Small Saccades Research Articles

Kinematically distinct saccades are used in a context-dependent manner by larval zebrafish

Saccades are rapid eye movements that are used by all species with good vision. In this study, we set out to characterize the complete repertoire of larval zebrafish horizontal saccades to gain insight into their contributions to visually guided behavior and underlying neural control. We identified five saccade types, defined by systematic differences in kinematics and binocular coordination, which were differentially expressed across a variety of behavioral contexts. Conjugate saccades formed a large group that serves at least four functions. These include fast phases of the optokinetic nystagmus, visual scanning in stationary animals, and shifting gaze in coordination with body turns. In addition, we discovered a previously undescribed pattern of eye-body coordination in which small conjugate saccades partially oppose head rotation to maintain gaze during forward locomotion. Convergent saccades were coordinated with body movements to foveate prey targets during hunting. Detailed kinematic analysis showed that conjugate and convergent saccades differed in the millisecond coordination of the eyes and body and followed distinct velocity main sequence relationships. This challenges the prevailing view that all horizontal saccades are controlled by a common brainstem circuit and instead indicates saccade-type-specific neural control.

(Micro)saccade-related potentials during face recognition: A study combining EEG, eye-tracking, and deconvolution modeling.

Under natural viewing conditions, complex stimuli such as human faces are typically looked at several times in succession, implying that their recognition may unfold across multiple eye fixations. Although electrophysiological (EEG) experiments on face recognition typically prohibit eye movements, participants still execute frequent (micro)saccades on the face, each of which generates its own visuocortical response. This finding raises the question of whether the fixation-related potentials (FRPs) evoked by these tiny gaze shifts also contain psychologically valuable information about face processing. Here, we investigated this question by corecording EEG and eye movements in an experiment with emotional faces (happy, angry, neutral). Deconvolution modeling was used to separate the stimulus ERPs to face onset from the FRPs generated by subsequent microsaccades-induced refixations on the face. As expected, stimulus ERPs exhibited typical emotion effects, with a larger early posterior negativity (EPN) for happy/angry compared with neutral faces. Eye tracking confirmed that participants made small saccades in 98% of the trials, which were often aimed at the left eye of the stimulus face. However, while each saccade produced a strong response over visual areas, this response was unaffected by the face's emotional expression, both for the first and for subsequent (micro)saccades. This finding suggests that the face's affective content is rapidly evaluated after stimulus onset, leading to only a short-lived sensory enhancement by arousing stimuli that does not repeat itself during immediate refixations. Methodologically, our work demonstrates how eye tracking and deconvolution modeling can be used to extract several brain responses from each EEG trial, providing insights into neural processing at different latencies after stimulus onset.

Poster Session: Spatial-frequency-selective enhancement of visual sensitivity from saccade dynamics.

Eye movements transform a spatial scene into luminance modulations on the retina. Recent work has shown that this transformation is highly structured: within human temporal sensitivity, saccades deliver power that increases in proportion to spatial frequency (SF) up to a critical frequency and remains constant beyond that. Importantly, the critical SF increases with decreasing amplitude. Therefore, at sufficiently low SFs, larger saccades effectively deliver stronger input signals to the retina. Here we tested whether this input reformatting has the predicted perceptual consequences, by examining how large and small saccades (6o & 1o) affect contrast sensitivity. We measured relative sensitivity at two SFs: a reference (0.5 cpd), equal to the critical SF for the small saccade, and a probe at either a lower or higher SF (0.1/2.5 cpd). We predicted that large saccades enhance visibility only when the probe has a lower SF than the reference. Subjects (N=7) made instructed saccades while presented with a plaid of overlapping orthogonal gratings at the two SFs and reported which grating was more visible. Results closely follow theoretical predictions: psychometric functions following small and large saccades only differed with the lower SF probe, in which case the larger saccade significantly enhanced visibility. In sum, saccades enable selectivity not only in the spatial domain, but also in the spatial-frequency domain.

Patterns of small involuntary fixation saccades (SIFSs) in different neurodegenerative diseases: the role of noise

During the attempt to steadily fixate at a single spot, sequences of small involuntary fixation saccades (SIFSs, known also as microsaccades οr intrusions) occur which form spatio-temporal patterns such as square wave jerks (SWJs), a pattern characterised by alternating centrifugal and centripetal movements of similar magnitude. In many neurodegenerative disorders, SIFSs exhibit elevated amplitudes and frequencies. Elevated SIFS amplitudes have been shown to favour the occurrence of SWJs (“SWJ coupling”). We analysed SIFSs in different subject groups comprising both healthy controls (CTR) and patients with amyotrophic lateral sclerosis (ALS) and progressive supranuclear palsy (PSP), i.e. two neurodegenerative diseases with completely different neuropathological basis and different clinical phenotypes. We show that, across these groups, the relations between SIFS amplitude and the relative frequency of SWJ-like patterns and other SIFS characteristics follow a common law. As an explanation, we propose that physiological and technical noise comprises a small, amplitude-independent component that has little effect on large SIFSs, but causes considerable deviations from the intended amplitude and direction of small ones. Therefore, in contrast to large SIFSs, successive small SIFSs have a lower chance to meet the SWJ similarity criteria. In principle, every measurement of SIFSs is affected by an amplitude-independent noise background. Therefore, the dependence of SWJ coupling on SIFS amplitude will probably be encountered in almost any group of subjects. In addition, we find a positive correlation between SIFS amplitude and frequency in ALS, but none in PSP, suggesting that the elevated amplitudes might arise at different sites in the two disorders.

The influence of scene tilt on saccade directions is amplitude dependent

When exploring a visual scene, humans make more saccades in the horizontal direction than any other direction. While many have shown that the horizontal saccade bias rotates in response to scene tilt, it is unclear whether this effect depends on saccade amplitude. We addressed this question by examining the effect of image tilt on the saccade direction distributions recorded during freely viewing natural scenes. Participants (n = 20) viewed scenes tilted at −30°, 0°, and 30°. Saccade distributions during free viewing rotated by an angle of 12.1° ± 6.7° (t(19) = 8.04, p < 0.001) in the direction of the image tilt. When we partitioned the saccades according to their amplitude we found that small amplitude saccades occurred most in the horizontal direction while large amplitude saccades were more oriented to the scene tilt (p < 0.001). To further study the characteristics of small saccades and how they are affected by scene tilt, we looked at the effect of image tilt on small fixational saccades made while fixating a central target amidst a larger scene and found that fixational saccade distributions did not rotate with scene tilt (−0.3° ±1.7° degrees; t(19) = −0.8, p = 0.39). These results suggest a combined effect of two reference frames in saccade generation: one egocentric reference frame that dominates for small saccades, biases them horizontally, and may be common for different tasks, and another allocentric reference frame that biases larger saccades along the orientation of an image during free viewing.

Enhanced temporal resolution of vision in action video game players

Video game play has been suggested to improve visual and attention processing. Nevertheless, while action video game play is highly dynamic, there is scarce research on how information is temporally discriminated at the millisecond level. This cross-sectional study investigates whether temporal discrimination at the millisecond level in vision varies across action video game players (VGPs; N = 23) and non-video game players (NVGPs; N = 23). Participants discriminated synchronous from asynchronous onsets of two visual targets in virtual reality, while their EEG and oculomotor movements were recorded. Results show an increased sensitivity to short asynchronies (11, 33 and 66 ms) in VGPs compared with NVGPs, which was especially marked at the start of the task, suggesting better temporal discrimination abilities. Pre-targets oculomotor freezing – the inhibition of small fixational saccades – was associated with correct temporal discrimination, probably revealing attentional preparation. However, this parameter did not differ between groups. EEG and reconstruction analyses suggest that the enhancement of temporal discrimination in VGPs during temporal discrimination is related to parieto-occipital processing, and a reduction of alpha-band (8–14 Hz) power and inter-trial phase coherence. Overall, the study reveals an enhanced ability in action video game players to discriminate in time visual events in close temporal proximity combined with reduced alpha-band oscillatory activities. Consequently, playing action video games is associated with an improved temporal resolution of vision.

Muscles that move the retina augment compound eye vision in Drosophila.

Most animals have compound eyes, with tens to thousands of lenses attached rigidly to the exoskeleton. A natural assumption is that all of these species must resort to moving either their head ortheir body to actively change their visual input. However, classic anatomy has revealed that flies have muscles poised to move their retinas under the stable lenses of each compound eye1-3. Here we show that Drosophila use their retinal muscles to smoothly track visual motion, which helps to stabilize the retinal image, and also to perform small saccades when viewing a stationary scene. We show that when the retina moves, visual receptive fields shift accordingly, and that even the smallest retinal saccades activate visual neurons. Using a head-fixed behavioural paradigm, we find that Drosophila perform binocular, vergence movements of their retinas-which could enhance depth perception-when crossing gaps, and impairing the physiology of retinal motor neurons alters gap-crossing trajectories during free behaviour. That flies evolved an ability to actuate their retinas suggests that moving the eye independently of the head is broadly paramount for animals. The similarities of smooth and saccadic movements of the Drosophila retina and the vertebrate eye highlight a notable example of convergent evolution.

Analysis of video head impulse test saccades data in patients with vestibular migraine or probable vestibular migraine

ObjectiveSaccades accompanied by normal gain in video head impulse tests (vHIT) are often observed in patients with vestibular migraine (VM). However, they are not considered as an independent indicator, reducing their utility in diagnosing VM. To better understand clinical features of VM, it is necessary to understand raw saccades data. MethodsFourteen patients with confirmed VM, 45 patients with probable VM (p-VM) and 14 age-matched healthy volunteers were included in this study. Clinical findings related to spontaneous nystagmus (SN), positional nystagmus (PN), head-shaking nystagmus (HSN), caloric test and vHIT were recorded. Raw saccades data were exported and numbered by their sequences, and their features analyzed. ResultsVM patients showed no SN, PN or HSN, and less than half of them showed unilateral weakness (UW) on caloric test. The first saccades from lateral semicircular canal stimulation were the most predominant for both left and right sides. Neither velocity nor time parameters were significantly different when compared between the two sides. Most VM patients (86%) exhibited small saccades, around 35% of the head peak velocity, with a latency of 200–400 ms. Characteristics of saccades were similar in patients with p-VM. Only four normal subjects showed saccades, all unilateral and seemingly random. ConclusionsSmall saccades involving bilateral semicircular canals with a scattered distribution pattern are common in patients with VM and p-VM.

The amplitude of small eye movements can be accurately estimated with video-based eye trackers

Estimating the gaze direction with a digital video-based pupil and corneal reflection (P-CR) eye tracker is challenging partly since a video camera is limited in terms of spatial and temporal resolution, and because the captured eye images contain noise. Through computer simulation, we evaluated the localization accuracy of pupil-, and CR centers in the eye image for small eye rotations (≪ 1 deg). Results highlight how inaccuracies in center localization are related to 1) how many pixels the pupil and CR span in the eye camera image, 2) the method to compute the center of the pupil and CRs, and 3) the level of image noise. Our results provide a possible explanation to why the amplitude of small saccades may not be accurately estimated by many currently used video-based eye trackers. We conclude that eye movements with arbitrarily small amplitudes can be accurately estimated using the P-CR eye-tracking principle given that the level of image noise is low and the pupil and CR span enough pixels in the eye camera, or if localization of the CR is based on the intensity values in the eye image instead of a binary representation.

Properties of rapid eye movements.

This chapter describes the metrics and kinematics of saccades and quick phases of nystagmus, including microsaccades and large eye-head saccades. Small saccades often display dynamic overshoot, predominantly in the abducting eye. Although the function of these overshoots is unclear, the return movement is saccadic in nature. The saccade kinematics can be quantified by stereotyped relations between amplitude, duration, peak eye velocity, and peak acceleration, which vary somewhat with the initial eye position and saccade direction (centripetal vs centrifugal), possibly due to ocular plant characteristics. Saccades in a structured light environment are considerably faster than when executed in total darkness, although the origin for this facilitation is not known. The horizontal and vertical components of slant saccades are coupled, approximately matching their durations, for which possible underlying neural mechanisms are discussed. The chapter closes with a cross-species comparison of saccade characteristics.

The eyes reflect an internal cognitive state hidden in the population activity of cortical neurons.

Decades of research have shown that global brain states such as arousal can be indexed by measuring the properties of the eyes. The spiking responses of neurons throughout the brain have been associated with the pupil, small fixational saccades, and vigor in eye movements, but it has been difficult to isolate how internal states affect the eyes, and vice versa. While recording from populations of neurons in the visual and prefrontal cortex (PFC), we recently identified a latent dimension of neural activity called "slow drift," which appears to reflect a shift in a global brain state. Here, we asked if slow drift is correlated with the action of the eyes in distinct behavioral tasks. We recorded from visual cortex (V4) while monkeys performed a change detection task, and PFC, while they performed a memory-guided saccade task. In both tasks, slow drift was associated with the size of the pupil and the microsaccade rate, two external indicators of the internal state of the animal. These results show that metrics related to the action of the eyes are associated with a dominant and task-independent mode of neural activity that can be accessed in the population activity of neurons across the cortex.

Reading Specific Small Saccades Predict Individual Phonemic Awareness and Reading Speed.

Small fixational eye-movements are a fundamental aspect of vision and thought to reflect fine shifts in covert attention during active viewing. While the perceptual benefits of these small eye movements have been demonstrated during a wide range of experimental tasks including during free viewing, their function during reading remains surprisingly unclear. Previous research demonstrated that readers with increased microsaccade rates displayed longer reading speeds. To what extent increased fixational eye movements are, however, specific to reading and might be indicative of reading skill deficits remains, however, unknown. To address this topic, we compared the eye movement scan paths of 13 neurotypical individuals and 13 subjects diagnosed with developmental dyslexia during short story reading and free viewing of natural scenes. We found that during reading only, dyslexics tended to display small eye movements more frequently compared to neurotypicals, though this effect was not significant at the population level, as it could also occur in slow readers not diagnosed as dyslexics. In line with previous research, neurotypical readers had twice as many regressive compared to progressive microsaccades, which did not occur during free viewing. In contrast, dyslexics showed similar amounts of regressive and progressive small fixational eye movements during both reading and free viewing. We also found that participants with smaller fixational saccades from both neurotypical and dyslexic samples displayed reduced reading speeds and lower scores during independent tests of reading skill. Slower readers also displayed greater variability in the landing points and temporal occurrence of their fixational saccades. Both the rate and spatio-temporal variability of fixational saccades were associated with lower phonemic awareness scores. As none of the observed differences between dyslexics and neurotypical readers occurred during control experiments with free viewing, the reported effects appear to be directly related to reading. In summary, our results highlight the predictive value of small saccades for reading skill, but not necessarily for developmental dyslexia.

Visual Strategies Underpinning the Spatiotemporal Demands During Visuomotor Tasks in Predicting Ball Direction.

We investigated gaze behavior of expert goalkeepers during the prediction of penalty kicks in different spatiotemporal constraints: penalties taken from 11 and 6m. From 11m, goalkeepers were more successful in predicting ball direction, with longer movement time initiation and a visual strategy with more fixations and greater saccade rates than penalties from 6m, where they exhibited fewer fixations with higher microsaccade rates. As long as the opponent's distance is large and time pressure low, gaze can be frequently shifted between the kicker's body and the ball, due to the low cost of saccades. Conversely, when the objects are close, there is increased reliance on foveal and parafoveal information. In conclusion, when the spatiotemporal constraint is less severe, goalkeepers adopt a visual strategy with more fixations and small saccades. When the spatiotemporal constraint is more severe, they rely on peripheral vision to monitor kickers' movements through the use of microsaccades.

The role of fixation disengagement and oculomotor preparation in gap saccade task is gap-duration dependent.

The influence of internal brain state on behavioral performance is well illustrated by the gap saccade task, in which saccades might be initiated with short latency (express saccade) or with long latency (regular saccade) even though the external visual condition is identical. Accumulated evidence has demonstrated that the internal brain state is different before the initiation of an express saccade than a regular saccade. However, the reported origin of the fluctuation of internal brain state is disputed among previous studies, e.g., the fixation disengagement theory versus the oculomotor preparation theory. The present study examined these two theories by analyzing the rate and direction of fixational saccades, i.e., small amplitude saccades during fixation, because they could be modulated by the internal brain state. Since fixation disengagement is not spatially tuned, it might affect the rate but not the direction of fixational saccades. In contrast, oculomotor preparation can contain spatial information for the upcoming saccade and thus affect fixational saccade direction. We found that the different spatiotemporal characteristics of fixational saccades among tasks with different gap durations reveal diverse driving force to change the internal brain state. Under short gap duration (100 ms), fixation disengagement plays a primary role in switching internal brain states. Conversely, oculomotor preparation plays a primary role under medium (200 ms) and long (400 ms) gap durations. These results suggest that both fixation disengagement and oculomotor preparation can change the internal brain state, but their relative contributions are gap-duration dependent.NEW & NOTEWORTHY While performing the gap saccade task, the role of fixation disengagement and oculomotor preparation in modulating the internal brain state is gap-duration dependent. Fixation disengagement plays a primary role when gap duration is shorter (100 ms), whereas oculomotor preparation plays a primary role when gap duration is longer (200 ms and 400 ms).

Cerebellar complex spikes multiplex complementary behavioral information.

Purkinje cell (PC) discharge, the only output of cerebellar cortex, involves 2 types of action potentials, high-frequency simple spikes (SSs) and low-frequency complex spikes (CSs). While there is consensus that SSs convey information needed to optimize movement kinematics, the function of CSs, determined by the PC’s climbing fiber input, remains controversial. While initially thought to be specialized in reporting information on motor error for the subsequent amendment of behavior, CSs seem to contribute to other aspects of motor behavior as well. When faced with the bewildering diversity of findings and views unraveled by highly specific tasks, one may wonder if there is just one true function with all the other attributions wrong? Or is the diversity of findings a reflection of distinct pools of PCs, each processing specific streams of information conveyed by climbing fibers? With these questions in mind, we recorded CSs from the monkey oculomotor vermis deploying a repetitive saccade task that entailed sizable motor errors as well as small amplitude saccades, correcting them. We demonstrate that, in addition to carrying error-related information, CSs carry information on the metrics of both primary and small corrective saccades in a time-specific manner, with changes in CS firing probability coupled with changes in CS duration. Furthermore, we also found CS activity that seemed to predict the upcoming events. Hence PCs receive a multiplexed climbing fiber input that merges complementary streams of information on the behavior, separable by the recipient PC because they are staggered in time.

Characteristics of saccades during the near point of convergence test

The near point of convergence test is widely used to evaluate binocular vision. It assesses the ability of the eyes to converge at short distances. Although the test consists of a pure symmetrical vergence task, small involuntary saccades occur concurrently. The main goal of this study was to analyze saccadic characteristics as a function of vergence demand when testing the near point of convergence. To this purpose, the eye movements of 11 participants were registered with an eye-tracker while they performed the near point of convergence test by following a target that traveled forward and backward on a motorized bench. Saccade amplitude increased and, on average, saccade rate decreased with vergence demand. In general, the direction of the concurrent vergence movement had no significant effect on saccade characteristics. However, each individual subject showed idiosyncratic behavior. Most saccades tended to be corrective in terms of both binocular disparity and individual fixation position errors. In particular, most participants tended to correct the fixation position error of the dominant eye.

Dissociable Cortical and Subcortical Mechanisms for Mediating the Influences of Visual Cues on Microsaccadic Eye Movements.

Visual selection in primates is intricately linked to eye movements, which are generated by a network of cortical and subcortical neural circuits. When visual selection is performed covertly, without foveating eye movements toward the selected targets, a class of fixational eye movements, called microsaccades, is still involved. Microsaccades are small saccades that occur when maintaining precise gaze fixation on a stationary point, and they exhibit robust modulations in peripheral cueing paradigms used to investigate covert visual selection mechanisms. These modulations consist of changes in both microsaccade directions and frequencies after cue onsets. Over the past two decades, the properties and functional implications of these modulations have been heavily studied, revealing a potentially important role for microsaccades in mediating covert visual selection effects. However, the neural mechanisms underlying cueing effects on microsaccades are only beginning to be investigated. Here we review the available causal manipulation evidence for these effects’ cortical and subcortical substrates. In the superior colliculus (SC), activity representing peripheral visual cues strongly influences microsaccade direction, but not frequency, modulations. In the cortical frontal eye fields (FEF), activity only compensates for early reflexive effects of cues on microsaccades. Using evidence from behavior, theoretical modeling, and preliminary lesion data from the primary visual cortex and microstimulation data from the lower brainstem, we argue that the early reflexive microsaccade effects arise subcortically, downstream of the SC. Overall, studying cueing effects on microsaccades in primates represents an important opportunity to link perception, cognition, and action through unaddressed cortical-subcortical neural interactions. These interactions are also likely relevant in other sensory and motor modalities during other active behaviors.

Learning to silence saccadic suppression

Perceptual stability is facilitated by a decrease in visual sensitivity during rapid eye movements, called saccadic suppression. While a large body of evidence demonstrates that saccadic programming is plastic, little is known about whether the perceptual consequences of saccades can be modified. Here, we demonstrate that saccadic suppression is attenuated during learning on a standard visual detection-in-noise task, to the point that it is effectively silenced. Across a period of 7 days, 44 participants were trained to detect brief, low-contrast stimuli embedded within dynamic noise, while eye position was tracked. Although instructed to fixate, participants regularly made small fixational saccades. Data were accumulated over a large number of trials, allowing us to assess changes in performance as a function of the temporal proximity of stimuli and saccades. This analysis revealed that improvements in sensitivity over the training period were accompanied by a systematic change in the impact of saccades on performance-robust saccadic suppression on day 1 declined gradually over subsequent days until its magnitude became indistinguishable from zero. This silencing of suppression was not explained by learning-related changes in saccade characteristics and generalized to an untrained retinal location and stimulus orientation. Suppression was restored when learned stimulus timing was perturbed, consistent with the operation of a mechanism that temporarily reduces or eliminates saccadic suppression, but only when it is behaviorally advantageous to do so. Our results indicate that learning can circumvent saccadic suppression to improve performance, without compromising its functional benefits in other viewing contexts.

Concomitant improvement in anti-saccade success rate and postural instability gait difficulty after rTMS treatment for Parkinson\u2019s disease

Parkinson’s disease (PD) is a progressive neurological disorder characterised by motor and non-motor deficits. Repetitive transcranial magnetic stimulation (rTMS) over the bilateral primary motor cortex at a high frequency (5 Hz or higher) is reported to be a potential treatment of PD. We aimed to assess the effect of rTMS on eye movement control in patients with PD in their ‘on’ state. We enrolled 14 patients with PD and assessed motor symptoms (Movement Disorder Society-Sponsored Unified Parkinson’s Disease Rating Scale; MDS-UPDRS) and eye movement performances (visually guided saccades, volitional anti-saccades, and small involuntary saccades during fixation) at baseline and after administering bilateral 10 Hz rTMS on leg region of the motor cortex. We confirmed that rTMS improved the MDS-UPDRS motor scores and found that rTMS improved the anti-saccade success rate, which requires adequate inhibition of the reflexive response. The improvement in anti-saccade success rate was correlated with that of the postural instability gait difficulty (PIGD) sub-scores of MDS-UPDRS and lower baseline Japanese version of the Montreal Cognitive Assessment scores. This result is consistent with previous findings that PIGD and inhibitory control deficits share common brain dysfunctions in PD. rTMS may alleviate dysfunctions of that circuit and have a clinical effect.

Regression-based analysis of combined EEG and eye-tracking data: Theory and applications.

Fixation-related potentials (FRPs), neural responses aligned to the end of saccades, are a promising tool for studying the dynamics of attention and cognition under natural viewing conditions. In the past, four methodological problems have complicated the analysis of such combined eye-tracking/electroencephalogram experiments: (1) the synchronization of data streams, (2) the removal of ocular artifacts, (3) the condition-specific temporal overlap between the brain responses evoked by consecutive fixations, and (4) the fact that numerous low-level stimulus and saccade properties also influence the postsaccadic neural responses. Although effective solutions exist for the first two problems, the latter two are only beginning to be addressed. In the current paper, we present and review a unified regression-based framework for FRP analysis that allows us to deconvolve overlapping potentials while also controlling for both linear and nonlinear confounds on the FRP waveform. An open software implementation is provided for all procedures. We then demonstrate the advantages of this proposed (non)linear deconvolution modeling approach for data from three commonly studied paradigms: face perception, scene viewing, and reading. First, for a traditional event-related potential (ERP) face recognition experiment, we show how this technique can separate stimulus ERPs from overlapping muscle and brain potentials produced by small (micro)saccades on the face. Second, in natural scene viewing, we model and isolate multiple nonlinear effects of saccade parameters on the FRP. Finally, for a natural sentence reading experiment using the boundary paradigm, we show how it is possible to study the neural correlates of parafoveal preview after removing spurious overlap effects caused by the associated difference in average fixation time. Our results suggest a principal way of measuring reliable eye movement-related brain activity during natural vision.