Saccade Initiation Research Articles

Saliency response in superior colliculus at the future saccade goal predicts fixation duration during free viewing of dynamic scenes.

Eye movements in daily life occur in rapid succession and often without a predefined goal. Using a free viewing task, we examined how fixation duration prior to a saccade correlates to visual saliency and neuronal activity in the superior colliculus (SC) at the saccade goal. Rhesus monkeys (three male) watched videos of natural, dynamic, scenes while eye movements were tracked and, simultaneously, neurons were recorded in the superficial and intermediate layers of the superior colliculus (SCs and SCi respectively), a midbrain structure closely associated with gaze, attention, and saliency coding. Saccades that were directed into the neuron's receptive field (RF) were extrapolated from the data. To interpret the complex visual input, saliency at the RF location was computed during the pre-saccadic fixation period using a computational saliency model. We analyzed if visual saliency and neural activity at the saccade goal predicted pre-saccadic fixation duration. We report three major findings: 1) Saliency at the saccade goal inversely correlated with fixation duration, with motion and edge information being the strongest predictors. 2) SC visual saliency responses in both SCs and SCi were inversely related to fixation duration. 3) SCs neurons, and not SCi neurons, showed higher activation for two consecutive short fixations, suggestive of concurrent saccade processing during free viewing. These results reveal a close correspondence between visual saliency, SC processing, and the timing of saccade initiation during free viewing and are discussed in relation to their implication for understanding saccade initiation during real-world gaze behavior.Significance statement Contrary to traditional controlled stimuli/task studies, eye movements in day-to-day life are not discrete events but occur in (rapid) succession and often without a predefined goal. Therefore, the study of visual processing during free viewing of dynamic scenes is an essential step in understanding visual processing in its functional context. We present an investigation into saliency and visual responses in the superior colliculus (SC) during task-free viewing of dynamic videos and their correspondence to saccade initiation. In short, these results show the correspondence between fixation duration, pre-saccadic visual saliency at the saccade goal and SC processing and provide first evidence of a neural correlate of concurrent visual processing across a chain of saccades in the SC during free viewing.

Early visual modulation and selection predict saccadic timing during visual search: An ERP study.

Saccadic eye movements, a critical aspect of real-world visual behavior, are preceded by an initial accumulation of visual information followed by the selection of a single location to move one's eyes. However, it is currently unclear how each of these stages uniquely affects saccadic timing. In this study, participants searched for a contour integration target while EEG was used to measure posterior cortical activity between search display onset and first saccade initiation. The goal was to determine whether saccade timing could be attributed to differences in early ERP amplitudes, with the P1 reflecting the magnitude of early perceptual information accumulation and the N1 reflecting the strength of selection leading to the saccadic decision. EOG was used to measure saccade timing, and trials were divided into fast, middle, and slow bins. The N1 response was smallest in the slow saccade tertile, relative to both the fast and middle tertiles, suggesting weak selection. In contrast, the P1 response was largest for this same slow saccadic tertile relative to the middle saccadic tertile, suggesting vigorous information accumulation. Therefore, delays in saccadic behavior may occur when the visual system is overwhelmed with visual input, thus increasing the time to reach a saccadic decision. These findings reconcile models of eye movement behavior which often prioritize either the impact of information accrual or selection, rather than regarding both as an integrated whole.

Dynamics of Saccade Trajectory Modulation by Distractors: Neural Activity Patterns in the Frontal Eye Field.

The sudden appearance of a visual distractor shortly before saccade initiation can capture spatial attention and modulate the saccade trajectory in spite of the ongoing execution of the initial plan to shift gaze straight to the saccade target. To elucidate the neural correlates underlying these curved saccades, we recorded from single neurons in the frontal eye field of two male rhesus monkeys shifting gaze to a target while a distractor with the same eccentricity appeared either left or right of the target at various delays after target presentation. We found that the population level of presaccadic activity of neurons representing the distractor location encoded the direction of the saccade trajectory. Stronger activity occurred when saccades curved toward the distractor, and weaker when saccades curved away. This relationship held whether the distractor was ipsilateral or contralateral to the recorded neurons. Meanwhile, visually responsive neurons showed asymmetrical patterns of excitatory responses that varied with the location of the distractor and the duration of distractor processing relating to attentional capture and distractor inhibition. During earlier distractor processing, neurons encoded curvature toward the distractor. During later distractor processing, neurons encoded curvature away from the distractor. This was observed when saccades curved away from distractors contralateral to the recording site and when saccades curved toward distractors ipsilateral to the recording site. These findings indicate that saccadic motor planning involves dynamic push-pull hemispheric interactions producing attraction or repulsion for potential but unselected saccade targets.

Case report on interesting cases of congenital ocular motor apraxia

Congenital ocular motor apraxia (COMA) is an uncommon disorder of ocular motility characterized by inability to initiate voluntary horizontal saccades with associated head thrusts during horizontal fixation shifts. In addition, when ocular movements typical of COMA are observed, detailed systemic and neurologic examination should be performed to rule out any associated congenital neurologic or metabolic disorder. We report two cases of COMA, a 14-year-old male with diminution of vision, esotropia, and his face turned to the right without any systemic disease, and a 9-year-old male with esotropia, no face turn, acute lymphocytic leukemia, and isolated complete agenesis of the corpus callosum. Both cases had abnormal saccadic initiation for horizontal gaze with head thrusts to take fixation. Both cases underwent Medial rectus (MR) recession, which improved face turn and strabismus, respectively.

Fast saccades to faces during the feedforward sweep.

Saccadic choice tasks use eye movements as a response method, typically in a task where observers are asked to saccade as quickly as possible to an image of a prespecified target category. Using this approach, face-selective saccades have been observed within 100ms poststimulus. When taking into account oculomotor processing, this suggests that faces can be detected in as little as 70 to 80ms. It has therefore been suggested that face detection must occur during the initial feedforward sweep, since this latency leaves little time for feedback processing. In the current experiment, we tested this hypothesis using backward masking-a technique shown to primarily disrupt feedback processing while leaving feedforward activation relatively intact. Based on minimum saccadic reaction time, we found that face detection benefited from ultra-fast, accurate saccades within 110 to 160ms and that these eye movements are obtainable even under extreme masking conditions that limit perceptual awareness. However, masking did significantly increase the median SRT for faces. In the manual responses, we found remarkable detection accuracy for faces and houses, even when participants indicated having no visual experience of the test images. These results provide evidence for the view that the saccadic bias to faces is initiated by coarse information used to categorize faces in the feedforward sweep but that, in most cases, additional processing is required to quickly reach the threshold for saccade initiation.

Bidirectional brain-body interactions during natural story listening

Narratives can synchronize neural and physiological signals between individuals, but the relationship between these signals, and the underlying mechanism, is unclear. We hypothesized a top-down effect of cognition on arousal and predicted that auditory narratives will drive not only brain signals but also peripheral physiological signals. We find that auditory narratives entrained gaze variation, saccade initiation, pupil size, and heart rate. This is consistent with a top-down effect of cognition on autonomic function. We also hypothesized a bottom-up effect, whereby autonomic physiology affects arousal. Controlled breathing affected pupil size, and heart rate was entrained by controlled saccades. Additionally, fluctuations in heart rate preceded fluctuations of pupil size and brain signals. Gaze variation, pupil size, and heart rate were all associated with anterior-central brain signals. Together, these results suggest bidirectional causal effects between peripheral autonomic function and central brain circuits involved in the control of arousal.

Abnormal Oculomotor Corollary Discharge Signaling as a Trans-diagnostic Mechanism of Psychosis.

Corollary discharge (CD) signals are "copies" of motor signals sent to sensory areas to predict the corresponding input. They are a posited mechanism enabling one to distinguish actions generated by oneself vs external forces. Consequently, altered CD is a hypothesized mechanism for agency disturbances in psychosis. Previous studies have shown a decreased influence of CD signals on visual perception in individuals with schizophrenia-particularly in those with more severe positive symptoms. We therefore hypothesized that altered CD may be a trans-diagnostic mechanism of psychosis. We examined oculomotor CD (using the blanking task) in 49 participants with schizophrenia or schizoaffective disorder (SZ), 36 bipolar participants with psychosis (BPP), and 40 healthy controls (HC). Participants made a saccade to a visual target. Upon saccade initiation, the target disappeared and reappeared at a horizontally displaced position. Participants indicated the direction of displacement. With intact CD, participants can make accurate perceptual judgements. Otherwise, participants may use saccade landing site as a proxy of pre-saccadic target to inform perception. Thus, multi-level modeling was used to examine the influence of target displacement and saccade landing site on displacement judgements. SZ and BPP were equally less sensitive to target displacement than HC. Moreover, regardless of diagnosis, SZ and BPP with more severe positive symptoms were more likely to rely on saccade landing site. These results suggest that altered CD may be a trans-diagnostic mechanism of psychosis.

Delayed ocular disengagement from arousing scenes.

Visual exploration of the world is supported by eye movements which can be speeded up or delayed depending on bottom-up stimulation, top-down goals, and prior associations. Previous studies observed faster initiation of saccades toward emotional than neutral natural scenes; however, less is known concerning saccades which originate from emotional, compared with neutral, scenes. Here, we addressed this issue by examining a task in which participants continuously moved their gaze from and toward pictures (natural scenes), which could be emotional or neutral, and changed position in every trial. Saccades were initiated later when the starting picture was emotional compared to neutral, and this slowing was associated with the arousal value of the picture, suggesting that ocular disengagement does not vary with stimulus valence but is affected by engaging picture contents such as erotica and threat/injuries.

Neuronal activity in posterior parietal cortex area LIP is not sufficient for saccadic eye movement production

It is widely recognized that the posterior parietal cortex (PPC) plays a role in active exploration with eye movements, arm reaching, and hand grasping. Whether this role is causal in nature is largely unresolved. One region of the PPC appears dedicated to the control of saccadic eye movement—lateral intraparietal (LIP) area. This area LIP possesses direct projections to well-established oculomotor centers and contains neurons with movement-related activity. In this study, we tested whether these neurons are implicated in saccade initiation and production. The movement-related activity of LIP neurons was tested by recording these neurons while monkeys performed a countermanding task. We found that LIP neuronal activity is not different before the execution or the cancelation of commanded saccades and thereby is not sufficient for the initiation and production of saccades. Consistent with the evolutionarily late emergence of the PPC, this finding relegates the role of this PPC area to processes that can regulate but not trigger eye movements.

Resilience of FEF neuronal saccade code to V4 perturbations.

The frontal eye field (FEF) plays a key role in initiating rapid eye movements known as saccades. Accumulation models have been proposed to explain the dynamic of these neurons and how they may enable the initiation of saccades. To update the scope of the viability of this model, we studied single neurons recorded from the FEF of two rhesus monkeys while they performed a memory-guided saccade task. We evaluated the degree to which each type of FEF neuron complied with these models by quantifying how precisely their discharge predicted an imminent saccade based on their immediate presaccadic activity. We found that decoders trained on single neurons with a stronger motor component performed better than decoders trained on neurons with a stronger visual component in predicting the saccade. Importantly, despite a dramatic effect on the reaction times, the perturbations delivered to the FEF neurons via area V4 did not impact their saccade predictability. Our results demonstrate a high degree of resilience of the FEF neuronal presaccadic discharge patterns, fulfilling the predictions of accumulation models.NEW & NOTEWORTHY We studied neurons in the brain's frontal eye field (FEF) to understand how these neurons predict swift eye shifts called saccades. We found that neurons with more movement-related activity were better at predicting saccades than those with sensory-related activity. Interestingly, electrical disruptions of this region strongly impacted saccade onset times but did not affect the individual neuron's saccade predictability, consistent with models suggesting that a specific threshold in neural activity triggers the saccade.

Inferential eye movement control while following dynamic gaze.

Attending to other people's gaze is evolutionary important to make inferences about intentions and actions. Gaze influences covert attention and triggers eye movements. However, we know little about how the brain controls the fine-grain dynamics of eye movements during gaze following. Observers followed people's gaze shifts in videos during search and we related the observer eye movement dynamics to the time course of gazer head movements extracted by a deep neural network. We show that the observers' brains use information in the visual periphery to execute predictive saccades that anticipate the information in the gazer's head direction by 190-350 ms. The brain simultaneously monitors moment-to-moment changes in the gazer's head velocity to dynamically alter eye movements and re-fixate the gazer (reverse saccades) when the head accelerates before the initiation of the first forward gaze-following saccade. Using saccade-contingent manipulations of the videos, we experimentally show that the reverse saccades are planned concurrently with the first forward gaze-following saccade and have a functional role in reducing subsequent errors fixating on the gaze goal. Together, our findings characterize the inferential and functional nature of social attention's fine-grain eye movement dynamics.

The effect of impaired velocity signals on goal-directed eye and hand movements

Information about position and velocity is essential to predict where moving targets will be in the future, and to accurately move towards them. But how are the two signals combined over time to complete goal-directed movements? We show that when velocity information is impaired due to using second-order motion stimuli, saccades directed towards moving targets land at positions where targets were ~ 100 ms before saccade initiation, but hand movements are accurate. Importantly, the longer latencies of hand movements allow for additional time to process the sensory information available. When increasing the period of time one sees the moving target before making the saccade, saccades become accurate. In line with that, hand movements with short latencies show higher curvature, indicating corrections based on an update of incoming sensory information. These results suggest that movements are controlled by an independent and evolving combination of sensory information about the target’s position and velocity.

Visual Development in a Patient with Infantile-Onset Saccade Initiation Delay

Congenital ocular motor apraxia (COMA), also known as infantile-onset saccade initiation delay (ISID), is characterized by the inability to initiate volitional horizontal saccades. This five-month-old girl showed constant rapid movements to right or left one week after birth. After four months, she began thrusting her head from left to right when following target objects. She was diagnosed with COMA/ISID. At 4 years of age, head thrust became less evident. She first uttered meaningful words at 5 years of age, and spoke in sentences by 6 years of age. At 6 years of age, visual acuity was 1.0 in both eyes at 1 m. Head thrust disappeared at almost 6 years of age. She could not start crawling and sustaining a sitting position due to hypotonia. Clinicians should be aware of visual development in COMA/ISID and should look for them during the assessment of these children.

POLG-Related Ataxia and Ophthalmoplegia: A Case Report (P9-11.008)

<h3>Objective:</h3> Not applicable <h3>Background:</h3> Genetic variants of the POLG gene result in mitochondrial DNA defects and account for a large proportion of mitochondrial diseases. These disorders encompass a spectrum of phenotypes, with onset ranging from infancy to late adulthood. In addition to hepatic dysfunction and myopathy in younger populations, and hormone derangements with cataracts in older patients, patients may also present with prominent neurologic symptoms, including ataxia, parkinsonism, epilepsy, and ophthalmoplegia. More than 300 pathogenic mutations of POLG have been reported and there are no clear genotype-phenotype correlations. Given the rarity of these syndromes and wide range of clinical phenotypes, there is often delay in diagnosis. <h3>Design/Methods:</h3> Not applicable <h3>Results:</h3> We present a 47-year old woman with symptoms of difficulty speaking and development of tremor, vision loss, dizziness, and gait difficulty progressing over several months. Initial exam was significant for ataxia and choreiform movements. Extraocular movement (EOM) testing revealed delayed initiation of saccades and slow, saccadic pursuits with nystagmus. Bloodwork, which included extensive workup for causes of chorea and ataxia, electromyography/nerve conduction studies, and imaging (MRI brain and malignancy work-up) were unremarkable. She continued to decline with development of ophthalmoplegia, ptosis, numbness, weakness, and pain, severely limiting her functional capacity. Whole exome sequencing was performed and showed a heterozygous variant of uncertain significance (VUS) in the POLG gene, which may explain her presentation. <h3>Conclusions:</h3> This case adds to a growing collection of case reports describing the various phenotypes of POLG-related disorders. Early detection of POLG variants and phenotype/genotype characterization may help guide development of future treatments for patients with similar conditions. <b>Disclosure:</b> Dr. Peabody has nothing to disclose. An immediate family member of Dr. Shpiner has received personal compensation for serving as an employee of University of Miami.

Neurologically Healthy Humans’ Ability to Make Saccades Toward Unseen Targets

Some patients with a visual field loss due to a lesion in the primary visual cortex (V1) can shift their gaze to stimuli presented in their blind visual field. The extent to which a similar “blindsight” capacity is present in neurologically healthy individuals remains unknown. Using retinotopically navigated transcranial magnetic stimulation (TMS) of V1 (Experiment 1) and metacontrast masking (Experiment 2) to suppress conscious vision, we examined neurologically healthy humans’ ability to make saccadic eye movements toward visual targets that they reported not seeing. In the TMS experiment, the participants were more likely to initiate a saccade when a stimulus was presented, and they reported not seeing it, than in trials which no stimulus was presented. However, this happened only in a very small proportion (∼8%) of unseen trials, suggesting that saccadic reactions were largely based on conscious perception. In both experiments, saccade landing location was influenced by unconscious information: When the participants denied seeing the target but made a saccade, the saccade was made toward the correct location (TMS: 68%, metacontrast: 63%) more often than predicted by chance. Signal detection theoretic measures suggested that in the TMS experiment, saccades toward unseen targets may have been based on weak conscious experiences. In both experiments, reduced visibility of the target stimulus was associated with slower and less precise gaze shifts. These results suggest that saccades made by neurologically healthy humans may be influenced by unconscious information, although the initiation of saccades is largely based on conscious vision.

Binocular, accommodative and oculomotor alterations in multiple sclerosis

AbstractMultiple sclerosis (MS) is an acquired demyelinating and inflammatory neurodegenerative disease affecting the central nervous system (CNS). Clinical and subclinical ocular disturbances occur in almost all patients with MS. Among them, oculomotor, accommodative and binocular alterations have been reported in MS. Our research group has conducted a complete analysis of the visual and oculomotor function in several researches with MS patients conducted in the last 4 years, obtaining a characterization of some visual aspects of the patient with MS. Concerning the binocular function, several alterations have been detected in MS patients, with divergence at near distance and stereopsis the most affected parameters. Likewise, MS patients with previous optic neuritis present worse binocular vision. Symptomatology compatible with convergence insufficiency measured with the CISS (Convergence Insufficiency Symptom Survey) is frequently reported by MS patients, but not supported by signs that show clear trend to esophoria at near. This discrepancy between the signs and symptoms could be due to the peculiarities of the CISS questionnaire, including items that are also related to the cognitive function and other ocular abnormalities.Oculomotor alterations have been reported also in patients with multiple sclerosis (MS), especially saccadic dysfunctions. The indirect analysis of horizontal saccades with the Developmental Eye Movement (DEM) and King‐Devick tests shows that are affected by MS, with more level of alteration if there has been a previous episode of optic neuritis. Similar findings have been also reported by other authors using objective eye tracking or video‐oculographic systems. Specifically, the most reported saccadic alterations in advanced MS and clinically isolated syndrome (CIS) are saccadic dysmetry (41.7%) and impaired smooth pursuit (42.3%), which relates to prolonged saccadic latency, no matter if they have suffered or not optic neuritis or internuclear ophthalmoplegia (INO). An increase in saccadic latency has been even found in children with MS. Saccadic initiation time (SI time) and average inter‐saccadic intervals (ISI) are also higher in patients with MS. Furthermore, the first cause of acquired pendular nystagmus is MS.Finally, macular sensitivity measured by microperimetry has been found to be altered in MS patients, especially in eyes with previous optic neuritis. Likewise, a fixational instability characterized also my microperimetry is present in MS patients with optic neuritis, with more increase of the vertical axis of the fixation area than of the horizontal. The ratio of the disease also affects the patient fixation pattern.

Saccade suppression of displacements, but not of contrast, depends on context.

Saccades let the visual scene sweep with high speed across the retina, thus producing a massive motion stimulus. Yet, in natural vision, we never perceive motion that is produced by saccades. The absence of perisaccadic motion perception might be caused by a transient reduction of visual sensitivity at the time of saccade initiation, so-called saccadic suppression. Saccade suppression occurs for contrast, displacement, and motion stimuli. Saccade suppression of displacements has been shown to be context sensitive. After performing saccades in sessions without perisaccadic stimulation, saccade suppression magnitude is drastically decreased (Zimmermann, 2020). Here, we aimed to test whether saccade suppression of contrast is similarly modulated by context. To this end, we projected stimuli on a homogeneously white wall such that we could establish a ganzfeld-like environment that, depending on the experimental session, did or did not contain any visible contrast stimuli. We first successfully replicated the context sensitivity of saccade suppression of displacements. Then, we tested context sensitivity of contrast suppression by asking subjects to perform several saccades either across the uniform white wall or across a background consisting of a sinusoidal grating. In contrast to perisaccadic context sensitivity for displacement suppression, we did not find context sensitivity for suppression of contrast.

Saccadic initiation biased by fixational activity

Both the gap and overlap paradigm may reveal the interaction between fixating and moving the eyes, but the effects of the overlap paradigm have not been fully characterized yet. Here we present a series of experiments probing how an overlap paradigm, combined with the manipulation of stimuli durations, saliency and transient changes might modulate saccadic reaction time distributions.We recorded saccadic reaction time in four participants in six experiments in which a saccade-target appeared at a pseudo-random amplitude after a fixation period. First, we parametrically manipulated the duration of the overlap using a range of intervals (from 0 to 200 ms). In a second experiment we probed the interaction of various foreperiod intervals (i.e. the duration of the fixation period prior to saccade-target onset) and overlap using two overlap intervals (20 or 140 ms). In two additional experiments we manipulated either the stimuli sizes or their contrast ratio in overlap paradigms (20 or 140 ms). Lastly, we introduced a visual transient during the overlap interval via two manipulations (both with a range of SOA): either a distractor ring appeared around the fixation-target, or a dynamic random noise patch replaced the fixation-target.Results show reliable modifications in the latency distributions depending on the overlap interval as well as idiosyncratic differences. Additional experimental manipulations also affected the latency distributions revealing strong interacting inhibitory processes. We conclude that the effects of overlap intervals may combine with the influence of other stimuli properties affecting decision process.

Saccadic premotor burst neurons and histochemical correlates of their firing patterns in rhesus monkey

Bursting behavior of brainstem premotor burst neurons (BNs) is essential for initiation of saccades and calibrating their metrics. Several ion channel families such as voltage-gated potassium (Kv) channels, low-voltage-activated calcium (Cav3) channels and hyperpolarization-activated cyclic nucleotide–gated (HCN) channels are major regulators of the bursting in neurons. Therefore, it was speculated that ion channels with rapid kinematics are essential for characteristic firing patterns of the BNs and rapid saccade velocities. However, the expression patterns of ion channels are yet to be confirmed. Confirmation would not only support the neuromimetic model predictions for saccade generation in brainstem, but also contemporary views that channelopathies can cause saccade disorders in humans. As proof of concept, we examined excitatory BNs in the rostral interstitial nucleus of medial longitudinal fasciculus (RIMLF, vertical saccades) and inhibitory BNs in nucleus paragigantocellularis dorsalis (PGD, horizontal saccades) histochemically in macaque monkeys. We found strong expression of Kv channels, which enable rapid-firing, as well as HCN1&2 and Cav3.2&3.3, which enable post-inhibitory rebound bursting, in both BN populations. Moreover, PGD was found to host multiple neuron groups in terms of calretinin immunoreactivity. Our results provide histochemical evidence that supports models proposing post-inhibitory rebound facilitates bursting in BNs. Furthermore, our findings support the notion that deductions can be made about electrophysiological firing properties by histochemical examination of functional groups within the brainstem saccadic circuitry. This development is an important building block supporting the concept of channelopathies in saccadic disorders. Future histological studies in humans will confirm this approach for saccadic disorders.

LAG-1: A dynamic, integrative model of learning, attention, and gaze.

It is clear that learning and attention interact, but it is an ongoing challenge to integrate their psychological and neurophysiological descriptions. Here we introduce LAG-1, a dynamic neural field model of learning, attention and gaze, that we fit to human learning and eye-movement data from two category learning experiments. LAG-1 comprises three control systems: one for visuospatial attention, one for saccadic timing and control, and one for category learning. The model is able to extract a kind of information gain from pairwise differences in simple associations between visual features and categories. Providing this gain as a reentrant signal with bottom-up visual information, and in top-down spatial priority, appropriately influences the initiation of saccades. LAG-1 provides a moment-by-moment simulation of the interactions of learning and gaze, and thus simultaneously produces phenomena on many timescales, from the duration of saccades and gaze fixations, to the response times for trials, to the slow optimization of attention toward task relevant information across a whole experiment. With only three free parameters (learning rate, trial impatience, and fixation impatience) LAG-1 produces qualitatively correct fits for learning, behavioural timing and eye movement measures, and also for previously unmodelled empirical phenomena (e.g., fixation orders showing stimulus-specific attention, and decreasing fixation counts during feedback). Because LAG-1 is built to capture attention and gaze generally, we demonstrate how it can be applied to other phenomena of visual cognition such as the free viewing of visual stimuli, visual search, and covert attention.