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Existence in the nucleus incertus of the cat of horizontal-eye-movement-related neurons projecting to the cerebellar flocculus.

1. Properties of nucleus incertus (NIC) neurons projecting to the cerebellar flocculus were studied in alert cats by using chronic unit and eye movement recording and antidromic activation. Projection of these neurons onto the flocculus was verified with retrograde transport of horseradish peroxidase after injections in the flocculus. 2. Bipolar stimulation electrodes were implanted into the "middle" zone of each flocculus because this zone is known to be involved in the control of horizontal eye movements. The dorsomedial aspect of the pontine tegmentum was explored with microelectrodes during stimulation of both flocculi. The majority of neurons antidromically activated from the flocculus were found in the caudal part of the NIC. 3. Of the 69 neurons activated from the flocculus, 44 were classified as burst-tonic (BT) neurons; 34 discharged in relation with horizontal movements of the eye, 10 in relation with vertical movements. Of the 14 remaining neurons, 6 were not related to eye movements and 8 were classified as burst neurons. The BT neurons of the NIC displayed a great sensitivity to both horizontal eye position and horizontal eye velocity. 4. This study demonstrates the presence of a new group of horizontal eye movement related BT neurons situated in the NIC. The fact that they project to the horizontal floccular zone emphasizes the importance of the functional specialization of the different Purkinje cell zones.

Animals↗

Effects of frontal eye field and superior colliculus ablations on eye movements.

Two parallel neural pathways are primarily responsible for the control of saccadic eye movements--one mediated through the frontal eye fields and the other through the superior colliculus. When both pathways are disrupted, control of saccadic eye movements is lost. Disruption of either pathway alone produces only subtle deficits.

Animals↗

Eye movements and picture processing during recognition.

Eye movements were monitored during a recognition memory test of previously studied pictures of full-color scenes. The test scenes were identical to the originals, had an object deleted from them, or included a new object substituted for an original object. In contrast to a prior report (Parker, 1978), we found no evidence that object deletions or substitutions could be recognized on the basis of information acquired from the visual periphery. Deletions were difficult to recognize peripherally, and the eyes were not attracted to them. Overall, the amplitude of the average saccade to the critical object during the memory test was less than 4.5 degrees of visual angle in all conditions and averaged 4.1 degrees across conditions. We conclude that information about object presence and identity in a scene is limited to a relatively small region around the current fixation point.

Color Perception↗

Selective activation of visual cortex neurons by fixational eye movements: implications for neural coding.

During normal vision, when subjects attempt to fix their gaze on a small stimulus feature, small fixational eye movements persist. We have recorded the impulse activity of single neurons in primary visual cortex (V1) of macaque monkeys while their fixational eye movements moved the receptive-field activating region (AR) over and around a stationary stimulus. Three types of eye movement activation were found. (1) Saccade cells discharged when a fixational saccade moved the AR onto the stimulus, off the stimulus, or across the stimulus. (2) Position/drift cells discharged during the intersaccadic (drift) intervals and were not activated by saccades that swept the AR across the stimulus without remaining on it. To activate these neurons, it was essential that the AR be placed on the stimulus and many of these cells were selective for the sign of contrast. They had smaller ARs than the other cell types. (3) Mixed cells fired bursts of activity immediately following saccades and continued to fire at a lower rate during intersaccadic intervals. The tendency of each neuron to fire transient bursts or sustained trains of impulses following saccades was strongly correlated with the transiency of its response to stationary flashed stimuli. For one monkey, an extraretinal influence accompanying fixational saccades was identified. During natural viewing, the different eye movement classes probably make different contributions to visual processing. Position/drift neurons are well suited for coding spatial details of the visual scene because of their small AR size and their selectivity for sign of contrast and retinal position. However, saccade neurons transmit information that is ambiguous with respect to the spatial details of the retinal image because they are activated whether the AR lands on a stimulus contour, or the AR leaves or crosses the contour and lands in another location. Saccade neurons may be involved in constructing a stable world in spite of incessant retinal image motion, as well as in suppressing potentially confusing input associated with saccades.

Action Potentials↗

Shared response preparation for pursuit and saccadic eye movements.

Tracking a single target in the visual world requires coordination between pursuit and saccadic eye movements. The constraints imposed on pursuit and saccade decisions by visual processing and response preparation are difficult to compare because latency differences between the two movements provide different amounts of visual sampling time. The present study compares pursuit and saccade decisions when visual processing was directly manipulated. Human observers were asked to select between two stationary stimuli presented simultaneously at two different locations based on which had the higher contrast. The stimuli were presented for a brief, variable interval and then occluded by masks. Because the masks moved horizontally and were offset vertically, subjects were obliged to make both pursuit and saccadic eye movements to track the mask covering the target stimulus. For each of the exposure durations, we constructed oculometric curves for pursuit and saccades. We found that both systems had similar oculometric thresholds and response biases. The initial pursuit decisions differed from the subsequent saccade decisions on 1-13% of the trials but were the same more often than predicted by independent mechanisms. Moreover, pursuit reversed direction on discordant trials, so that the pursuit decision always matched that of the saccade by the time the saccade was started. These results support the view that, in addition to overlap in early visual areas and the final motor pathways, the pursuit and saccadic systems share processing at the level of response preparation. This shared processing may help ensure the coordination of pursuit and saccadic eye movements in selecting a single target.

Adult↗

Directional organization of eye movement and visual signals in the floccular lobe of the monkey cerebellum.

The floccular lobe of the monkey is critical for the generation of visually-guided smooth eye movements. The present experiments reveal physiological correlates of the directional organization in the primate floccular lobe by examining the selectivity for direction of eye motion and visual stimulation in the firing of individual Purkinje cells (PCs) and mossy fibers. During tracking of sinusoidal target motion along different axes in the frontoparallel plane, PCs fell into two classes based on the axis that caused the largest modulation of simple-spike firing rate. For "horizontal" PCs, the response was maximal during horizontal eye movements, with increases in firing rate during pursuit toward the side of recording (ipsiversive). For "vertical" PCs, the response was maximal during eye movement along an axis just off pure vertical, with increases in firing rate during pursuit directed downward and slightly contraversive. During pursuit of target motion at constant velocity, PCs again fell into horizontal and vertical classes that matched the results from sinusoidal tracking. In addition, the directional tuning of the sustained "eye velocity" and transient "visual" components of the neural responses obtained during constant velocity tracking were very similar. PCs displayed very broad tuning approximating a cosine tuning curve; the mean half-maximum bandwidth of their tuning curves was 170-180 degrees. Other cerebellar elements, related purely to eye movement and presumed to be mossy fibers, exhibited tuning approximately 40 degrees narrower than PCs and had best directions that clustered around the four cardinal directions. Our data indicate that the motion signals encoded by PCs in the monkey floccular lobe are segregated into channels that are consistent with a coordinate system defined by the vestibular apparatus and eye muscles. The differences between the tuning properties exhibited by PCs compared with mossy fibers indicate that a spatial transformation occurs within the floccular lobe.

Animals↗

[Eye movement: experimental and clinical study using cine mode MRI].

As a special lecture at the 96th Annual Congress of the Japanese Ophthalmological Society in 1992, we presented experimental and clinical studies on eye movement using magnetic resonance imaging (MRI). MRI is more valuable than X-ray CT in defining external muscle anatomy in the orbit in a variety of pathological changes, and is free from the danger of radiation. Cine mode MRI, which was originally developed to observe cardiovascular function, was utilized experimentally for observation of eye movement. We invented two methods to produce a series of photographs. In the first method, "the moving eye method", the subjects were asked to perform 256 or 512 vertical and horizontal eye movements in synchronization with a sound trigger. In the second, "the fixed eye method", the subjects were asked to gaze at individual points in a 5 to 7 point sequence as indicated by the investigator. In both methods MRI data was generated and the resulting series of photographs were observed in cine mode on a television monitor. Displaying the MRI generated photographs in cine mode enables direct observation of the movement of the eye, optic nerve and extraocular muscles within the orbit. The moving eye method was more accurate in its portrayal of the actual movement of the eye, but the large number of eye movements, at least 256 movements, required about 5 minutes, and caused physical and mental fatigue for the subjects. The fixed eye method did not show actual movement of the eye, but was less fatiguing than the former method, and avoided several of its problems. A large number of ophthalmoplegia cases such as abducens palsy, oculomotor palsy, fracture of the orbital wall, foreign bodies in the orbit, Duane's syndrome, endocrine myopathy, esotropia, exotropia, etc., were examined using the fixed eye method of cine mode MRI. Using this technique, anatomical changes, such as atrophy or hypertrophy and the functional state demonstrated by the contraction and relaxation of the extraocular muscles could be investigated. Transposition of the extraocular muscles, an effective treatment for paralytic squint, was reexamined and a new surgical procedure was developed for abducens palsy. The measurement of the length, thickness and weight of the extraocular muscles from the MRI-generated photographs allowed for more accurate analysis. The three dimensional reconstructive method of MRI is very valuable for diagnosis in the orbit and is expected to contribute to future progress.

Adult↗

Experience-dependent changes in cerebral functional connectivity during human rapid eye movement sleep.

One function of sleep is hypothesized to be the reprocessing and consolidation of memory traces (Smith, 1995; Gais et al., 2000; McGaugh, 2000; Stickgold et al., 2000). At the cellular level, neuronal reactivations during post-training sleep in animals have been observed in hippocampal (Wilson and McNaughton, 1994) and cortical (Amzica et al., 1997) neuronal populations. At the systems level, using positron emission tomography, we have recently shown that some brain areas reactivated during rapid-eye-movement sleep in human subjects previously trained on an implicit learning task (a serial reaction time task) (Maquet et al., 2000). These cortical reactivations, located in the left premotor area and bilateral cuneus, were thought to reflect the reprocessing--possibly the consolidation--of memory traces during post-training rapid-eye-movement sleep. Here, the experience-dependent functional connectivity of these brain regions is examined. It is shown that the left premotor cortex is functionally more correlated with the left posterior parietal cortex and bilateral pre-supplementary motor area during rapid-eye-movement sleep of subjects previously trained to the reaction time task compared to rapid-eye-movement sleep of untrained subjects. The increase in functional connectivity during post-training rapid-eye-movement sleep suggests that the brain areas reactivated during post-training rapid-eye-movement sleep participate in the optimization of the network that subtends subject's visuo-motor response. The optimization of this visuo-motor network during sleep could explain the gain in performance observed during the following day.

Brain Mapping↗

Visual-tracking neurons in area MST are activated during anticipatory pursuit eye movements.

Here, I report that rhesus monkeys are able to generate anticipatory smooth pursuit eye movements in the transient absence of a moving target, if this target moves periodically. The eye velocity before target reappearance was significantly larger if the target trajectory was predictable compared with the control condition consisting of unpredictable target trajectory. Parallel to the registration of the eye movements, single-unit activity was recorded from neurons in the middle superior temporal (MST) area of the two monkeys. The neuronal activity of visual-tracking neurons resembled the observed eye movements, i.e. these neurons increased their activity earlier if the movement of the target was predictable compared with the unpredictable control. These results provide further evidence for the existence of extra-retinal signals in the activity of visual-tracking neurons located in area MST.

Action Potentials↗

Smooth pursuit eye movements and neuropsychological tests in schizophrenic patients: possible involvement of attentional components.

The relationships among different components of frontal lobe dysfunction and voluntary attention were studied. Drug-free schizophrenic patients and matched normal controls were recruited and assessed for smooth pursuit eye movements, voluntary saccadic eye movements and by means of neuropsychological tests (Toluse-Pieron test and Wisconsin Card Sorting Test). No clear-cut relationship was found between eye movement performance and neuropsychological impairment.

Adult↗

Lorazepam-induced modifications of saccadic and smooth-pursuit eye movements in humans: attentional and motor factors.

In a placebo-controlled, double-blind study, we measured the effects of low dose lorazepam on attentional and motor factors involved in saccadic and smooth pursuit eye movements. We manipulated the temporal interval between the extinction of the central fixation target and the appearance of a second eccentric target (gap/overlap step paradigm). The second target was either stationary (saccade trial) or moving in a direction opposite to the step (pursuit trial). Gap/overlap effects on the latency of saccadic and smooth pursuit eye movements were measured before and after oral intake of either lorazepam or placebo. Pharmacological effects on the dynamics and the accuracy of both types of eye movements were also investigated. In 14 healthy volunteers, we found that the temporal interval between fixation target offset and eccentric target onset modulates the latency of saccadic and smooth pursuit eye movements in a similar way. As compared to placebo, lorazepam significantly increased the latency of both types of eye movements, but did not modify the gap/overlap effect. Moreover, lorazepam significantly decreased the peak velocity of the first saccade towards the eccentric stationary target, as well as the gain of tracking towards the eccentric moving target. However, the overall accuracy of both behaviors was not significantly affected, indicating that systematic errors in foveating or tracking were detected and corrected by appropriate corrective or catch-up saccades, respectively. Results are discussed in terms of shared/different mechanisms for saccadic and pursuit systems in primates.

Adult↗

A quantitative analysis of eye movements during the cover test--a preliminary report.

The aim of this study was to carry out quantitative analyses of eye movements during the cover test on a group (n = 57) of asymptomatic emmetropes. Eye movements were recorded during an automated cover test while subjects fixated a distance (3.4 m) and near (0.4 m) target. There was a significant difference between the amplitude of phoria measured after 2 s of occlusion compared to 10 s (P < 0.01). The mean phoria after 10 s of occlusion was 0.1 degree (eso) for distance fixation and 1.6 degrees (exo) for near fixation. The distribution was approximately normal for both distance and near fixation but the standard deviation was greater for near (2.0 degrees) than distance fixation (1.4 degrees). The pattern of eye movements during the recovery phase was more complex than is often assumed, in many cases consisting of a variety of saccadic and vergence movements involving both the covered and 'fixing' eye. The latency of the first recovery movement was significantly shorter for exophoric than esophoric deviations (P < 0.01) and the latency of some recovery movements were very short (< 150 ms).

Adolescent↗

Inhibitory inefficiency and failures of intention activation: age-related decline in the control of saccadic eye movements.

Young and older adults' control of saccadic eye movements was compared using an antisaccade task, which requires the inhibition of a reflexive saccade toward a peripheral onset cue followed by an intentional saccade in the opposite direction. In 2 experiments, an age-related decline was found in the suppression of reflexive eye movements, as indicated by an increased proportion of saccades toward the cue, and a longer time needed to initiate correct antisaccades. The results from Experiment 2 suggested that older adults' slower antisaccades may be explained partly in terms of increased failures to maintain the cue-action representation at a sufficient activation level. The results suggest that the notion of selective preservation with age of the ability to inhibit spatial responses does not apply to the active inhibition of prepotent spatial responses.

Adolescent↗

Eye movements and processing stages in reading: relative contribution of visual, lexical, and contextual factors.

The independent and the combined influence of word length, word frequency, and contextual predictability on eye movements in reading was examined across processing stages under two priming-context conditions. Length, frequency, and predictability were used as predictors in multiple regression analyses, with parafoveal, early, late, and spillover eye movement measures as the dependent variables. There were specific effects of: (a) length, both on where to look (how likely a word was fixated and in which location) and how long to fixate, across all processing stages; (b) frequency, on how long to fixate a word, but not on where to look, at an early processing stage; and (c) predictability, both on how likely a word was fixated and for how long, in late processing stages. The source of influence for predictability was related to global rather than to local contextual priming. The contribution of word length was independent of contextual source. These results are relevant to determine both the time course of the influence of visual, lexical, and contextual factors on eye movements in reading, and which main component of eye movements, that is, location or duration, is affected.

Eye Movements↗

Object motion computation for the initiation of smooth pursuit eye movements in humans.

Pursuing an object with smooth eye movements requires an accurate estimate of its two-dimensional (2D) trajectory. This 2D motion computation requires that different local motion measurements are extracted and combined to recover the global object-motion direction and speed. Several combination rules have been proposed such as vector averaging (VA), intersection of constraints (IOC), or 2D feature tracking (2DFT). To examine this computation, we investigated the time course of smooth pursuit eye movements driven by simple objects of different shapes. For type II diamond (where the direction of true object motion is dramatically different from the vector average of the 1-dimensional edge motions, i.e., VA not equal IOC = 2DFT), the ocular tracking is initiated in the vector average direction. Over a period of less than 300 ms, the eye-tracking direction converges on the true object motion. The reduction of the tracking error starts before the closing of the oculomotor loop. For type I diamonds (where the direction of true object motion is identical to the vector average direction, i.e., VA = IOC = 2DFT), there is no such bias. We quantified this effect by calculating the direction error between responses to types I and II and measuring its maximum value and time constant. At low contrast and high speeds, the initial bias in tracking direction is larger and takes longer to converge onto the actual object-motion direction. This effect is attenuated with the introduction of more 2D information to the extent that it was totally obliterated with a texture-filled type II diamond. These results suggest a flexible 2D computation for motion integration, which combines all available one-dimensional (edge) and 2D (feature) motion information to refine the estimate of object-motion direction over time.

Analysis of Variance↗

Deficits in saccadic eye-movement control in Parkinson's disease.

In contrast to their slowed limb movements, individuals with Parkinson's disease (PD) produce rapid automatic eye movements to sensory stimuli and show an impaired ability to generate voluntary eye movements in cognitive tasks. Eighteen PD patients and 18 matched control volunteers were instructed to look either toward (pro-saccade) or away from (anti-saccade) a peripheral stimulus as soon as it appeared (immediate, gap and overlap conditions) or after a variable delay; or, they made sequential saccades to remembered targets after a variable delay. We found that PD patients made more express saccades (correct saccades in the latency range of 90-140 ms) in the immediate pro-saccade task, more direction errors (automatic pro-saccades) in the immediate anti-saccade task, and were less able to inhibit saccades during the delay period in all delay tasks. PD patients also made more directional and end-point errors in the memory-guided sequential task. Their inability to plan eye movements to remembered target locations suggests that PD patients have a deficit in spatial working memory which, along with their deficit in automatic saccade suppression, is consistent with a disorder of the prefrontal-basal ganglia circuit. Impairment of this pathway may release the automatic saccade system from top-down inhibition and produce deficits in volitional saccade control. Parallel findings across various motor, cognitive and oculomotor tasks suggest a common mechanism underlying a general deficit in automatic response suppression.

Adult↗

Lossy compression of eye movement and auditory brainstem response signals.

Eye movement and auditory brainstem response signals recorded for balance and hearing investigations were used as a medical test battery for several types of lossy compression techniques. These signals are associated with the function of the ears. The former signals are used to assess the balance problems (especially vertigo) of a subject and the latter his or her hearing problems. New technique is also presented based on successive approximation quantization. The effect of information loss on medical parameters computed from the signals in the course of compression was evaluated for brainstem response signals. It is important to ensure that lossy compression techniques of these biomedical signals do not impair medical parameter values computed from the signals.

Algorithms↗

Effect of grammatical gender on visual word recognition: evidence from lexical decision and eye movement experiments.

Lexical decision times and eye movements were recorded to determine whether grammatical gender can influence the visual recognition of isolated French nouns. This issue was investigated by assessing the use of two types of regularities between a noun's form and its gender--namely ending-to-gender regularities (e.g., the final letter sequence -at appears only in masculine nouns and, thus, is predictive of masculine gender) and gender-to-ending regularities (e.g., feminine gender would predict the final letter e, whereas masculine gender would not). Previous studies have shown that noun endings are used by readers when they have to identify gender. However, the influence of ending-to-gender predictiveness has never been investigated in a lexical decision task, and the effect of gender-to-ending regularities has never been evaluated at all. The results suggest that gender information can influence both the activation stage (Experiments 1 and 3) and the selection stage (Experiments 2 and 3) of the word recognition process.

Adult↗