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Visual scene memory and the guidance of saccadic eye movements.

An unresolved question is how much information can be remembered from visual scenes when they are inspected by saccadic eye movements. Subjects used saccadic eye movements to scan a computer-generated scene, and afterwards, recalled as many objects as they could. Scene memory was quite good: it improved with display duration, it persisted over time long after the display was removed, and it continued to accumulate with additional viewings of the same display (Melcher, D. (2001) The persistance of memory for scenes. Nature 412, 401). The occurrence of saccadic eye movements was important to ensure good recall performance, even though subjects often recalled non-fixated objects. Inter-saccadic intervals increased with display duration, showing an influence of duration on global scanning strategy. The choice of saccadic target was predicted by a Random Selection with Distance Weighting (RSDW) model, in which the target for each saccade is selected at random from all available objects, weighted according to distance from fixation, regardless of which objects had previously been fixated. The results show that the visual memory that was reflected in the recall reports was not utilized for the immediate decision about where to look in the scene. Visual memory can be excellent, but it is not always reflected in oculomotor measures, perhaps because the cost of rapid on-line memory retrieval is too great.

Attention↗

A quantitative study of auditory-evoked saccadic eye movements in two dimensions.

We investigated the properties of human saccadic eye movements evoked by acoustic stimuli in the two-dimensional frontal plane. These movements proved to be quite accurate, both in azimuth and in elevation, provided the sound source spectrum had a broad bandwidth and a sufficiently long duration. If the acoustic target was a tone, the azimuth of the saccadic end points remained equally accurate, whereas the elevation of the response was related to the frequency of the tone, rather than to the physical position of the target. Saccade elevation accuracy also declined substantially for short-duration noise bursts, although response elevation remained highly correlated with target elevation. The latencies of auditory saccades depended on the amplitude, but not on the direction of the eye movement, suggesting a polar coordinate origin of auditory saccade initiation. We also observed that the trajectories of auditory saccades were often substantially curved. Both a qualitative and a model-based analysis showed that this curvature corrected for errors in the initial direction of the saccade. The latter analysis also suggested that the kinematic properties of auditory saccades could be described by the superposition of two overlapping saccadic eye movements, hypothesized to be based on binaural difference cues and monaural spectral cues in the auditory signal, respectively. It is argued that, although the audio-oculomotor system has to operate in a feedforward way, it must nevertheless have access to an accurate representation of actual and desired eye position. Different models underlying the generation of auditory saccades are discussed.

Adult↗

Slow correcting eye movements of head-fixed, trained cats toward stationary targets.

Inspection of eye saccades made by head-fixed, trained cats revealed the existence of many eye shifts at an approximately constant velocity during the deceleratory phase of the saccade or at the end of it. Slow eye movements occurring at the end of a saccade are usually referred to as "postsaccadic drifts". It is shown that the duration and mean velocity of these "drifts" are related to the amplitude of the movement. The kinematics of these slow eye movements are nevertheless different from those of saccades. Slow movements at the end of the gaze shift have longer durations than those occurring during the intersaccadic interval between a saccade and a reacceleration of the eye. A closer study of the drifts of three trained cats showed that they play an important corrective role in reducing the residual error at the end of a saccade or during an intersaccadic interval. This functional corrective role was demonstrated by relating the amplitude of the slow movement to the amplitude of the residual error when the slow velocity eye shift began. It is therefore proposed to name these eye shifts "slow correcting movements".

Animals↗

Evaluation of eye movement variables of Chinese and American readers. Information abstraction by Chinese and American readers.

Eye movements were recorded while competent readers of English and Chinese read in their respective languages. There were no important differences in oculometric patterns of fixation pause durations for Chinese and American readers, suggesting that cognitive aspects of reading are more important than perceptual ones in determining eye movement patterns and fixation pauses of competent readers. An analysis of undershoot saccades associated with return of the eyes from the end of one line to the beginning of the next line suggests that these occur more frequently during reading than during the performance of similar eye movements that are more perceptually determined, however, the amount of undershoot does not discriminate between readers of alphabetic and logographic texts.

China↗

The induction and compensation of asymmetric eye movements following unilateral blockage of a horizontal semicircular canal in the rabbit.

The influence of unilateral plugs of the left horizontal semicircular canal (LHC plugs) of rabbits on the development and compensation of asymmetric eye movements evoked by horizontal vestibular stimulation was studied. LHC plugs caused an immediate reduction of 50-65% in the gain of the horizontal vestibuloocular reflex (HVOR). This reduction in gain was achieved without altering the symmetry of the HVOR, and was accompanied by a change in the axial alignment of eye movements evoked by vestibular stimulation about the vertical (HVOR) and longitudinal (VVOR) axes. Postoperative asymmetry of eye movements developed 12-48 hr after the plugging operation. The development of asymmetry was reduced if the rabbit was restrained for 24 hr, thereby minimizing vestibular stimulation following the plugging operation. Over a 3-4 week period, the normal symmetry of eye movements was restored and the axial alignments of the HVOR and VVOR returned to the preoperative values. The gain of the HVOR did not recover. The horizontal cervicoocular reflex (HCOR) was examined before the plugging operation and after compensation of asymmetry was complete. The gain and phase of the HCOR were not altered. A relatively simple set of explanations at a cellular level is proposed to account for the induction and compensation of asymmetric eye movements following a unilateral plug of the horizontal semicircular canal.

Adaptation, Physiological↗

Principles underlying real-time nystagmus analysis of horizontal and vertical eye movements recorded with electro-, infra-red-, or video-oculographic techniques.

New methods for separating fast and slow phases of human nystagmus in real time are presented that are adaptive to the time-dependent noise properties of the input eye movement signal and therefore applicable to different recording techniques and directions. The methods employ a statistical filter technique to track slow-phase eye movements, uninfluenced by fast phases and blink artifacts, and fuzzy-logic techniques to identify fast-phase eye movements. Because these two techniques are decoupled from one another, highly accurate phase separation and slow-phase velocity profiles are achieved. In addition, the tracking of the variance of slow-phase and fast-phase eye movement recording permits a quality control of the analysis for different recording techniques and a variety of ocular nystagmus responses. Because blinks impose different eye velocity profiles on the recordings, depending on the type of recording technique and direction (horizontal, vertical), blink detection and its effect on fast-phase amplitude must be individually adjusted to each recording technique. Results are illustrated in the context of simultaneously recorded video-oculographic, infra-red, and electro-oculographic recordings of vestibulo-ocular reflex and optokinetic reflex responses causing horizontal or vertical eye movements.

Algorithms↗

Higher control mechanisms of saccadic eye movements.

The higher control of saccadic eye movements depends on a complex interaction between a number of different neural centres. Inputs from the frontal eye field and superior colliculus both project to the brainstem and appear to provide parallel and interacting inputs. The former is more concerned with voluntary shifts of gaze, whereas the latter mediates involuntary reflexive gaze shifts to suddenly appearing visual stimuli. The basal ganglia through the substantia nigra, pars reticularis has direct inputs to the colliculus and appears to have a gating role. These concepts, derived from experimental studies on primates, are discussed in relation to clinical material.

Animals↗

Effects of digital filtering on the parameters of sinusoidal tracking eye movements.

In the present study linear nonrecursive (FIR) filters designed with the Hamming window and hybrid median filters were examined in connection with computer analysis of sinusoidal tracking eye movements. It is important that digital filters which do not distort sensitive properties of eye movement signals are employed in order to prevent changes caused by too strict filtering in average parameter values of eye movement signals, indicators of some otoneurological disorders. It turned out that within reasonable limits these filters are valid in filtering of sinusoidal tracking eye movement signals.

Automation↗

Neurons in monkey parietal area LIP are tuned for eye-movement parameters in three-dimensional space.

1. A functional class of neurons in area LIP on the lateral bank of the intraparietal sulcus were shown previously (Gnadt and Andersen 1988) to be related to the metrics of saccadic eye movements. In this study, we tested LIP neurons at different depths with respect to the plane of fixation. 2. Sixty-one neurons were identified for their increased activity before saccadic eye movements. While holding the location of the target constant at the center of the frontoparallel (saccadic) response field, the neurons were tested systematically during eye movements to target positions proximal (near) to the plane of fixation, at the plane of fixation, and distal (far) to the plane of fixation. By necessity, the movements of these targets required a combination of saccadic and vergence movements. 3. Seventy-two percent of the neurons were found to change their activity as a function of target depth relative to the plane of fixation. The neurons had broad tuning curves for depth. Some cells preferred "near" target positions, some preferred "far" positions, and others responded best in the frontoparallel plane of fixation. 4. The location of a neuron's response field in the frontoparallel plane remained constant regardless of target depth. However, the magnitude of the neuron's response increased when the target was positioned at the preferred depth and it decreased for targets positioned at nonpreferred depths. This indicated that the neurons always were related to the same frontoparallel coordinates, but responded more vigorously when the target was positioned at its preferred depth. 5. The visual display apparatus allowed independent presentation of two stimulus cues for depth: binocular disparity and accommodative demand whereas other cues were held constant. For many neurons, either cue was sufficient to tune the activity in depth, though most neurons responded best for the geometrically appropriate combination of the two cues. 6. Comparison of the binocular tuning for depth with the individual monocular responses showed that the tuning for depth was not produced by simple linear combination of two monocular response fields. 7. We tested a subset of the neurons in a double-movement task that dissociated the retinal coordinates of the visual stimuli from the eye-movement coordinates of the second movement. These tests confirmed earlier findings that this functional class of neurons are active when the eye-movement coordinates matched the neurons' response field. It was not necessary for a visual stimulus to fall within the neurons' response field for them to become active.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Exploratory eye movements in schizophrenia: effects of figure size and the instruction on visual search.

It has been reported that patients with schizophrenia show restricted eye-scanning in comparison with normal controls; however, the precise mechanism underlying the limited eye movement pattern remains unknown. The purpose of this study was to determine the factors affecting restricted eye-scanning in schizophrenic patients by examining exploratory eye movements during demonstration of two different sizes of the S-shaped figure. The second purpose was to determine the effect of the instruction for performance on the restricted viewing pattern in patients with schizophrenia. Eye movements during demonstration of the S-shaped figure of the original or half size were examined in 15 patients with schizophrenia and 15 normal controls using an infrared eye-mark recorder. The patients showed lower search scores than control subjects for both sizes of the figure. The subjects were then instructed to compare a slightly modified figure with the original one. Lower responsive search scores were found for the patients when "fixation point" was defined as a point at which a gaze was held for at least 200 ms, while the patients and control subjects performed equally at the 100-ms setting. Direct instruction to scrutinize the S-shape abolished the difference in the search scores between patients and control subjects at both the 100-ms and 200-ms settings. These findings suggest that the size of the S-figure is not a factor of restricted eye movements, and that the direct instruction improves the visual performance in patients with schizophrenia.

Adult↗

Head and eye movements following vestibular stimulus in squirrel monkeys.

Head and eye movements induced by vestibular stimuli (rotational velocity steps) were studied under several experimental conditions in squirrel monkeys. Monkeys showed prominent head nystagmus under a free head condition. Slow phase gaze velocity (SPGV) was calculated by summing the slow phase speed of coupled head and eye nystagmus. A statistical test showed no significant difference between slow phase eye velocities in a fixed head versus SPGV in a free head condition. This indicates that an accurate negative feedback control of eye movements is performed by the vestibular system when sensing active head movements. Bilateral lateral semicircular canal block eliminated these responses in subjects without amphetamine injection. However, in amphetamine-injected animals, 14-21% of preoperative response could be found even after this operation. Since the plane of rotation was not parallel to that of the lateral semicircular canals, the result was considered to be due to vertical semicircular canal stimulation.

Animals↗

Effects of alcohol and other psychotropic drugs on eye movements: relevance to traffic safety.

The effects of alcohol and other psychotropic drugs on eye movements are reviewed with particular attention to the possible relevance of these effects for traffic safety. Alcohol has been shown to have diverse effects, including reduction of the velocity of both saccadic and smooth pursuit eye movements, increased saccadic latency, impairment of convergence and induction of nystagmus. These effects probably contribute to impaired visual information processing, which reduces driving ability. Barbiturates have been reported to produce effects similar to alcohol, and the effects of benzodiazepines and opioids seem to be more limited but still substantial. Marihuana has relatively little effect on eye movements.

Accidents, Traffic↗

The effects of anterior arcuate and dorsomedial frontal cortex lesions on visually guided eye movements in the rhesus monkey: 1. Single and sequential targets.

This study examined the effects of anterior arcuate and dorsomedial frontal cortex lesions on visually guided eye movements in three rhesus monkeys. Lesions of the anterior bank of the arcuate, where the frontal eye fields reside, produced major deficits in the execution of saccadic eye movements to sequentially presented targets that did not recover even after 1 year after the lesions. Ablations of the dorsomedial frontal cortex, wherein the medial eye fields reside, produced much smaller and shorter-duration deficits on this task. Deficits after paired lesions of the anterior arcuate and dorsomedial frontal cortex were of approximately the same magnitude as after single anterior arcuate lesions. Anterior arcuate lesions also increased saccadic reaction times to single visual targets and decreased saccadic velocities that recovered gradually over a period of 2-5 months. Dorsomedial frontal cortex lesions produced only small increases in saccadic latencies that recovered rapidly. None of the lesions produced deficits in executing combined saccadic and pursuit eye movements to moving targets. The results suggest that the anterior arcuate area plays a central role in the execution of sequences of eye movements to successively appearing targets.

Animals↗

Improvement in smooth pursuit eye movements after cigarette smoking in schizophrenic patients.

This study examined whether schizophrenics' cigarette smoking normalized smooth pursuit eye movement abnormalities. Fifteen schizophrenic and 15 nonschizophrenic subjects abstained from their usual cigarette smoking for an average of 10 h. Their baseline performance during a constant velocity smooth pursuit task was then assessed. The subjects smoked as much as they desired in a 10-min period and then were retested immediately postsmoking, and 10 and 20 min later. Smooth pursuit gain and the percentage of total eye movement due to various saccadic subtypes were computed using infrared oculography and computerized pattern recognition software. After smoking, smooth pursuit gain increased and the percentage of total eye movements due to leading saccades decreased significantly in the schizophrenic patients. There were no changes in the gain or leading saccades of nonschizophrenic subjects after smoking. Nicotinic receptor dysfunction may be a candidate mechanism for smooth pursuit eye movement abnormalities in schizophrenia.

Adult↗

Topography of saccadic eye movements evoked by microstimulation in rabbit cerebellar vermis.

1. We investigated saccadic eye movements elicited by microstimulation in the vermis of the rabbit. Scleral search coils were implanted under the conjunctiva of both eyes and a recording chamber was placed over the cerebellar vermis. 2. Conjugate saccadic eye movements were evoked in lobules VIa, b and c and VII of the vermis by currents ranging from 4 to 60 microA. All movements were horizontal with no apparent vertical component. 3. The cortex on both sides of the vermal mid-line could be divided in two zones, dependent on the direction of elicited saccades. In the medial zone saccades were directed ipsilaterally, in the lateral zone contralaterally. 4. We conclude that the topography of saccadic eye movements in the rabbit cerebellar vermis is, unlike in monkey and cat, organized in parasagittal zones.

Animals↗

Rapid eye movement sleep deprivation in post-critical period, adolescent rats alters the balance between inhibitory and excitatory mechanisms in visual cortex.

Suppression of rapid eye movement sleep (REMS) in developing animals has both anatomical and physiological consequences. We have recently shown that initiating REMS deprivation (REMSD) prior to the end of the critical period in young rats delays termination of the critical period (CP) in visual cortex, and, consequently, the synaptic plasticity mechanisms that support a developmentally regulated form of long-term potentiation (LTP) are maintained in an immature state [J.P. Shaffery, C.M. Sinton, G. Bisset, H.P. Roffwarg, G.A. Marks, Rapid eye movement sleep deprivation modifies expression of long-term potentiation in visual cortex of immature rats, Neuroscience, 110 (2002) 431-443]. In CP animals, high-frequency, theta burst stimulation (TBS) directed at the white matter (WM) below visual cortex produces LTP in the post-synaptic cells in layer II/III (LTPWM-III). However, LTPWM-III can be induced in cortical tissue taken from REMS-deprived animals for up to a week beyond the usual end of the CP [J.P. Shaffery, C.M. Sinton, G. Bisset, H.P. Roffwarg, G.A. Marks, Rapid eye movement sleep deprivation modifies expression of long-term potentiation in visual cortex of immature rats, Neuroscience, 110 (2002) 431-443]. Further, in post-CP, adolescent animals (as late as postnatal day 60), REMSD appears to unmask synaptic plasticity mechanisms that allow for production of developmentally regulated LTPWM-III [J.P. Shaffery, J. Lopez, G. Bissette, H.P. Roffwarg, Rapid eye movement sleep deprivation revives a form of developmentally regulated synaptic plasticity in the visual cortex of post-critical period rats, Neurosci Lett., (2005), in press]. It has been proposed that REMSD's effects on production of LTPWM-III result from a reduction in efficiency of the inhibitory mechanisms thought to precipitate termination of the CP of brain development [J.P. Shaffery, J. Lopez, G. Bissette, H.P. Roffwarg, Rapid eye movement sleep deprivation revives a form of developmentally regulated synaptic plasticity in the visual cortex of post-critical period rats, Neurosci Lett., (2005), in press]. In this study we tested the hypothesis that low-frequency stimulation (LFS) of the fibers of the WM, which usually produces the related form of synaptic plasticity, long-term depression (LTD), will also reflect the reduction in inhibitory tone. We report here that LFS protocols, which in normally sleeping, adolescent rats usually produce either LTD or no change in response magnitude, in REMS-deprived, adolescent rats are more likely to produce LTP.

Age Factors↗

Reading without saccadic eye movements.

To assess the limitation on reading speed imposed by saccadic eye movements, we measured reading speed in 13 normally-sighted observers using two modes of text presentations: PAGE text which presents an entire passage conventionally in static, paragraph format, and rapid serial visual presentation (RSVP) which presents text sequentially, one word at a time at the same location in the visual field. In Expt 1, subjects read PAGE and RSVP text orally across a wide range of letter sizes (2X to 32X single-letter acuity) and reading speed was computed from the number of correct words read per minute. Reading speeds were consistently faster for RSVP compared to PAGE text at all letter sizes tested. The average speeds for text of an intermediate letter size (8X acuity) were 1171 words/min for RSVP and 303 words/min for PAGE text. In Expt 2 subjects read PAGE and RSVP text silently and a multiple-choice comprehension test was administered after each passage. All subjects continued to read RSVP text faster, and 6 subjects read at the maximum testable rate (1652 words/min) with at least 75% correct on the comprehension tests. Experiment 3 assessed the minimum word exposure time required for decoding text using RSVP to minimize potential delays due to saccadic eye movement control. Successive words were presented for a fixed duration (word duration) with a blank interval (ISI) between words. The minimum word duration required for accurate oral reading averaged 69.4 msec and was not reduced by increasing ISI. We interpret these results as an indication that the programming and execution of saccadic eye movements impose an upper limit on conventional reading speed.

Adult↗

Eye tracking dysfunction in schizophrenia: characterization of component eye movement abnormalities, diagnostic specificity, and the role of attention.

To characterize oculomotor components and diagnostic specificity of eye tracking abnormalities in schizophrenia, we examined a large consecutively admitted series of psychotic patients and matched controls. The most common abnormality in schizophrenic patients was low gain (slow) pursuit eye movements (47% of cases). Pursuit and saccadic eye movement abnormalities were no more severe in schizophrenic Ss than in those with affective psychoses, except that high rates of catch-up saccades were unique to schizophrenic Ss (17% of cases). These findings indicate that impaired pursuit eye movements are a major cause of eye tracking impairments in schizophrenia, that tracking dysfunctions commonly occur in affective psychoses, and that markedly high rates of catch-up saccades during eye tracking may be specific to schizophrenia.

Adult↗