Amplitude Of Microsaccades Research Articles

Foveolar Drusen Decrease Fixation Stability in Pre-Symptomatic AMD.

This study aims at linking subtle changes of fixational eye movements (FEM) in controls and in patients with foveal drusen using adaptive optics retinal imaging in order to find anatomo-functional markers for pre-symptomatic age-related macular degeneration (AMD). We recruited 7 young controls, 4 older controls, and 16 patients with presymptomatic AMD with foveal drusen from the Silversight Cohort. A high-speed research-grade adaptive optics flood illumination ophthalmoscope (AO-FIO) was used for monocular retinal tracking of fixational eye movements. The system allows for sub-arcminute resolution, and high-speed and distortion-free imaging of the foveal area. Foveal drusen position and size were documented using gaze-dependent imaging on a clinical-grade AO-FIO. FEM were measured with high precision (RMS-S2S = 0.0015degrees on human eyes) and small foveal drusen (median diameter = 60 µm) were detected with high contrast imaging. Microsaccade amplitude, drift diffusion coefficient, and ISOline area (ISOA) were significantly larger for patients with foveal drusen compared with controls. Among the drusen participants, microsaccade amplitude was correlated to drusen eccentricity from the center of the fovea. A novel high-speed high-precision retinal tracking technique allowed for the characterization of FEM at the microscopic level. Foveal drusen altered fixation stability, resulting in compensatory FEM changes. Particularly, drusen at the foveolar level seemed to have a stronger impact on microsaccade amplitudes and ISOA. The unexpected anatomo-functional link between small foveal drusen and fixation stability opens up a new perspective of detecting oculomotor signatures of eye diseases at the presymptomatic stage.

Temporal Changes in Fixational Eye Movements After Concussion in Adolescents and Adults: Preliminary Findings.

Concussions often involve ocular impairment and symptoms such as convergence insufficiency, accommodative insufficiency, blurred vision, diplopia, eye strain, and pain. Current clinical assessments of ocular function and symptoms rely on subjective symptom reporting and/or involve lengthy administration time. More objective, brief assessments of ocular function following concussion are warranted. The purpose of this study was to evaluate changes in fixational eye movements (FEMs) and their association with clinical outcomes including recovery time, symptoms, cognitive and vestibular/ocular motor impairment. Thirty-three athletes (13-27 years of age; 54.5% female) within 21 days of a diagnosed concussion participated in the study. A tracking scanning laser ophthalmoscope (TSLO) evaluated FEMs metrics during fixation on a center and corner target. Participants completed symptom (Post-Concussion Symptom Scale [PCSS]), cognitive (Immediate Post-concussion Assessment and Cognitive Testing [ImPACT], and Vestibular/Ocular Motor Screening (VOMS) evaluations. All measures were administered at the initial visit and following medical clearance, which was defined as clinical recovery. Changes in FEMs were calculated using paired-samples t tests. Linear regression (LR) models were used to evaluate the association of FEMs with clinical recovery. Pearson product-moment correlations were used to evaluate the associations among FEMs and clinical outcomes. On the center task, changes across time were supported for average microsaccade amplitude (p = 0.005; Cohen's d = 0.53), peak velocity of microsaccades (p = 0.01; d = 0.48), peak acceleration of microsaccades (p = 0.02; d = 0.48), duration of microsaccade (p < 0.001; d = 0.72), and drift vertical (p = 0.017; d = -0.154). The LR model for clinical recovery was significant (R2 = 0.37; p = 0.023) and retained average instantaneous drift amplitude (β = 0.547) and peak acceleration of microsaccade (β = 0.414). On the corner task, changes across time were supported for drift proportion (p = 0.03; d = 0.43). The LR model to predict clinical recovery was significant (R2 = 0.85; p = 0.004) and retained average amplitude of microsaccades (β = 2.66), peak velocity of microsaccades (β = -15.11), peak acceleration of microsaccades (β = 12.56), drift horizontal (β = 7.95), drift vertical (β = 1.29), drift amplitude (β = -8.34), drift proportion (β = 0.584), instantaneous drift direction (β = -0.26), and instantaneous drift amplitude (β = 0.819). FEMs metrics were also associated with reports of nausea and performance within the domain of visual memory. The FEMs metric were also associated with PCSS, ImPACT, and VOMS clinical concussion outcomes, with the highest magnitude correlations between average saccade amplitude and VOMS symptoms of nausea and average instantaneous drift speed and ImPACT visual memory, respectively. FEMs metrics changed across time following concussion, were useful in predicting clinical recovery, and were correlated with clinical outcomes. FEMs measurements may provide objective data to augment clinical assessments and inform prognosis following this injury.

Poster Session: Fixational Eye Motion Measured with Tracking Scanning Laser Ophthalmoscopy in ABI/TBI and Control Subjects.

This study compared fixational eye motion (FEM) in acquired/traumatic brain injury (ABI/TBI) subjects presenting with clinical symptoms (reading text, losing place while reading, words moving around the page, decreased reading comprehension) to control subjects as measured with Tracking Scanning Laser Ophthalmoscopy (TSLO). Six ABI/TBI subjects (4M, 2F) aged 29-62 and six control subjects (4M, 2F) aged 23-42 participated in the study. A 5°x5° square 840nm TSLO imaging raster was used to guide fixation in 3 conditions: upper right corner (UR), freely in the center (CF), and a dot in the center (DOT). At least three 10sec videos were collected for all subjects in both eyes for each condition. Offline, custom MATLAB software was used to stabilize the videos, extract eye motion at 480Hz, and extract FEM characteristics. Significant differences were seen between the ABI/TBI and control subjects with t-test analysis in all fixation conditions. Different Microsaccade and drift parameters of FEM were found to be significantly different for the different conditions with the number of Microsaccade reaching significance only for UR (p<0.001) and Microsaccade amplitude and peak velocity only for CF (p<0.01). These results demonstrate FEM measurements with TSLO in ABI/TBI subjects are different from control and are highly variable between fixation tasks. This warrants further exploration to determine the mechanisms behind oculomotor deficiency and to guide future clinical treatments.

V1-bypassing suppression leads to direction-specific microsaccade modulation in visual coding and perception

Microsaccades play a critical role in refreshing visual information and have been shown to have direction-specific influences on human perception. However, the neural mechanisms underlying such direction-specific effects remains unknown. Here, we report the emergence of direction-specific microsaccade modulation in the middle layer of V2 but not in V1: responses of V2 neurons after microsaccades moved toward their receptive fields were stronger than those when microsaccades moved away. The decreased responses from V1 to V2, which are correlated with the amplitude of microsaccades away from receptive fields, suggest topographically location-specific suppression from an oculomotor source. Consistent with directional effects in V2, microsaccades function as a guide for monkeys’ behavior in a peripheral detection task; both can be explained by a dynamic neural network. Our findings suggest a V1-bypassing suppressive circuit for direction-specific microsaccade modulation in V2 and its functional influence on visual sensitivity, which highlights the optimal sampling nature of microsaccades.

The dynamics of microsaccade amplitude reflect shifting of covert attention

Attention flexibly shifts between spatial locations to accommodate task demands. The present study examined if the dynamics of attentional shifting are seen in microsaccades whose direction has been shown to accompany the shifts of covert attention. In a spatial cueing task, the cue predicted the target location on 100%, 75%, or 50% of the trials. The results revealed that microsaccade rate and amplitude were both reduced following cue onset and then rebounded. Both microsaccade rate and amplitude were biased towards the opposite direction of the cue and then returned to the cued direction. Importantly, the cue validity modulated the temporal profile of microsaccade amplitude but had little impact on the temporal profile of microsaccade rate. In line with this, the cueing effect measured with target response accuracy was correlated with the microsaccade amplitude only. These results indicate that the temporal dynamics of microsaccade amplitude reflect shifting of covert attention.

Reduced fixation stability induced by peripheral viewing does not contribute to crowding.

Attending to peripheral visual targets while maintaining central fixation, a process that involves covert attention, reduces fixation stability. Here, we tested the hypothesis that changes in fixation stability induced by peripheral viewing contribute to crowding in peripheral vision by increasing positional uncertainty. We first assessed whether fixation was less stable during peripheral versus central (foveal) viewing for both crowded and uncrowded stimuli. We then tested whether fixation stability during peripheral viewing was associated with the extent of crowding. Fourteen participants performed a tumbling E orientation discrimination task at three different eccentricities (0°, 5°, 10°). The target was presented with or without flankers. Fixational eye movements were measured using an infrared video-based eyetracker. A central fixation cross was provided for the two peripheral viewing conditions, and optotype size was scaled for each eccentricity. Discrimination of appropriately scaled uncrowded stimuli was unaffected by eccentricity, whereas discrimination of crowded stimuli deteriorated dramatically with eccentricity, despite scaling. Both crowded and uncrowded peripheral stimuli were associated with reduced fixation stability, increased microsaccadic amplitude, and a greater proportion of horizontal microsaccades relative to centrally presented stimuli. However, these effects were not associated with the magnitude of crowding. This suggests that reduced fixation stability due to peripheral viewing does not contribute to crowding in peripheral vision.

Microsaccades: Empirical Research and Methodological Advances - Introduction to Part 1 of the Thematic Special Issue.

Microsaccades: Empirical Research and Methodological Advances - Introduction to Part 1 of the Thematic Special Issue.

Object detection based on saturation of visual perception

This paper presents a framework to detect salient objects from natural images through simulating human perception. We think perception saturation, generated by microsaccades in gaze, is the primary reason why human brains export consciousness of salient objects. Perception saturation is represented by using amplitude of microsaccades (AOM). When the AOM or changes of the AOM tend to zero, human perception becomes saturated. Motivated by this analysis, we construct a group of learning-based models to detect a salient object in a coarse-to-fine sequence. Firstly, a small image is selected to minimize the AOM so as to saturate the perception. Then, neural networks with random weights (NNRW) are chosen to simulate the received visual stimuli. In order to examine the changes of AOM, a positive feedback loop is constructed which executes the procedure of “pixel sampling-learning classification” iteratively. The final fixation area after iterations is regarded as a salient object. The proposed algorithm is based on unsupervised learning and data-driven completely. Our results based on open image datasets show that the proposed method achieves better performance compared to those existing unsupervised algorithms.

Effects of visual blur on microsaccades during visual exploration.

Microsaccades shift the image on the fovea and counteract visual fading. They also serve as an optimal sampling strategy while viewing complex visual scenes. Microsaccade production relies on the amount of retinal error or acuity demand of a visual task. The goal of this study was to assess the effects of blur induced by uncorrected refractive error on visual search. Eye movements were recorded in fourteen healthy subjects with uncorrected and corrected refractive error while they performed a) visual fixation b) blankscene viewing c) visual search (spot the difference) tasks. Microsaccades, saccades, correctly identified differences and reaction times were analyzed. The frequency of microsaccades and correctly identified differences were lower in the uncorrected refractive error during visual search. No similar change in microsaccades was seen during blank-scene viewing and gaze holding tasks. These findings suggest that visual blur, hence the precision of an image on the fovea, has an important role in calibrating the amplitude of microsaccades during visual scanning.

Changes in Fixational Eye Movements following Concussion

Concussions can affect smooth pursuit, saccadic, and vergence eye movements. Fixational eye movements (FEMs) - the small, involuntary motion of the eye that occurs while focusing on a target - are affected by Alzheimer’s, Parkinson’s, and mild cognitive impairment. However, little is known about changes in FEMs following concussion. PURPOSE: To compare FEMs in concussed patients to controls using a retinal image-based eye tracking device. METHODS: Participants included 50 patients with a concussion and 39 age- and gender-matched controls aged 13-27 years. FEMs were measured with a tracking scanning laser ophthalmoscope (TSLO) that tracks retinal image motion at 480 Hz with accuracy of ~0.2 arcmin. Eye traces were analyzed offline to compute microsaccadic amplitude, peak velocity, and peak acceleration. Fixational spread, using bivariate contour ellipse area (BCEA), intersaccadic intervals, blink rate, and total blink time were also analyzed. Concussed patients completed the Vestibular Oculomotor Screening (VOMS), Post-concussion Symptom Scale (PCSS), and Immediate Post-Concussion Assessment and Cognitive Test (ImPACT) for comparison with FEMs. RESULTS: Microsaccades were larger (amplitude – controls: 0.397° SD 0.32, concussion: 0.597° SD: 0.45; p <0.001) and faster (peak velocity: control: 27.9°/sec SD: 22.2; concussion: 39.7°/sec SD: 30.3; p<0.001; peak acceleration: control: 6.27°/sec2 SD: 9.29; concussion: 9.47°/sec2 SD: 14.1) in concussed participants. Although concussed patients and controls made equal numbers of microsaccades during a 30 sec recording, concussion patients had a greater proportion of larger, faster microsaccades - with 19% more microsaccades at amplitudes greater than 0.75° and 22% more microsaccades ≥30°/sec. The BCEA was 221% larger in concussed patients (0.56°) compared to controls (0.26°). CONCLUSIONS: These findings support changes in FEMs following concussion as measured using retinal image-based eye tracking. Microsaccades in concussed patients were larger in amplitude, peak velocity, and peak acceleration compared to controls. Specifically, fixation is less precise, with a larger spread (i.e., increased BCEA). Retinal imaging and eye-tracking of FEMs may be useful in identifying and monitoring recovery following concussion.

Reduced amblyopic eye fixation stability cannot be simulated using retinal-defocus-induced reductions in visual acuity

Amblyopia is associated with impaired visual acuity (VA) and reduced fixation stability (FS). To assess whether impaired VA may cause reduced FS, the effects of retinal-defocus-induced visual acuity reductions on FS were measured in observers with amblyopia and controls. Fixational eye movements were measured in 8 patients with amblyopia and 12 controls. Monocular near VA of a subset of controls (n = 5) was then varied from 20/20 to 20/100 using convex lenses. The amblyopia group completed three monocular conditions; 1) amblyopic eye fixating, 2) fellow eye fixating and 3) fellow eye fixating with VA reduced to match the amblyopic eye. Fixational eye movements were quantified using bivariate contour ellipse area (BCEA) and microsaccadic amplitude. Amblyopic eye BCEA was significantly larger with increased microsaccadic amplitude compared to the fellow eye and control eyes. BCEA and VA were positively correlated for amblyopic eyes. VA impairments induced by retinal defocus did not reduce FS in controls or the fellow eye of observers with amblyopia, even when fellow eye VA was matched to that of the amblyopic eye. This suggests that reduced FS in amblyopic eyes cannot be simulated by acute VA reductions. Therefore, reduced amblyopic eye FS may not be a direct consequence of the VA loss alone. As in previous studies, a correlation between BCEA and VA for amblyopic eyes was observed. This relationship could be due to a third, mediating variable or an effect of fixational eye movements on VA.

Dynamic modulation of the perceptual load on microsaccades during a selective spatial attention task

Selective spatial attention enhances task performance at restricted regions within the visual field. The magnitude of this effect depends on the level of attentional load, which determines the efficiency of distractor rejection. Mechanisms of attentional load include perceptual selection and/or cognitive control involving working memory. Recent studies have provided evidence that microsaccades are influenced by spatial attention. Therefore, microsaccade activities may be exploited to help understand the dynamic control of selective attention under different load levels. However, previous reports in humans on the effect of attentional load on microsaccades are inconsistent, and it is not clear to what extent these results and the dynamic changes of microsaccade activities are similar in monkeys. We trained monkeys to perform a color detection task in which the perceptual load was manipulated by task difficulty with limited involvement of working memory. Our results indicate that during the task with high perceptual load, the rate and amplitude of microsaccades immediately before the target color change were significantly suppressed. We also found that the occurrence of microsaccades before the monkeys’ detection response deteriorated their performance, especially in the hard task. We propose that the activity of microsaccades might be an efficacious indicator of the perceptual load.

The proportion of microsaccadic overshoot and the influence of accomodation on the quantitative measures of microsaccades in fourteen normal test persons

PurposeIn normal test persons to (1) quantify the proportion of microsacadic overshoot, and (2) to investigate if accomodation have an influence on the quantitative measure of microsaccades.MethodsFourteen healthy adults (mean age 31.2 years, range 16–50 years) all with a visual acuity of 1.0 or better with no previous eye diseases were investigated. Mean ametropia was −1.0 diopter. Microsaccades were quantified binocularly using an infrared recording technique (Eye Link 1000) during fixation at a distance of 40 cm and 6 m. Accomodation was measured at 40 cm and 6 m using an open field autorefractor (Shin Nippon SRW‐5000).ResultsThe mean proportion of microsaccadic overshoot during fixation at 40 cm and 6 m was 75.8% and 146.1% larger than the normal “classic” microsaccadic amplitude, respectively. The relation between the “classic” microsaccadic amplitude (x) and the proportion of overshoot (f) could be fitted to a negative exponential curve. The best curve fit at 40 cm (R2 = 0.88) was f = 3.02*exp(−6.00*x) and at 6 m f = 3.10*exp(−6.21*x) (R2=0.98). The mean “classic” microsaccadic amplitude (±SD) at 40 cm was 0.334 ± 0.140 and 0.256 ± 0.073° at 6 m. The microsaccadic amplitude ± SD including overshoot at 40 cm and 6 m was 0.486 ± 0.209 and 0.411 ± 0.101°, respectively. Accomodation (in diopters) at 40 centimeters was positively correlated to the mean “classic” microsaccadic amplitude.ConclusionsThe proportion of microsaccadic overshoot is larger for smaller microsaccades when fixating at both near and far targets. Microsaccades seems to be influenced by accomodation.

Fixational eye movements in Tourette syndrome.

Studies of saccadic eye movements in subjects with Tourette syndrome (TS) have provided additional evidence that there is a link between TS symptoms and deficits in fronto-striato-thalamic networks. These studies revealed impaired timing and inhibition of saccades. We compared fixational eye movements, such as microsaccades and ocular drifts, in subjects with TS and healthy controls.We measured horizontal and vertical eye positions with video-oculography in 14 subjects with Tourette syndrome. We found reduced microsaccade amplitude but increased time between adjacent microsaccades (intersaccadic interval). Hence, the rate of microsaccades was reduced in subjects with TS compared to controls. Measure of ocular stability during intersaccadic intervals revealed increased drift velocity and increased variance in eye position. We hypothesize that increased activity of the direct fronto-striatal pathway and the resulting reduction in basal ganglia outflow targeting the superior colliculus fixation zone affect the rate and amplitude of microsaccades in subjects with TS. The resulting impairment in frontal eye field fixation leads to increased drifts during intersaccadic interval in subjects with TS. Possible clinical implication for these results is that fixational eye movements can be objective biological markers of TS.

Microsaccade dysfunction and adaptation in hemianopia after stroke

Besides the reduction of visual field size, hemianopic patients may also experience other poorly understood symptoms such as blurred vision, diplopia, or reduced visual acuity, which may be related to microsaccade function. To determine (i) if microsaccades are altered in hemianopia; (ii) how altered microsaccade features correlate with visual performances; and (iii) how their direction relates to visual field defect topography. In this case-control study, microsaccades of hemianopic stroke patients (n = 14) were assessed with high-resolution eye-tracking technique, compared with those of healthy controls (n = 14), and correlated with visual performances, visual field defect parameters and lesion age. Patients' microsaccades had (i) larger amplitude (P = 0.027), (ii) longer duration (P = 0.042), and (iii) impaired binocular microsaccade conjugacy (horizontal: P = 0.002; vertical: P = 0.035). Older lesions were associated with poorer binocular conjugacy (horizontal: r(14) = 0.67, P = 0.009; vertical: r(14) = 0.75, P = 0.002) and larger microsaccade amplitudes (r(14) = 0.55, P = 0.043). (iv) Half of the patients had a microsaccade bias towards the seeing field (monocular: P = 0.002; binocular: P < 0.001) which was associated with faster reactions to super-threshold visual stimuli in areas of residual vision (P = 0.042). Finally, (v) patients with more binocular microsaccades (r(14) = 0.59, P = 0.027) and lower microsaccade velocity (r(14) = -0.66, P = 0.011) had better visual acuity. Hemianopia leads not only to the loss of visual field but also to microsaccade enlargement and impaired binocular conjugacy, suggesting malfunctioning microsaccade control circuits which worsen over time. But a microsaccade bias towards the seeing field, which suggests greater allocation of attention, accelerates stimulus detection. Microsaccades may play a role to compensate for vision impairment and provide a basis for vision restoration and plasticity, which deserves further exploration.

Alteration of the microsaccadic velocity-amplitude main sequence relationship after visual transients: implications for models of saccade control.

Microsaccades occur during gaze fixation to correct for miniscule foveal motor errors. The mechanisms governing such fine oculomotor control are still not fully understood. In this study, we explored microsaccade control by analyzing the impacts of transient visual stimuli on these movements' kinematics. We found that such kinematics can be altered in systematic ways depending on the timing and spatial geometry of visual transients relative to the movement goals. In two male rhesus macaques, we presented peripheral or foveal visual transients during an otherwise stable period of fixation. Such transients resulted in well-known reductions in microsaccade frequency, and our goal was to investigate whether microsaccade kinematics would additionally be altered. We found that both microsaccade timing and amplitude were modulated by the visual transients, and in predictable manners by these transients' timing and geometry. Interestingly, modulations in the peak velocity of the same movements were not proportional to the observed amplitude modulations, suggesting a violation of the well-known "main sequence" relationship between microsaccade amplitude and peak velocity. We hypothesize that visual stimulation during movement preparation affects not only the saccadic "Go" system driving eye movements but also a "Pause" system inhibiting them. If the Pause system happens to be already turned off despite the new visual input, movement kinematics can be altered by the readout of additional visually evoked spikes in the Go system coding for the flash location. Our results demonstrate precise control over individual microscopic saccades and provide testable hypotheses for mechanisms of saccade control in general.NEW & NOTEWORTHY Microsaccadic eye movements play an important role in several aspects of visual perception and cognition. However, the mechanisms for microsaccade control are still not fully understood. We found that microsaccade kinematics can be altered in a systematic manner by visual transients, revealing a previously unappreciated and exquisite level of control by the oculomotor system of even the smallest saccades. Our results suggest precise temporal interaction between visual, motor, and inhibitory signals in microsaccade control.

Modulation of microsaccades by spatial frequency during object categorization

The organization of visual processing into a coarse-to-fine information processing based on the spatial frequency properties of the input forms an important facet of the object recognition process. During visual object categorization tasks, microsaccades occur frequently. One potential functional role of these eye movements is to resolve high spatial frequency information. To assess this hypothesis, we examined the rate, amplitude and speed of microsaccades in an object categorization task in which participants viewed object and non-object images and classified them as showing either natural objects, man-made objects or non-objects. Images were presented unfiltered (broadband; BB) or filtered to contain only low (LSF) or high spatial frequency (HSF) information. This allowed us to examine whether microsaccades were modulated independently by the presence of a high-level feature – the presence of an object – and by low-level stimulus characteristics – spatial frequency. We found a bimodal distribution of saccades based on their amplitude, with a split between smaller and larger microsaccades at 0.4° of visual angle. The rate of larger saccades (⩾0.4°) was higher for objects than non-objects, and higher for objects with high spatial frequency content (HSF and BB objects) than for LSF objects. No effects were observed for smaller microsaccades (<0.4°). This is consistent with a role for larger microsaccades in resolving HSF information for object identification, and previous evidence that more microsaccades are directed towards informative image regions.

Uncorrected Myopic Refractive Error Increases Microsaccade Amplitude.

Human brain generates miniature eye movements, such as microsaccades, to counteract image fading due to visual adaptation. Generation of microsaccade relies on the amount of retinal error or acuity demand for a desired visual task. The goal of this study was to assess the influence of visual blur, induced by uncorrected refractive error on microsaccades and saccades. Ten subjects with myopia held their gaze on a visual target during two experiment conditions: corrected refractive error and uncorrected refractive error. Eye movements were measured with high-resolution video oculography under binocular viewing conditions during both tasks. Gaze holding function, microsaccades, and visually guided saccades were analyzed and compared during both tasks. We found an increase in the amplitude of microsaccades in the presence of uncorrected refractive error, but the microsaccade frequency and velocity remained unchanged. The microsaccade amplitude systematically increased with an increase in uncorrected refractive error. The main sequence relationship relating the saccade amplitude with respective peak velocity was not significantly different between two conditions. The onset latency, peak velocities, and accuracy of visually guided saccades also were unchanged between the two conditions. These results suggest that visual blur, hence the precision of an image on the fovea, has an important role in calibrating the amplitude of fixational eye movements, such as microsaccades.

Characteristics of fixational eye movements in amblyopia: Limitations on fixation stability and acuity?

Persons with amblyopia, especially those with strabismus, are known to exhibit abnormal fixational eye movements. In this paper, we compared six characteristics of fixational eye movements among normal control eyes (n=16), the non-amblyopic fellow eyes and the amblyopic eyes of anisometropic (n=14) and strabismic amblyopes (n=14). These characteristics include the frequency, magnitude of landing errors, amplitude and speed of microsaccades, and the amplitude and speed of slow drifts. Fixational eye movements were recorded using retinal imaging while observers monocularly fixated a 1° cross. Eye position data were recovered using a cross-correlation procedure. We found that in general, the characteristics of fixational eye movements are not significantly different between the fellow eyes of amblyopes and controls, and that the strabismic amblyopic eyes are always different from the other groups. Next, we determined the primary factors that limit fixation stability and visual acuity in amblyopic eyes by examining the relative importance of the different oculomotor characteristics, adding acuity (for fixation stability) or fixation stability (for acuity), and the type of amblyopia, as predictive factors in a multiple linear regression model. We show for the first time that the error magnitude of microsaccades, acuity, amplitude and frequency of microsaccades are primary factors limiting fixation stability; while the error magnitude, fixation stability, amplitude of drifts and amplitude of microsaccades are the primary factors limiting acuity. A mediation analysis showed that the effects of error magnitude and amplitude of microsaccades on acuity could be explained, at least in part, by their effects on fixation stability.

Experimental tests of hypotheses for microsaccade generation.

Fixational eye movements such as microsaccades are important to prevent fading. These miniature eye movements are also necessary to redirect gaze to the target after a drift. Generation of saccades and microsaccades utilizes common neural substrates. We, therefore, hypothesized that physiological modulators of saccades should also affect microsaccades. Test of this hypothesis will also provide support for the models of a microsaccade generation. We performed two experiments. In the first experiment, complete darkness led to a decrease in the frequency and velocity, but increased the amplitude of microsaccades. In the second experiment, active eyelid closure further reduced the velocity and frequency of microsaccades, but increased their amplitude. Darkness reduces the superior colliculus activity leading to a reduction in the velocity and frequency of microsaccades. Eye closure might cause sustained inhibition of the omnipause neurons. Subsequent disinhibition of the burst neurons might cause a reduction in the post-inhibitory rebound firing resulting in a decreased velocity of microsaccades. Sustained inhibition of the omnipause neurons could also reduce the inhibitory drive that would otherwise abort microsaccades. Hence, by inhibiting the activity of omnipause neurons, the eye closure could increase the amplitude of microsaccades.