗

Activity of substantia nigra pars reticulata neurons during smooth pursuit eye movements in monkeys.

Nuclei within the basal ganglia (BG), in particular the substantia nigra pars reticulata (SNr), subthalamic nucleus (STN) and caudate nucleus, are known to be involved in the generation of rapid or saccadic eye movements. Neurons in the SNr are active tonically and generally show a pause, but also increase, in discharge rate, for the appearance of visual stimuli and the generation of saccades. Recent experimental results in oculomotor regions of the brainstem reveal overlap in the neuronal pathways used for saccades and smooth pursuit, or slow tracking, eye movements. Whether the overlap of processing for saccades and pursuit extends to the oculomotor BG is unknown. In the present report, we were interested in whether the overlap between the pursuit and saccadic systems extends into the oculomotor BG. Using single-neuron recording and electrical stimulation techniques, we tested whether neurons within the saccade portion of the BG, the SNr, could be involved in smooth pursuit eye movements. Monkeys were required to follow visual targets with either a smooth eye movement or a saccade while we recorded from SNr neurons. We report here on SNr neuronal activity that was modulated during the performance of visually guided saccades and also during the initiation and the maintenance of smooth pursuit eye movements. Importantly, the modulation of neuronal activity during pursuit was present even when catch-up saccades were absent. The majority of SNr neurons was active tonically and their discharge ceased during pursuit, although some neurons also increased their discharge rate during smooth pursuit, similar to the behaviour reported for saccades. We also found that electrical stimulation of the SNr during the initiation of pursuit suppressed ipsiversive and, in some cases, enhanced contraversive pursuit. Our combined recording and stimulation results are consistent with the hypothesis that the overlap between the pursuit and saccadic systems extends, at least somewhat, into the BG and that the signal conveyed by the SNr can be used by the pursuit system. Like the signal for saccades, the SNr may provide a permissive disinhibition for pursuit eye movements. We hypothesize that alterations in this signal in BG diseased states such as Parkinson's may explain in part the deficits observed in smooth pursuit eye movements of these patients.

Action Potentials↗

The selective disruption of spatial working memory by eye movements.

In the late 1970s/early 1980s, Baddeley and colleagues conducted a series of experiments investigating the role of eye movements in visual working memory. Although only described briefly in a book, these studies have influenced a remarkable number of empirical and theoretical developments in fields ranging from experimental psychology to human neuropsychology to nonhuman primate electrophysiology. This paper presents, in full detail, three critical studies from this series, together with a recently performed study that includes a level of eye movement measurement and control that was not available for the older studies. Together, the results demonstrate several facts about the sensitivity of visuospatial working memory to eye movements. First, it is eye movement control, not movement per se, that produces the disruptive effects. Second, these effects are limited to working memory for locations and do not generalize to visual working memory for shapes. Third, they can be isolated to the storage/maintenance components of working memory (e.g., to the delay period of the delayed-recognition task). These facts have important implications for models of visual working memory.

Adolescent↗

Cancelling of pursuit and saccadic eye movements in humans and monkeys.

The countermanding paradigm provides a useful tool for examining the mechanisms responsible for cancelling eye movements. The key feature of this paradigm is that, on a minority of trials, a stop signal is introduced some time after the appearance of the target, indicating that the subject should cancel the incipient eye movement. If the delay in giving the stop signal is too long, subjects fail to cancel the eye movement to the target stimulus. By modeling this performance as a race between a go process triggered by the appearance of the target and a stop process triggered by the appearance of the stop signal, it is possible to estimate the processing interval associated with cancelling the movement. We have now used this paradigm to analyze the cancelling of pursuit and saccades. For pursuit, we obtained consistent estimates of the stop process regardless of our technique or assumptions--it took 50-60 ms to cancel pursuit in both humans and monkeys. For saccades, we found different values depending on our assumptions. When we assumed that saccade preparation was under inhibitory control up until movement onset, we found that saccades took longer to cancel (humans: approximately 110, monkeys: approximately 80 ms) than pursuit. However, when we assumed that saccade preparation includes a final "ballistic" interval not under inhibitory control, we found that the same rapid stop process that accounted for our pursuit results could also account for the cancelling of saccades. We favor this second interpretation because cancelling pursuit or saccades amounts to maintaining a state of fixation, and it is more parsimonious to assume that this involves a single inhibitory process associated with the fixation system, rather than two separate inhibitory processes depending on which type of eye movement will not be made. From our behavioral data, we estimate that this ballistic interval has a duration of 9-25 ms in monkeys, consistent with the known physiology of the final motor pathways for saccades, although we obtained longer values in humans (28-60 ms). Finally, we examined the effect of trial sequence during the countermanding task and found that pursuit and saccade latencies tended to be longer if the previous trial contained a stop signal than if it did not; these increases occurred regardless of whether the preceding trial was associated with the same or different type of eye movement. Together, these results suggest that a common inhibitory mechanism regulates the initiation of pursuit and saccades.

Adolescent↗

Saccadic eye movement programming in unilateral neglect.

The present study examined the eye movements made by patients with unilateral neglect under fixation gap and overlap conditions. The prior offset of fixation in a + 100 msec gap condition did not produce an increase in the numbers of contralesional saccades made by 3 out of 4 patients. This finding is incompatible with the view that the deficit in producing contralesional saccades reflects an inability to disengage visual attention from fixation. A significant reduction in saccade latency was, however, obtained in the gap condition ('gap effect'). The latency reduction in the gap condition is consistent with models which attribute the gap effect to warning signal effects and the release of an ocular rather than an attentional disengagement mechanism. Saccade latency was not increased when two targets were presented bilaterally and simultaneously in both hemifields (in contrast to the increase in latency shown by normal subjects). The lack of a normal 'bilateral target effect' in neglect is attributed to an imbalance in the level of activity in the saccadic system. Three patients showed visual 'extinction' and did not make saccades to the contralesional bilateral targets. In contrast R.R. who shows object-based neglect did not show extinction and made saccades to the contralesional bilateral targets. This suggests that visual extinction may be influenced by the form of neglect shown by the patient. The effects on saccade amplitude of presenting two targets in the same hemifield were also examined in a global effect task. One patient showed a much greater global effect than normal when pairs of targets were presented in his ipsilesional hemifield. In contrast R.R. showed a normal magnitude global effect. It appears that the form of neglect shown by a patient is a factor that influences their eye movement behaviour on simple saccade tasks and these eye movement abnormalities cannot be accounted for by a deficit of attentional disengagement.

Cognition Disorders↗

Properties of cerebellar fastigial neurons during translation, rotation, and eye movements.

The most medial of the deep cerebellar nuclei, the fastigial nucleus (FN), receives sensory vestibular information and direct inhibition from the cerebellar vermis. We investigated the signal processing in the primate FN by recording single-unit activities during translational motion, rotational motion, and eye movements. Firing rate modulation during horizontal plane translation in the absence of eye movements was observed in all non-eye-movement-sensitive cells and 26% of the pursuit eye-movement-sensitive neurons in the caudal FN. Many non-eye-movement-sensitive cells recorded in the rostral FN of three fascicularis monkeys exhibited convergence of signals from both the otolith organs and the semicircular canals. At low frequencies of translation, the majority of these rostral FN cells changed their firing rates in phase with head velocity rather than linear acceleration. As frequency increased, FN vestibular neurons exhibited a wide range of response dynamics with most cells being characterized by increasing phase leads as a function of frequency. Unlike cells in the vestibular nuclei, none of the rostral FN cells responded to rotational motion alone, without simultaneously exhibiting sensitivity to translational motion. Modulation during earth-horizontal axis rotation was observed in more than half (77%) of the neurons, although with smaller gains than during translation. In contrast, only 47% of the cells changed their firing rates during earth-vertical axis rotations in the absence of a dynamic linear acceleration stimulus. These response properties suggest that the rostral FN represents a main processing center of otolith-driven information for inertial motion detection and spatial orientation.

Action Potentials↗