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Saccade

A saccade is a rapid, ballistic eye movement that shifts the point of fixation from one location to another in the visual field, enabling high-acuity vision by aligning the fovea with objects of interest.^[1] These movements are conjugate, meaning both eyes move together in the same direction, and they occur multiple times per second during normal visual scanning, with durations typically ranging from 20 to 100 milliseconds and peak velocities up to 700 degrees per second.^[2] Unlike smooth pursuit, vision is suppressed during the saccade itself to prevent perceptual blurring, a phenomenon known as saccadic masking.^[3] Saccades are essential for exploring the visual environment and are classified into several types based on their triggers and purposes. Reflexive saccades are involuntary responses to sudden stimuli, such as a flashing light, with latencies around 150-250 milliseconds.^[2] Voluntary saccades, in contrast, are goal-directed and include prosaccades (direct shifts to a target), antisaccades (shifts in the opposite direction to suppress reflexive responses), memory-guided saccades (to recalled locations), and predictive saccades (anticipating a target's movement).^[1] Smaller variants, known as microsaccades (less than 0.5 degrees), occur during fixation to counteract visual fading and maintain perceptual stability, happening about once per second.^[2] The neural control of saccades involves a distributed network in the brain, integrating sensory input with motor output for precise execution. High-level planning occurs in cortical areas like the frontal eye fields (Brodmann's area 8), which initiate voluntary saccades, while the superior colliculus in the midbrain serves as a key integrator for both reflexive and voluntary movements, mapping sensory stimuli to motor commands.^[3] Burst neurons in brainstem structures, such as the paramedian pontine reticular formation for horizontal saccades and the rostral interstitial nucleus of the medial longitudinal fasciculus for vertical ones, generate the high-frequency signals needed for rapid eye acceleration.^[1] The amplitude and direction are encoded by the duration and pattern of activity in oculomotor nuclei, ensuring conjugate movement without mid-flight corrections due to the ballistic nature of the process.^[3] Clinically, saccade abnormalities provide diagnostic insights into neurological disorders, as their metrics like latency, velocity, and accuracy reflect integrity of oculomotor pathways. Hypometric (shortened) saccades are common in Parkinson's disease, while slow or absent vertical saccades characterize progressive supranuclear palsy.^[4] Impairments also appear in neuropsychiatric conditions, such as increased latencies in attention-deficit/hyperactivity disorder or erratic patterns in schizophrenia, underscoring saccades' role as a biomarker for brain function.^[1]

Fundamentals

Definition and Function

A saccade is a rapid, ballistic movement of both eyes that abruptly shifts the point of fixation from one location to another in the visual field.^[5] These movements are conjugate, meaning the eyes move together in the same direction, and they typically last between 20 and 200 milliseconds, depending on their size.^[6] Saccades range in amplitude from less than 1 degree for small shifts, such as during reading, to up to 90 degrees for large gaze changes when scanning a broad environment.^[5] The primary function of saccades is to direct the fovea—the central, high-acuity region of the retina—toward objects or features of interest, enabling detailed visual processing where peripheral vision lacks resolution.^[3] This compensates for the retina's nonuniform sensitivity, allowing efficient sampling of the visual scene without relying solely on head or body movements.^[7] Evolutionarily, saccades facilitate active exploration of the visual environment, supporting survival by quickly orienting gaze to potential threats, rewards, or relevant stimuli in a dynamic world.^[8] Key characteristics distinguish saccades from other eye movements, such as smooth pursuit or vestibular reflexes. Saccades exhibit peak velocities reaching up to 700 degrees per second for larger amplitudes, with acceleration and deceleration phases that follow a stereotyped, nonlinear profile known as the main sequence.^[6] This ballistic nature ensures precise, high-speed repositioning, executed approximately three to four times per second during active vision.^[7]

Types of Saccades

Saccades are categorized into distinct types based on their triggers, cognitive involvement, and functional roles, reflecting the diversity of oculomotor control mechanisms. Reflexive saccades, also known as exogenous saccades, are rapid, automatic responses elicited by the sudden onset of a peripheral visual stimulus, such as a flashing light or abrupt target appearance. These movements orient the eyes toward novel or salient environmental cues with relatively short latencies, typically around 150-250 ms, facilitating immediate visual exploration.^[2] In contrast, voluntary or endogenous saccades are goal-directed movements initiated by internal cognitive intent, such as shifting gaze to a remembered location or following an instructed direction, without direct sensory prompting. These saccades involve higher-level planning and exhibit longer latencies, generally 200-250 ms, due to the integration of top-down attentional processes.^[2]^[9] Microsaccades represent small-amplitude, involuntary eye movements, typically less than 1 degree of visual angle, that occur during attempted visual fixation. Functioning to counteract retinal image adaptation and prevent perceptual fading, they occur at a rate of about 1-2 per second and help maintain visual stability by subtly repositioning the gaze. Express saccades are a specialized subset of reflexive saccades characterized by ultra-fast latencies of 70-120 ms, often triggered in paradigms involving a temporal gap between the offset of a central fixation point and the onset of a peripheral target. This rapid response is facilitated by direct collicular pathways, bypassing some cortical processing, and is more prevalent with repeated exposure or predictable stimuli.^[2]90760-6) Other notable variants include memory-guided saccades, which direct the eyes to a previously viewed but no longer visible target based on working memory representations, often showing reduced accuracy compared to visually guided ones; anti-saccades, which require voluntary suppression of a reflexive response to a stimulus and instead shifting gaze to the opposite direction, testing inhibitory control with latencies around 250-350 ms and error rates of 10-20%; and predictive saccades, anticipatory movements made in advance of an expected target appearance, such as in rhythmic or learned sequences, where timing relies on temporal expectations rather than immediate visual input. These latency differences among saccade types are tied to distinct neural processing pathways.^[2]90218-3)

Physiological Mechanisms

Timing and Kinematics

Saccadic eye movements exhibit characteristic temporal and dynamic properties that ensure rapid reorientation of gaze. The duration of a saccade, defined as the time from initiation to completion, typically ranges from 30 to 120 milliseconds, depending on the amplitude of the movement.^[10] This duration increases approximately linearly with saccade amplitude, following the main sequence relationship: duration ≈ 2.2 + 2.6 × amplitude (in degrees).^[2] Larger saccades thus take longer to execute, reflecting the ballistic nature of the movement where the eye accelerates and decelerates without mid-flight corrections. The velocity profile of a saccade forms a triangular waveform, characterized by an initial rapid acceleration phase followed by deceleration to zero velocity. Acceleration can reach up to 10,000 degrees per second squared, enabling quick attainment of peak velocity, which occurs roughly midway through the movement.^[11] Peak velocity (V_max) also adheres to the main sequence, increasing with amplitude but saturating for larger movements. More generally, this relationship can be modeled logarithmically as log(V_max) = log(k) + b × log(amplitude), where k and b are empirical constants derived from experimental data, capturing the nonlinear saturation observed in human saccades.^[12] Saccadic latency, the interval from stimulus onset to movement initiation, varies by task demands and typically measures 150-250 milliseconds for voluntary saccades.^[2] This delay encompasses visual processing and motor planning. The gap effect, where latency shortens when the central fixation point disappears before target onset, can reduce this time by 50-100 milliseconds, facilitating faster responses in certain paradigms.^[2] Saccades are not always perfectly accurate, often undershooting the target by 10-20% of the intended amplitude, which is subsequently corrected by smaller corrective saccades.^[13] In some contexts, dynamic overshoot may occur, where the eye briefly exceeds the target before settling. These errors highlight the trade-off between speed and precision inherent in saccadic control. These kinematic parameters—duration, velocity, latency, and accuracy—are measured using high-resolution eye-tracking devices, such as video-based systems or scleral search coils, which record eye position over time to compute velocity (via differentiation) and acceleration (second derivative).^[2] Such techniques allow precise quantification of saccade dynamics in both laboratory and clinical settings.

Neural Control

The neural control of saccades involves a distributed network of cortical and subcortical structures that plan, select, and execute rapid eye movements, ensuring precise foveation of visual targets. This system operates hierarchically, with higher cortical areas processing sensory inputs and cognitive demands to generate saccade commands, which are then relayed through subcortical pathways for motor implementation. Feedback mechanisms within this network refine accuracy and adapt to ongoing visual updates, integrating voluntary intention with reflexive responses.^[14] At the core of reflexive saccades lies the superior colliculus (SC), a midbrain structure that integrates multisensory inputs, including visual, auditory, and somatosensory signals, to trigger orienting movements. Neurons in the intermediate layers of the SC encode target locations in retinotopic coordinates and generate burst activity that directly projects to brainstem motor circuits, facilitating quick, stimulus-driven saccades. Electrical stimulation or lesion studies in primates confirm the SC's essential role, as its inactivation impairs reflexive but spares voluntary saccades.^[15]^[16] Voluntary saccades and their planning are primarily orchestrated by cortical regions, including the frontal eye fields (FEF) and supplementary eye fields (SEF). The FEF, located in the prefrontal cortex, plays a pivotal role in target selection during visual search tasks, where visuomotor neurons accumulate evidence about stimulus salience until reaching a threshold that initiates the saccade command. Projections from the FEF descend to the SC and brainstem, influencing both reflexive and goal-directed movements. In contrast, the SEF, situated on the medial frontal gyrus, contributes to sequencing multiple saccades and monitoring performance, with neurons showing sustained activity during planned eye movement chains.^[14]^[17] The parietal eye fields, particularly the lateral intraparietal area (LIP), support spatial attention and perceptual stability by remapping visual representations ahead of impending saccades. LIP neurons shift their receptive fields to anticipate the postsaccadic gaze position, enabling trans-saccadic continuity of object features and attention allocation to potential targets. This remapping process integrates with FEF and SC activity, ensuring that attentional priorities guide saccade metrics without disrupting visual perception.^[15]^[18] Execution of saccades relies on brainstem circuits that translate cortical and collicular commands into coordinated ocular motor output. The paramedian pontine reticular formation (PPRF) contains excitatory burst neurons that generate the high-velocity pulse for horizontal saccades, while the reticular formation integrates velocity and position signals to control eye trajectory. The medial longitudinal fasciculus (MLF) interconnects these brainstem nuclei with the oculomotor and abducens nuclei, synchronizing contractions of extraocular muscles for conjugate gaze shifts. Omnipause neurons in the raphe interpositus pause their tonic inhibition during saccades, gating the burst to prevent unwanted movements.^[15]^[16] Inhibitory control prevents reflexive saccades in tasks requiring suppression, such as the anti-saccade paradigm, where subjects look away from a sudden stimulus. The basal ganglia, via the substantia nigra pars reticulata (SNr), exert tonic inhibition on the SC; reduced SNr firing disinhibits SC neurons to permit saccades, while sustained activity blocks erroneous responses. Fixation-related neurons in the SC further enhance suppression by increasing discharge during anti-saccade preparation, countering visuomotor bursts. This dual mechanism—prestimulus top-down gating and postsaccade override—ensures flexible behavioral control.^[19]^[20] The overall hierarchical model posits that cortical planning in the FEF, SEF, and LIP feeds descending signals to the SC for target selection and amplitude specification, which then converge on brainstem circuits for final motor burst generation. Feedback loops, including corollary discharge from the brainstem to the SC and cortex, allow real-time corrections for accuracy, as evidenced by adaptive adjustments in neuronal thresholds during error trials. This architecture balances speed and precision, with disruptions in any level altering saccade dynamics.^[14]^[21]

Perceptual Integration

Role in Visual Perception

Saccades play a central role in visual scanning by sequencing brief periods of fixation to construct a coherent representation of the visual environment. During natural viewing, humans typically execute three to four saccades per second, each redirecting gaze to points of interest and allowing the accumulation of detailed information across successive fixations.^[22]^[23] A key benefit of saccades lies in their ability to shift high-resolution foveal vision toward selected targets, thereby enhancing visual acuity beyond the coarse resolution provided by peripheral detection. This foveation process enables precise examination of objects or features initially detected in the low-acuity periphery, optimizing the use of the retina's central high-density photoreceptor region.^[24] Through saccades, the visual system integrates disparate fragments of information acquired during separate fixations into a stable and unified percept of the scene, compensating for the discontinuous nature of eye movements. This integration supports the perception of a continuous world despite the rapid shifts in retinal input that occur with each saccade.^[25] Saccades frequently align with shifts in covert attention, where attentional focus precedes and guides the eye movement to improve target detection and processing efficiency. This linkage, rooted in shared neural mechanisms, ensures that saccades are directed toward attended locations, enhancing overall perceptual selectivity.^[26] Recent studies from the 2020s highlight how saccades facilitate perception in dynamic environments, such as driving, by dynamically prioritizing salient features like moving vehicles or road hazards through modulated saccadic patterns.^[27]

Saccadic Masking

Saccadic masking, also known as saccadic suppression, refers to the temporary reduction in visual sensitivity that occurs during the execution of saccadic eye movements, effectively preventing the perception of motion blur resulting from the high-velocity sweep of images across the retina. This phenomenon ensures perceptual stability by inhibiting the processing of perisaccadic visual input, which would otherwise produce a disorienting smear due to retinal slip speeds exceeding 500 degrees per second in larger saccades. The suppression begins approximately 50-100 ms before saccade onset and persists through the movement, aligning closely with typical saccade durations of 30-120 ms.^[28]^[29]^[30] The underlying mechanism involves neural inhibition of visual processing pathways, primarily through corollary discharge signals from oculomotor centers that modulate activity in the visual cortex, reducing neuronal firing rates and contrast sensitivity by 50-80% in regions affected by the eye movement. This creates a transient functional scotoma—a blind spot—centered on the saccadic trajectory, where sensitivity to luminance, motion, and other stimuli is markedly diminished. Suppression occurs at multiple levels, starting in the retina with reduced ganglion cell responses to sequential stimuli and extending to cortical areas like V1 and V4, where inhibitory interneurons and feedback from higher centers amplify the effect.^[24]^[31]^[32]^[33] The adaptive purpose is to maintain a coherent visual world despite constant refixations, avoiding conflicts between pre- and post-saccadic scenes.^[34]^[35] Classic experimental evidence demonstrates elevated detection thresholds for brief visual flashes presented during saccades, with sensitivity dropping to as low as 10-20% of baseline levels, as thresholds can rise by 6-10 times compared to fixation conditions. More recent investigations using electrophysiological recordings have tied this suppression to enhanced oscillatory activity in the visual cortex; for instance, 2023 studies show increased alpha-band (7-13 Hz) power in V4 during saccades, which correlates with reduced visual responsiveness and contributes to the inhibitory gating.^[36]^[37]^[38]^[39]^[40]

Trans-saccadic Perception

Trans-saccadic perception encompasses the neural processes that preserve visual continuity across saccadic eye movements, ensuring the world appears stable despite the eyes' rapid shifts. During natural viewing, humans make approximately 3 saccades per second, displacing the retinal image by several degrees each time, yet compensatory mechanisms like corollary discharge signals from oculomotor commands counteract this instability by updating visual representations in advance. These signals, originating from pathways involving the superior colliculus and frontal eye fields, shift neuronal receptive fields to maintain a consistent perceptual map.^[41]^[42] A core component is the integration of visual features, where pre-saccadic information—such as object identity and color—is briefly stored in a limited trans-saccadic memory buffer with a capacity of about 3–4 items, akin to visual working memory. This stored information is then matched and fused with post-saccadic input through efference-copy-based remapping in cortical regions like the parietal and frontal eye fields, allowing synthesis of features from ventral and dorsal streams. Such integration relies on egocentric saccade metrics to align features spatially, preventing perceptual fragmentation.^[43]^[44] Perceptual continuity is further aided by a subjective compression of time during saccades, where visual intervals around saccade onset are underestimated by up to 50%—for instance, a 100 ms gap perceived as roughly 50 ms—peaking at saccade initiation and spanning a 300 ms window. This temporal distortion, specific to visual stimuli and independent of saccade amplitude, minimizes disruptions in the flow of perception without inverting order entirely. Despite these mechanisms, trans-saccadic memory exhibits clear limits, with poor retention of fine details like precise object positions; fidelity declines sharply as memory load increases from 1 to 4 items, evidenced by rising response variability (from ~22° to ~37°) and bias toward post-saccadic cues. The system compensates via heuristics, such as prioritizing recent sensory input or attentional cues to allocate limited resources efficiently, rather than maintaining high-resolution storage across all features.^[45]^[46] Recent advances highlight dynamic influences on these processes: a 2025 study demonstrated that short-term priors (from immediate prior stimuli) induce behavioral oscillations in orientation judgments during saccades, at ~9–10 Hz and synchronized to saccade onset, with bias amplitudes up to 1°. These oscillations, absent for long-term priors, align with predictive coding frameworks, where alpha-range neural rhythms facilitate Bayesian integration of pre- and post-saccadic signals for enhanced stability.^[47]

Spatial Updating

Spatial updating during saccades involves the brain's predictive remapping of visual receptive fields to compensate for the impending shift in gaze. Neurons in the parietal cortex, particularly the lateral intraparietal area (LIP), and frontal eye fields adjust their receptive fields prior to saccade onset, shifting them by the vector of the planned eye movement. This remapping ensures that neural representations of space remain stable despite the retinal displacement caused by the saccade. A key mechanism underlying this process is the corollary discharge, or efference copy, which originates from oculomotor commands in the brainstem and midbrain. This internal signal is transmitted via pathways such as the projection from the superior colliculus to the frontal eye fields and then to parietal areas, informing sensory regions about the expected change in eye position. The corollary discharge allows for anticipatory adjustments in visual processing, preventing perceptual disruptions from self-generated eye movements. The primary purpose of spatial updating is to maintain coherent spatial representations across gaze shifts, supporting behaviors like selective attention and visuomotor actions such as grasping objects. For instance, it enables the tracking of targets during sequential saccades, ensuring that attentional resources and motor plans align with the updated world coordinates. In parietal regions, this integration facilitates the remapping of attentional maps, minimizing disruptions to ongoing tasks. Evidence from single-cell recordings in monkeys demonstrates predictive remapping in LIP neurons, where responses to stimuli shift to future receptive fields before the saccade executes. Human functional magnetic resonance imaging (fMRI) studies confirm similar processes in the posterior parietal cortex, showing gaze-centered updating during double-step saccade tasks.^[48]^[49]^[50] Errors in spatial updating can lead to perceptual mislocalization, particularly when multiple objects are present, as the system struggles to remap all locations accurately. Such inaccuracies manifest as systematic shifts in perceived positions, especially in complex scenes with competing stimuli, highlighting the limits of predictive mechanisms under high cognitive load. Computational models suggest that these errors arise from incomplete integration of remapping signals across neural populations.^[51]

Applications

In Reading

During reading, the eyes make rapid saccadic movements interspersed with brief fixations, allowing the extraction of visual information from text. Forward saccades typically span 7-9 character spaces, advancing the gaze to the next word or within a word, while fixations last 200-250 milliseconds, during which most linguistic processing occurs.^[52] Regressions, which are backward saccades comprising 10-15% of all eye movements, return the gaze to previously fixated text to resolve comprehension difficulties or integrate information.^[53] The perceptual span—the region from which useful information is acquired during a fixation—is asymmetrical in left-to-right languages, extending approximately 7-8 characters to the right of the fixation point but only 3-4 characters to the left.^[54] This rightward bias facilitates previewing upcoming words, aiding in word identification and sentence prediction without disrupting the forward flow of reading.^[55] Saccade length and the frequency of regressions are influenced by linguistic properties of the text. Longer words elicit shorter forward saccades and more fixations within them, while high-frequency and predictable words promote longer saccades and fewer regressions by enabling efficient parafoveal processing.^[56]^[57] Low predictability, in contrast, increases regression rates as readers seek clarification from earlier material.^[58] Developmentally, children's reading eye movements differ markedly from those of adults. Young readers produce more regressions—often exceeding 20% of saccades—and shorter forward saccades due to immature linguistic and oculomotor control, leading to less efficient text processing.^[59] With reading skill acquisition, individuals make fewer regressions and longer, more targeted saccades, optimizing the balance between information uptake and cognitive load.^[60] Eye-tracking research has linked saccade efficiency to dyslexia, where affected individuals exhibit prolonged fixations, shorter saccades, and higher regression rates, reflecting disrupted text integration.^[61]

In Neurological Assessment

Saccades serve as valuable biomarkers in neurological assessment, where deviations from normal parameters—such as latencies typically ranging from 150-250 ms, peak velocities up to 500 degrees per second, and absence of intrusions—can signal underlying dysfunction in key brain regions.^[62] Abnormal saccadic latencies, velocities, or the presence of intrusions often indicate cerebellar involvement, as seen in slow or hypometric saccades due to impaired coordination; basal ganglia disorders, which may prolong latencies through disrupted initiation; or cortical dysfunction, leading to inaccurate targeting or inhibitory errors.^[63]^[64] These metrics, measured via eye-tracking, enable clinicians to localize lesions and monitor progression non-invasively.^[65] In Parkinson's disease (PD), saccades reveal characteristic impairments that aid early diagnosis and treatment evaluation. Patients commonly exhibit prolonged saccadic latencies and hypometric saccades, where eye movements undershoot targets, reflecting dopaminergic deficits in the basal ganglia.^[66] Eye-tracking studies from 2022-2025 have demonstrated that pro-saccade deficits, including increased latency and reduced velocity, can aid in early PD detection.^[67] For amyotrophic lateral sclerosis (ALS), saccadic intrusions—unintended small eye movements during fixation—emerge as a reliable progression biomarker. A 2024 longitudinal study of 28 ALS patients found that intrusion frequency and amplitude increased over 12 months, correlating with declines in the ALS Functional Rating Scale-Revised (ALSFRS-R) bulbar subscale (r ≈ -0.45), highlighting their utility in tracking bulbar-onset disease.^[68] This non-invasive measure outperforms subjective scales for early detection of upper motor neuron involvement, as intrusions reflect progressive loss of oculomotor control.^[69] Beyond neurodegenerative conditions, anti-saccade tasks, which require suppressing reflexive gazes to look away from a stimulus, quantify executive dysfunction in psychiatric disorders. In attention-deficit/hyperactivity disorder (ADHD), children show elevated anti-saccade error rates, indicating impaired inhibitory control, as validated by 2024 eye-tracking protocols using portable devices for screening.^[70] Similarly, in schizophrenia, anti-saccade error rates are elevated in patients, serving as a biomarker of prefrontal cortex deficits.^[71] Automated analysis algorithms, incorporating machine learning on eye-tracking data, enable non-invasive assessment of reading issues in children.^[72] Video-oculography (VOG) remains the gold standard for quantitative saccade assessment, using infrared cameras to capture high-resolution metrics like latency and velocity with sub-degree precision.^[73] Emerging devices, such as the 2025 EYE ROLL system, facilitate targeted saccadic training by delivering controlled visual stimuli, improving symmetry and speed.^[74]

Abnormalities and Adaptation

Pathophysiologic Saccades

Pathophysiologic saccades refer to abnormal eye movements arising from disruptions in the neural pathways controlling saccadic function, often manifesting in various neurological disorders. These abnormalities can include slowed velocities, hypometria (undershooting the target), intrusions (unwanted saccades interrupting fixation), or impaired conjugacy (coordinated movement of both eyes), reflecting underlying pathology in brainstem, cerebellar, or cortical structures. Unlike normal saccades, which are rapid and precise, pathophysiologic variants impair visual stability and contribute to symptoms like oscillopsia or gaze instability. In ocular motor disorders, cerebellar ataxia is associated with nystagmus-like saccadic intrusions, where involuntary saccades mimic the fast phases of nystagmus, disrupting steady fixation due to cerebellar dysfunction in modulating saccade accuracy. ^[75] Progressive supranuclear palsy (PSP), a tauopathy affecting midbrain structures, characteristically features slow saccades, particularly vertical ones, with peak velocities significantly reduced compared to normals, stemming from degeneration of the rostral interstitial nucleus of the medial longitudinal fasciculus. ^[76] Neurodegenerative conditions further exemplify saccade pathologies. In Parkinson's disease (PD), hypometric saccades often require multiple corrective steps to reach the target, a pattern linked to basal ganglia dopamine depletion and common in PD patients, contrasting with the single-step saccades in healthy individuals. ^[77] Multiple system atrophy (MSA), involving olivopontocerebellar atrophy, presents with frequent square-wave jerks—small horizontal saccadic intrusions (1-5 degrees) that interrupt fixation every few seconds—correlating with cerebellar involvement and observed in 64% of MSA patients versus 15% in PD, helping to distinguish MSA from PD. ^[78] Vascular and traumatic insults commonly produce gaze palsies that abolish or impair saccades. Post-stroke lesions in the pontine paramedian reticular formation or frontal eye fields result in conjugate gaze palsies, preventing ipsilesional saccades and leading to a persistent deviation of eyes to the contralesional side, as seen in 20-30% of hemispheric strokes. ^[79] Internuclear ophthalmoplegia (INO), often from demyelination or ischemia in the medial longitudinal fasciculus, disrupts conjugate horizontal saccades, causing adduction failure in the ipsilesional eye with abducting nystagmus in the fellow eye. ^[80] Recent longitudinal studies from 2023-2025 highlight saccade metrics as biomarkers for early disease detection and progression. In PD, reductions in saccade velocity—particularly prosaccades dropping below 300 degrees/second—emerge as an early indicator, detectable up to two years before motor symptoms and outperforming traditional dopamine imaging in sensitivity. ^[81] For amyotrophic lateral sclerosis (ALS), increasing saccadic intrusions, such as square-wave jerks, track bulbar progression over 12-24 months, correlating with ALSFRS-R scores (r=0.65) and offering a noninvasive marker independent of respiratory decline. ^[82] Genetic conditions also yield distinct saccade deficits. Congenital nystagmus, arising from mutations in FRMD7 or other genes, features impaired saccadic inhibition, with quick phases showing altered latencies (around 80-140 ms) and reduced amplitudes, perpetuating the oscillatory cycle from infancy. ^[83] In Niemann-Pick disease type C, a lysosomal storage disorder, patients exhibit slowed vertical saccades and hypometric horizontal ones, reflecting cholesterol accumulation in brainstem nuclei and correlating with cognitive severity across age groups. ^[84]

Saccade Adaptation

Saccade adaptation refers to the brain's capacity to modify the amplitude and direction of saccades through experience-dependent plasticity, ensuring precise gaze shifts despite changes in the oculomotor system. This process maintains saccadic accuracy, typically landing within 0.5–1 degree of the target in healthy individuals, by adjusting motor commands based on post-saccadic visual errors. The primary mechanism involves error signals generated from retinal-target misalignment at saccade endpoint, which drive gain adjustments—scaling the saccade amplitude up or down—primarily through cerebellar climbing fibers. These fibers convey sensory error information from the inferior olive to Purkinje cells in the cerebellar cortex, triggering long-term depression or potentiation of synaptic weights to refine the saccadic command.^[85]^[86] Adaptation manifests in two main types: outward adaptation, which increases saccade amplitude when the target steps away from the fovea during the saccade, and inward adaptation, which decreases amplitude when the target steps closer. These are commonly induced using the double-step paradigm, where the target initially appears at one location and then jumps to a second position either during or immediately after the saccade onset, allowing selective modification of primary saccade metrics without altering secondary corrective movements.^[87]^[88] The neural basis centers on the oculomotor vermis of the posterior cerebellum (lobules VI-VII) and interconnected brainstem nuclei, such as the paramedian pontine reticular formation, where error-driven signals modulate burst neuron activity. Studies in primates show that lesions or inactivation of the cerebellar vermis abolish adaptation, while electrical stimulation can induce it, with healthy adaptation achieving 70–80% correction of imposed errors over repeated trials in a single session.^[89]^[90] Functionally, saccade adaptation compensates for transient perturbations like muscle fatigue during prolonged gaze shifts or prism-induced visual displacements, restoring accuracy without conscious effort. This plasticity is incomplete in cerebellar disorders, highlighting its reliance on intact vermal circuits for ongoing calibration. Recent research in the 2020s has explored virtual reality (VR)-based protocols for enhancing saccade adaptation in rehabilitation, demonstrating improved oculomotor plasticity in patients with neurological impairments through immersive error-feedback training. These approaches, including integration with sports vision devices for dynamic target tracking, show promise for targeted therapy, with studies from 2020-2022 reporting improved adaptation rates compared to traditional methods.^[91]^[92]

Comparative Physiology

In Non-Human Primates

Saccades in non-human primates, particularly rhesus macaques, exhibit kinematic properties closely resembling those in humans, establishing them as premier model organisms for oculomotor research. Visually guided saccades in macaques typically have latencies ranging from 150 to 250 ms, aligning with human values and reflecting shared visuomotor processing timelines.^[93] They adhere to the main sequence—a nonlinear relationship between amplitude and velocity—observed across primate species.^[94] These similarities extend to neural circuits, where the frontal eye fields (FEF) and lateral intraparietal area (LIP) encode saccade-related activity, as demonstrated by single-unit recordings in behaving monkeys that mirror human functional imaging findings.^[95] Invasive electrophysiological techniques, feasible in non-human primates due to ethical and technical advantages over human studies, have elucidated key mechanisms underlying saccade generation. Recordings from the superior colliculus reveal burst neurons that discharge at high frequencies during saccades, providing the motor command signals absent in non-invasive human data.^[96] Additionally, laboratory training enables the elicitation of express saccades in macaques, with latencies as short as 100-120 ms, which depend on target luminance and size and offer a window into preparatory neural states not easily studied in humans.^[97] Subtle differences in saccade characteristics exist between humans and non-human primates. Primate models also display more robust adaptation to oculomotor lesions; for example, following dorsal cerebellar vermis ablation, macaques recover saccade accuracy through compensatory mechanisms faster than observed in human lesion cases.^[98] Non-human primate research has been pivotal in validating human trans-saccadic perception, confirming that mechanisms like perisaccadic suppression operate similarly across species.^[99] In the 2020s, optogenetic tools have advanced causal understanding, such as by selectively activating corticotectal pathways in macaques to dissect their role in saccade targeting and perisaccadic visual stability.^[100]

In Other Animals

In birds, eye movements are typically limited in range, often to about 20 degrees, with the primary role of stabilizing gaze during larger head saccades rather than independent ocular shifts. This configuration arises from the relatively fixed position of the eyes in the skull, necessitating coordinated head movements to redirect gaze toward targets. For instance, in chickens, saccadic eye movements occur predominantly during the thrust phases of head bobbing while walking, ensuring image stabilization across a wide field of view.^[101]^[102] In species like barn owls, which possess high visual acuity despite these constraints, microsaccades—small, involuntary flicks—contribute to maintaining sharp focus by counteracting drift and enhancing resolution during fixation, particularly in low-light hunting scenarios.^[103] Reptiles and amphibians exhibit saccades that are generally constrained in amplitude, with eye rotations limited to 3–6 degrees in frogs, complemented by broader head movements up to 30–40 degrees for overall gaze redirection. These movements facilitate prey detection but are slower and less frequent than in mammals, prioritizing stability over rapid scanning. In frogs, saccades play a critical role in ballistic tongue projection during prey capture, where eye and head positioning establish a shared coordinate system with the tongue trajectory, allowing precise extrapolation of moving targets' paths in milliseconds.^[104]^[105] Fish and invertebrates rely heavily on optokinetic saccades to stabilize retinal images during locomotion or environmental motion. In zebrafish, these saccades form the fast phase of the optokinetic response, resetting eye position to counteract slow-phase drifts induced by visual stimuli like rotating gratings, thereby maintaining a stable view of the surroundings.^[106] Invertebrates such as flies demonstrate foveation-like scanning through stereotyped head saccades that interrupt smooth optokinetic tracking, resetting gaze to high-resolution regions of the compound eye for targeted inspection, akin to foveal shifts in vertebrates. This strategy supports rapid flight navigation and prey pursuit in species like robber flies.^[107]^[108] Evolutionary trends in saccades reflect adaptations to ecological niches, with mammals developing quick, high-velocity foveal shifts to enable the "saccade-and-fixate" strategy for detailed visual sampling. In contrast, non-mammalian vertebrates emphasize reflexive stabilization over voluntary exploration. Recent comparative studies, such as those on predatory insects, link saccade speed and predictability to predation success; for example, faster predictive saccades in robber flies allow interception of evasive beetle prey by estimating wingbeat frequencies from visual cues.^[109]^[108] Functional divergence across phylogeny shows reduced voluntary control in lower vertebrates, where saccades are predominantly reflexive and often disconjugate between eyes, driven by sensory triggers rather than cognitive intent, unlike the integrated voluntary-reflexive systems in higher taxa.^[102]

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Oct 15, 2015 · Saccades are rapid eye movements designed to shift the fovea to objects of visual interest. Abnormalities of saccades offer important clues ...
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Saccades are rapid, ballistic movements of the eyes that abruptly change the point of fixation. They range in amplitude from the small movements made while ...
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Saccade - an overview | ScienceDirect Topics
Eye Movements; Saccades Saccades typically have a peak velocity of 30–700°/s and a duration of 30–100 ms for amplitude ranging from 0.5° to 40° (Figure 1(a)).
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Effort Drives Saccade Selection - eLife
Feb 28, 2025 · Humans make fast, ballistic eye movements, called saccades, to explore the rich visual world [1]. Saccades are executed approximately three to ...
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The Evolution of Gaze Shifting Eye Movements | Request PDF
This saccade-and-fixate strategy arose early in fish evolution, when the original function of saccades was to re-centre the eye as the fish turned.
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Neurophysiology and Neuroanatomy of Reflexive and Volitional ...
This review provides a summary of the contributions made by human functional neuroimaging studies to the understanding of neural correlates of saccadic control.
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Saccade Amplitude - an overview | ScienceDirect Topics
For saccades from 3 to 7 °, the system is constrained by a minimum duration of the agonist pulse; saccade magnitude is dependent on the number of active neurons ...
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The parallel programming of landing position in saccadic eye ...
... saccades using a 22 degree per second velocity and 8,000 degrees per second squared acceleration criteria. Further analysis of saccade dynamics and metrics ...
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The Spectral Main Sequence of Human Saccades
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[PDF] Does Saccadic Undershoot Minimize Saccadic Flight-time ... - CORE
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We review a new computational model developed to understand how evidence about stimulus salience in visual search is translated into a saccade command.Missing: paper | Show results with:paper
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Saccade execution increases the preview effect with faces: An EEG ...
Nov 2, 2023 · Under naturalistic viewing conditions, humans conduct about three to four saccadic eye movements per second. These dynamics imply that in ...
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A Computational Dual-Process Model of Fixation-Duration Control in ...
Sep 1, 2021 · In typical tasks, humans make three to four saccadic eye movements per second (Rayner, 2009). Whenever the eyes shift to a new location, the ...
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Fast and nonuniform dynamics of perisaccadic vision in the ... - PNAS
These rapid eye movements (saccades) enable high visual acuity by redirecting the tiny high-resolution region of the retina (the foveola). But in doing so ...
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Near-optimal integration of orientation information across saccades
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Is covert attention necessary for programming accurate saccades ...
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https://arxiv.org/pdf/2511.02689
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Oct 7, 2009 · This aspect of perceptual stability is often referred to as saccadic suppression: a reduction of visual sensitivity around the time of saccades.
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Saccades and Fixation - an overview | ScienceDirect Topics
... ms to take in visual detail, while a saccade is a rapid eye movement that repositions the eyes to a new location, occurring within 30-120 ms. AI generated ...
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Neural Dynamics of Saccadic Suppression - PMC - PubMed Central
This aspect of perceptual stability is often referred to as saccadic suppression: a reduction of visual sensitivity around the time of saccades. Here, we ...
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Age effects on saccadic suppression of luminance and color | JOV
Jun 30, 2021 · Compared to fixation, average perisaccadic contrast sensitivity decreased significantly by 66% for luminance and by 36% for color. A significant ...
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Suppression without inhibition: how retinal computation contributes ...
Jul 12, 2022 · We find that sequential stimuli, like those that naturally occur during saccades, trigger three independent suppressive mechanisms in the retina.
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Our results demonstrate neural correlates of saccadic suppression in a significant number of V4 neurons. The nature of this suppression changes dynamically ...
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A dramatic decline in visual sensitivity when saccades take place. This reduction in visual sensitivity during saccades is called saccadic suppression.Eye Movement Research · Saccades · Functions And...
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Saccadic suppression as a perceptual consequence of efficient ...
We suggest that saccadic suppression originates from efficient sensorimotor processing, indicating that the brain shares neural resources for perception and ...
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Saccadic suppression and stimulus uncertainty
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The role of neural oscillations in visuo-motor communication at the ...
Nov 5, 2023 · We show that saccades lead to suppression of visual sensitivity at saccadic onset, and that this suppression is accompanied by endogenous neural oscillations ...
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Laminar mechanisms of saccadic suppression in primate visual cortex
Saccadic eye movements are known to cause saccadic suppression, a temporary reduction in visual sensitivity and visual cortical firing rates.
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Suppression and reversal of motion perception around the time of ...
Saccadic motion suppression must be active because it starts before saccade onset, at a time when the motion probe appears completely outside the saccade. No ...Missing: VR sickness fMRI
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This review considers the substantial advances in understanding the neuronal mechanisms underlying this visual stability.
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Saccadic Corollary Discharge Underlies Stable Visual Perception
Jan 6, 2016 · We conclude that this corollary discharge provides a critical signal that can be used to unite jumping retinal images into a consistent visual scene.Materials And Methods · Results · Perception Changes During Md...
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Nov 8, 2014 · In this article, we examine accounts of how such phenomenal stability may be achieved. We then discuss findings on visual memory and integration ...
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May 21, 2021 · Our results suggest that transsaccadic integration of an object's features requires allocation of limited working memory resources to that ...
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Mar 17, 2025 · Here we investigate the role of oscillations in integrating pre-saccadic information with the current sensory signals.
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Predictive updating of attentional maps could minimize the impact of saccades on behavior by ensuring attention is directed towards appropriate environmental ...
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Jun 3, 2020 · We hypothesized that inferior parietal cortex (specifically supramarginal gyrus [SMG]) integrates saccade and visual signals to update grasp plans.
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A circuit model for transsaccadic space updating and mislocalization
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Fixation durations are typically about 180–250 ms. They are traditionally classified as first-pass or rereading fixations.
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Regressive Saccades and Word Perception in Adult Reading
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Sep 7, 2023 · The present study set out to test the joint influences of word properties and individual reading skills on eye movements during reading among Chinese children.
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Word Predictability Affects Saccade Length in Chinese Reading
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A dynamic adjustment model of saccade lengths in reading for word ...
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[PDF] Children's Development of Oculomotor Control during Reading
shorter saccades, and more regressions than adults during reading. These differences between children and adults in oculomotor behaviour during reading.
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Eye movements during reading in beginning and skilled readers
We discuss the differences found in eye movements during reading between children in different age groups and with different reading levels and skilled adult ...
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Improving Reading and Eye Movement Control in Readers with ...
Jun 23, 2025 · These findings suggest that guiding the eyes toward the PVL may offer a novel way to improve reading efficiency, particularly for individuals ...
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Saccadic eye movements in neurological disease - PubMed Central
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Jan 17, 2019 · Reduced saccade velocity, frequently called “slow saccades” are typically seen in a classic disorder of the midbrain called progressive supranuclear palsy.Background · Disorders Of Cerebellum... · Syndrome Of Anti-Gad...Missing: assessment utility
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A review of pursuit and saccadic eye movements and their utility in ...
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Eye Tracking in Parkinson's Disease: A Review of Oculomotor ...
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Changes in saccadic intrusions over time as an objective biomarker ...
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Changes in saccadic intrusions over time as an objective biomarker ...
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Auxiliary Diagnosis of Children With Attention-Deficit/Hyperactivity ...
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Diagnosis of schizophrenia by integrated saccade scores and ...
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Using Eye-Tracking to Assess Dyslexia: A Systematic Review of ...
The findings underscore the potential of eye-tracking to enhance diagnostic accuracy through metrics such as fixation counts, saccadic patterns, and processing ...
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Saccadic eye movements in neurological disease: cognitive ...
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Application of a New Device for Saccadic Training in Athletes
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This article provides an overview of nystagmus and saccadic intrusions with the goal of facilitating recognition and differentiation of abnormal eye movements.
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The frequency of square wave jerks (SWJ) was compared in eight patients with progressive supranuclear palsy (PSP), 25 patients with multiple system atrophy ...
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validation of an iPad-based eye movement assessment system ...
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Changes in saccadic intrusions over time as an objective biomarker ...
Jul 8, 2024 · Evaluation of saccadic intrusions during fixation was able to detect disease progression over time, correlated with ALSFRS-R bulbar subscale.
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Quick Phases of Infantile Nystagmus Show the Saccadic Inhibition ...
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A reverberation of past errors in the cerebellar climbing fiber signal
Characteristics of responses of Golgi cells and mossy fibers to eye saccades and saccadic adaptation recorded from the posterior vermis of the cerebellum.
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How cerebellar motor learning keeps saccades accurate - PMC - NIH
The neural elements that fashion the command signal for the generation of accurate saccades involve subcortical structures in the brain stem and cerebellum.The Necessity For The... · Neuronal Circuit Of Saccade... · The Neuronal Sites Of...<|control11|><|separator|>
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Monkeys were initially trained to sit and drink water in the primate chair. They received a liquid reward when their eye position entered the invisible computer ...
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Express-saccades of the monkey: reaction times versus intensity ...
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Effects of lesions of the oculomotor vermis on eye movements in ...
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Laminar mechanisms of saccadic suppression in primate visual cortex
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Retrograde Optogenetics Reveals Functional Convergence within a ...
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Oculomotor behaviour in vertebrates and invertebrates
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The Role of Motion Extrapolation in Amphibian Prey Capture
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The optokinetic response in zebrafish and its applications
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Mechanisms of punctuated vision in fly flight - ScienceDirect.com
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Predictive saccades and decision making in the beetle-predating ...
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In animals with good eyesight most eye movements consist of saccades, which rapidly shift the direction of the eye's axis, and intervals between the saccades.