Individuals with Parkinson’s disease (PD) display deficits in voluntary saccade generation but improved automatic, visually-triggered saccade performance. This can be tested using prosaccades, saccades to visual stimuli, and antisaccades, saccades in the opposite direction from the visual stimuli. Voluntary saccade impairments resulting in antisaccade direction errors and longer saccadic reaction times (SRTs) are thought to be due to insufficient presetting of neural circuitry during saccade preparation in complex tasks involving suppression and selection. The basal ganglia, a major site of PD pathology, might be the cause of abnormalities in preparing for action selection in PD patients. Recently, microsaccade rates have been hypothesized to reflect the dual preparatory signals of saccade facilitation and suppression. In this thesis, we investigated the microsaccade behaviour of PD patients as they performed prosaccades and antisaccades. We hypothesized that deficits in voluntary movements in PD would result in impaired suppression of involuntary movements as reflected by increased microsaccade rates. Our findings demonstrate consistently elevated microsaccade rates in PD subjects compared to age-matched controls. Furthermore, positive correlations were found between antisaccade direction error rate and microsaccade rate as well as microsaccade rate and Hoehn-Yahr score, an indicator of disease severity in PD patients. We conclude that microsaccades reflect the impaired suppression of involuntary movements caused by voluntary movement deficits in PD pathology. Our findings indicate that microsaccades provide insight into action preparatory mechanisms and BG dysfunction. Therefore, measuring microsaccades in PD may provide a useful biomarker to follow disease progression and effectiveness of treatment therapies. / Thesis (Master, Neuroscience Studies) -- Queen's University, 2014-01-09 23:31:21.78

Yes / Purpose: Even during steady fixation, people make small eye movements such as microsaccades, whose rate is altered by presentation of salient stimuli. Our goal was to develop a practical method for objectively and robustly estimating contrast sensitivity from microsaccade rates in a diverse population. Methods: Participants, recruited to cover a range of contrast sensitivities, were visually normal (n = 19), amblyopic (n = 10), or had cataract (n = 9). Monocular contrast sensitivity was estimated behaviorally while binocular eye movements were recorded during interleaved passive trials. A probabilistic inference approach was used to establish the likelihood of observed microsaccade rates given the presence or absence of a salient stimulus. Contrast sensitivity was estimated from a function fitted to the scaled log-likelihood ratio of the observed microsaccades in the presence or absence of a salient stimulus across a range of contrasts. Results: Microsaccade rate signature shapes were heterogeneous; nevertheless, estimates of contrast sensitivity could be obtained in all participants. Microsaccade-estimated contrast sensitivity was unbiased compared to behavioral estimates (1.2% mean), with which they were strongly correlated (Spearman's ρ 0.74, P < 0.001, median absolute difference 7.6%). Measurement precision of microsaccade-based contrast sensitivity estimates was worse than that of behavioral estimates, requiring more than 20 times as many presentations to equate precision. Conclusions: Microsaccade rate signatures are heterogeneous in shape when measured across populations with a broad range of contrast sensitivities. Contrast sensitivity can be robustly estimated from rate signatures by probabilistic inference, but more stimulus presentations are currently required to achieve similarly precise estimates to behavioral techniques. / Supported by a Confidence in Concept grant from the Medical Research Council, a Fight for Sight Project Grant (5059/5060) and a Wellcome Trust Research Fellowship to NWR (WT097387).

Fixational eye movements occur involuntarily during visual fixation of stationary scenes. The fastest components of these miniature eye movements are microsaccades, which can be observed about once per second. Recent studies demonstrated that microsaccades are linked to covert shifts of visual attention [e.g., Engbert & Kliegl (2003), Vision Res 43:1035-1045]. Here,we generalized this finding in two ways. First, we used peripheral cues, rather than the centrally presented cues of earlier studies. Second, we spatially cued attention in vision and audition to visual and auditory targets. An analysis of microsaccade responses revealed an equivalent impact of visual and auditory cues on microsaccade-rate signature (i.e., an initial inhibition followed by an overshoot and a final return to the pre-cue baseline rate). With visual cues or visual targets,microsaccades were briefly aligned with cue direction and then opposite to cue direction during the overshoot epoch, probably as a result of an inhibition of an automatic saccade to the peripheral cue. With left auditory cues and auditory targets microsaccades oriented in cue direction. Thus, microsaccades can be used to study crossmodal integration of sensory information and to map the time course of saccade preparation during covert shifts of visual and auditory attention.

abstract: During attempted fixation, the eyes are not still but continue to produce so called "fixational eye movements", which include microsaccades, drift, and tremor. Microsaccades are thought to help prevent and restore vision loss during fixation, and to correct fixation errors, but how they contribute to these functions remains a matter of debate. This dissertation presents the results of four experiments conducted to address current controversies concerning the role of microsaccades in visibility and oculomotor control. The first two experiments set out to correlate microsaccade production with the visibility of foveal and peripheral targets of varied spatial frequencies, during attempted fixation. The results indicate that microsaccades restore the visibility of both peripheral targets and targets presented entirely within the fovea, as a function of their spatial frequency characteristics. The last two experiments set out to determine the role of microsaccades and drifts on the correction of gaze-position errors due to blinks in human and non-human primates, and to characterize microsaccades forming square-wave jerks (SWJs) in non-human primates. The results showed that microsaccades, but not drifts, correct gaze-position errors due to blinks, and that SWJ production and dynamic properties are equivalent in human and non-human primates. These combined findings suggest that microsaccades, like saccades, serve multiple and non-exclusive functional roles in vision and oculomotor control, as opposed to having a single specialized function. / Dissertation/Thesis / Doctoral Dissertation Neuroscience 2014

No description available.

The human eyes are always moving. Even during periods of fixation when visual information is acquired, a persistent jittering of the eyes (ocular drift) is occasionally interrupted by small rapid gaze shifts (microsaccades). Though much has been learned in the last 20 years about the perceptual roles of fixational eye movements, little is known about the consequences of their active control for fine pattern vision and depth perception. Using custom techniques for high-resolution eye-tracking and precise control of retinal stimulation, this dissertation describes three studies that investigated the consequences of controlled fixational eye movements for visual perception of fine patterns in two and three dimensions. The first study addresses whether fixational eye movements are controlled to meet the needs of a demanding visual task and their contributions to visual acuity. We show that in a standard acuity test, humans actively tune their drifts to enhance relevant spatial information and control their microsaccades to precisely place stimuli within the foveola. Together these eye movements contribute 0.15 logMAR to visual acuity, approximately two lines of an eye chart. The second study addresses the perceptual and computational impact of tuning ocular drift. We show that humans are sensitive to changes in visual flow generated by drifts of different sizes. Changes in sensitivity are fully predicted by changes in effective power of luminance modulations delivered by drift, suggesting that drift acts as a mechanism for controlling the effective contrast of the retinal stimulus. The third study addresses the impact of binocular fixational eye movements on fine depth perception. We show that these movements, specifically the opposing movements of the eyes (vergence), are beneficial for stereovision. In the absence of disparity modulations from fixational vergence, fine depth perception is significantly impaired. The research described in this dissertation advances the field in several fundamental ways by showing that (a) contrary to traditional assumptions, ocular drift is tuned to the demands of the visual task; (b) the precise spatiotemporal structure of the luminance changes from ocular drift predictably impacts visual sensitivity; and (c) stereoscopic vision is a dynamic process that uses temporal disparity modulations generated by fixational vergence. / 2024-02-02T00:00:00Z

It has recently been demonstrated that the presentation of a rare target in a visual oddball paradigm induces a prolonged inhibition of microsaccades. In the field of electrophysiology, the amplitude of the P300 component in event-related potentials (ERP) has been shown to be sensitive to the stimulus category (target vs. non target) of the eliciting stimulus, its overall probability, and the preceding stimulus sequence. In the present study we further specify the functional underpinnings of the prolonged microsaccadic inhibition in the visual oddball task, showing that the stimulus category, the frequency of a stimulus and the preceding stimulus sequence influence microsaccade rate. Furthermore, by co-recording ERPs and eye-movements, we were able to demonstrate that, despite being largely sensitive to the same experimental manipulation, the amplitude of P300 and the microsaccadic inhibition predict each other very weakly, and thus constitute two independent measures of the brain’s response to rare targets in the visual oddball paradigm.

Describing human eye movement behavior as an alternating sequence of saccades and fixations turns out to be an oversimplification because the eyes continue to move during fixation. Small-amplitude saccades (e.g., microsaccades) are typically observed 1-2 times per second during fixation. Research on microsaccades came in two waves. Early studies on microsaccades were dominated by the question whether microsaccades affect visual perception, and by studies on the role of microsaccades in the process of fixation control. The lack of evidence for a unique role of microsaccades led to a very critical view on the importance of microsaccades. Over the last years, microsaccades moved into focus again, revealing many interactions with perception, oculomotor control and cognition, as well as intriguing new insights into the neurophysiological implementation of microsaccades. In contrast to early studies on microsaccades, recent findings on microsaccades were accompanied by the development of models of microsaccade generation. While the exact generating mechanisms vary between the models, they still share the assumption that microsaccades are generated in a topographically organized saccade motor map that includes a representation for small-amplitude saccades in the center of the map (with its neurophysiological implementation in the rostral pole of the superior colliculus). In the present thesis I criticize that models of microsaccade generation are exclusively based on results obtained during prolonged presaccadic fixation. I argue that microsaccades should also be studied in a more natural situation, namely the fixation following large saccadic eye movements. Studying postsaccadic fixation offers a new window to falsify models that aim to account for the generation of small eye movements. I demonstrate that error signals (visual and extra-retinal), as well as non-error signals like target eccentricity influence the characteristics of small-amplitude eye movements. These findings require a modification of a model introduced by Rolfs, Kliegl and Engbert (2008) in order to account for the generation of small-amplitude saccades during postsaccadic fixation. Moreover, I present a promising type of survival analysis that allowed me to examine time-dependent influences on postsaccadic eye movements. In addition, I examined the interplay of postsaccadic eye movements and postsaccadic location judgments, highlighting the need to include postsaccadic eye movements as covariate in the analyses of location judgments in the presented paradigm. In a second goal, I tested model predictions concerning preparatory influences on microsaccade generation during presaccadic fixation. The observation, that the preparatory set significantly influenced microsaccade rate, supports the critical model assumption that increased fixation-related activity results in a larger number of microsaccades. In the present thesis I present important influences on the generation of small-amplitude saccades during fixation. These eye movements constitute a rich oculomotor behavior which still poses many research questions. Certainly, small-amplitude saccades represent an interesting source of information and will continue to influence future studies on perception and cognition. / Die Beschreibung des Blickbewegungsverhaltens als eine sich abwechselnde Folge von Sakkaden und Fixationen stellt eine starke Vereinfachung dar, denn auch während einer Fixation bewegen sich die Augen. Typischerweise treten Bewegungen von kleiner Amplitude (z.B. Mikrosakkaden), 1-2 mal pro Sekunde während einer Fixation auf. Frühe Studien zu Mikrosakkaden wurden von Fragen bezüglich des Einflusses von Mikrosakkaden auf die visuelle Wahrnehmung, und Studien zu der Rolle von Mikrosakkaden bei der Fixationskontrolle dominiert. Fehlende Evidenz für eine Rolle, die ausschließlich Mikrosakkaden zufällt, führten zu einer sehr kritischen Betrachtung von Mikrosakkaden. In den letzten Jahren rückten Mikrosakkaden wieder mehr in den Fokus. Vielerlei Zusammenhänge mit Wahrnehmung, okulomotorischer Kontrolle und Kognition, sowie neue Erkenntnisse bezüglich der neurophysiologischen Implementierung von Mikrosakkaden konnten aufgedeckt werden. In den letzten Jahren wurden verschiedene Modelle der Mikrosakkadengenerierung vorgestellt. Auch wenn sich diese in ihren exakten Mechanismen unterscheiden, so teilen sie doch die Annahme, dass Mikrosakkaden in einer topographisch organisierten motorischen Karte für Sakkaden ausgelöst werden. Diese Karten beinhalten eine Repräsentation für klein-amplitudige Sakkaden im Zentrum der Karte (mit dem rostralen Pol der colliculi superiores als neurophysiologische Implementierung). In der vorliegenden Arbeit kritisiere ich, dass Modelle der Mikrosakkadengenerierung ausschließlich auf Resultaten langanhaltender präsakkadischer Fixation beruhen. Ich führe an, dass Mikrosakkaden in einer natürlicheren Situation untersucht werden sollten, nämlich während der Fixation nach einer großen Sakkade. Die Untersuchung postsakkadischer Fixation bietet eine neue Möglichkeit Modelle der Mikrosakkadengenerierung zu falsifizieren. In den Studien zeige ich, dass Signale über den Fehler in der Sakkadenlandeposition (visuelle und extra-retinale), sowie fehler-unabhängige Signale, wie die Zielreiz-Exzentrizität, einen entscheidenden Einfluss auf kleine Sakkaden haben. Diese Resultate erfordern Modifikationen an dem kürzlich eingeführten Modell von Rolfs, Kliegl und Engbert (2008), um die Generierung von kleinen Sakkaden auch während der postsakkadischen Fixation erklären zu können. Darüber hinaus präsentiere ich eine viel versprechende Ereigniszeitanalyse, die uns erlaubt zeitabhängige Einflüsse auf das postsakkadische Blickbewegungsverhalten zu untersuchen. Außerdem untersuche ich das Zusammenspiel von postsakkadischen Augenbewegungen und postsakkadischen Positionsurteilen. Dabei wird die Bedeutung von postsakkadischen Augenbewegungen als Kovariate in den statistischen Analysen betont. Ein zweites Ziel dieser Arbeit besteht darin Modellvorhersagen bezüglich vorbereitender Einflüsse auf die Mikrosakkadengenerierung zu untersuchen. Die Ergebnisse, hinsichtlich eines signifikanten Einflusses des preparatory set auf die Mikrosakkadenrate unterstützt die wesentliche Modellannahme, dass erhöhte fixationsbezogene Aktivität zu einer größeren Anzahl an Mikrosakkaden führt. In der vorliegenden Arbeit präsentiere ich wichtige Einflüsse auf die Generierung von kleinen Sakkaden während der Fixation. Diese Augenbewegungen stellen ein vielseitiges okulomorisches Verhalten dar, welche weiterhin zahlreiche Fragen mit sich bringen und sicherlich zukünftige Studien zu Wahrnehmung und Kognition beeinflussen werden.

In normal everyday viewing, we perform large eye movements (saccades) and miniature or fixational eye movements. Most of our visual perception occurs while we are fixating. However, our eyes are perpetually in motion. Properties of these fixational eye movements, which are partly controlled by the brainstem, change depending on the task and the visual conditions. Currently, fixational eye movements are poorly understood because they serve the two contradictory functions of gaze stabilization and counteraction of retinal fatigue. In this dissertation, we investigate the spatial and temporal properties of time series of eye position acquired from participants staring at a tiny fixation dot or at a completely dark screen (with the instruction to fixate a remembered stimulus); these time series were acquired with high spatial and temporal resolution. First, we suggest an advanced algorithm to separate the slow phases (named drift) and fast phases (named microsaccades) of these movements, which are considered to play different roles in perception. On the basis of this identification, we investigate and compare the temporal scaling properties of the complete time series and those time series where the microsaccades are removed. For the time series obtained during fixations on a stimulus, we were able to show that they deviate from Brownian motion. On short time scales, eye movements are governed by persistent behavior and on a longer time scales, by anti-persistent behavior. The crossover point between these two regimes remains unchanged by the removal of microsaccades but is different in the horizontal and the vertical components of the eyes. Other analyses target the properties of the microsaccades, e.g., the rate and amplitude distributions, and we investigate, whether microsaccades are triggered dynamically, as a result of earlier events in the drift, or completely randomly. The results obtained from using a simple box-count measure contradict the hypothesis of a purely random generation of microsaccades (Poisson process). Second, we set up a model for the slow part of the fixational eye movements. The model is based on a delayed random walk approach within the velocity related equation, which allows us to use the data to determine control loop durations; these durations appear to be different for the vertical and horizontal components of the eye movements. The model is also motivated by the known physiological representation of saccade generation; the difference between horizontal and vertical components concurs with the spatially separated representation of saccade generating regions. Furthermore, the control loop durations in the model suggest an external feedback loop for the horizontal but not for the vertical component, which is consistent with the fact that an internal feedback loop in the neurophysiology has only been identified for the vertical component. Finally, we confirmed the scaling properties of the model by semi-analytical calculations. In conclusion, we were able to identify several properties of the different parts of fixational eye movements and propose a model approach that is in accordance with the described neurophysiology and described limitations of fixational eye movement control. / Während des alltäglichen Sehens führen wir große (Sakkaden) und Miniatur- oder fixationale Augenbewegungen durch. Die visuelle Wahrnehmung unserer Umwelt geschieht jedoch maßgeblich während des sogenannten Fixierens, obwohl das Auge auch in dieser Zeit ständig in Bewegung ist. Es ist bekannt, dass die fixationalen Augenbewegungen durch die gestellten Aufgaben und die Sichtbedingungen verändert werden. Trotzdem sind die Fixationsbewegungen noch sehr schlecht verstanden, besonders auch wegen ihrer zwei konträren Hauptfunktionen: Das stabilisieren des Bildes und das Vermeiden der Ermüdung retinaler Rezeptoren. In der vorliegenden Dissertation untersuchen wir die zeitlichen und räumlichen Eigenschaften der Fixationsbewegungen, die mit hoher zeitlicher und räumlicher Präzision aufgezeichnet wurden, während die Versuchspersonen entweder einen sichtbaren Punkt oder aber den Ort eines verschwundenen Punktes in völliger Dunkelheit fixieren sollten. Zunächst führen wir einen verbesserten Algorithmus ein, der die Aufspaltung in schnelle (Mikrosakkaden) und langsame (Drift) Fixationsbewegungen ermöglicht. Den beiden Typen von Fixationsbewegungen werden unterschiedliche Beiträge zur Wahrnehmung zugeschrieben. Anschließend wird für die Zeitreihen mit und ohne Mikrosakkaden das zeitliche Skalenverhalten untersucht. Für die Fixationsbewegung während des Fixierens auf den Punkt konnten wir feststellen, dass diese sich nicht durch Brownsche Molekularbewegung beschreiben lässt. Stattdessen fanden wir persistentes Verhalten auf den kurzen und antipersistentes Verhalten auf den längeren Zeitskalen. Während die Position des Übergangspunktes für Zeitreihen mit oder ohne Mikrosakkaden gleich ist, unterscheidet sie sich generell zwischen horizontaler und vertikaler Komponente der Augen. Weitere Analysen zielen auf Eigenschaften der Mikrosakkadenrate und -amplitude, sowie Auslösemechanismen von Mikrosakkaden durch bestimmte Eigenschaften der vorhergehenden Drift ab. Mittels eines Kästchenzählalgorithmus konnten wir die zufällige Generierung (Poisson Prozess) ausschließen. Des weiteren setzten wir ein Modell auf der Grundlage einer Zufallsbewegung mit zeitverzögerter Rückkopplung für den langsamen Teil der Augenbewegung auf. Dies erlaubt uns durch den Vergleich mit den erhobenen Daten die Dauer des Kontrollkreislaufes zu bestimmen. Interessanterweise unterscheiden sich die Dauern für vertikale und horizontale Augenbewegungen, was sich jedoch dadurch erklären lässt, dass das Modell auch durch die bekannte Neurophysiologie der Sakkadengenerierung, die sich räumlich wie auch strukturell zwischen vertikaler und horizontaler Komponente unterscheiden, motiviert ist. Die erhaltenen Dauern legen für die horizontale Komponente einen externen und für die vertikale Komponente einen internen Kontrollkreislauf dar. Ein interner Kontrollkreislauf ist nur für die vertikale Kompoente bekannt. Schließlich wird das Skalenverhalten des Modells noch semianalytisch bestätigt. Zusammenfassend waren wir in der Lage, unterschiedliche Eigenschaften von Teilen der Fixationsbewegung zu identifizieren und ein Modell zu entwerfen, welches auf der bekannten Neurophysiologie aufbaut und bekannte Einschränkungen der Kontrolle der Fixationsbewegung beinhaltet.

Until recently induced gamma-band activity (GBA) was considered a neural marker of cortical object representation. However, induced GBA in the electroencephalogram (EEG) is susceptible to artifacts caused by miniature fixational saccades. Recent studies have demonstrated that fixational saccades also reflect high-level representational processes. Do high-level as opposed to low-level factors influence fixational saccades? What is the effect of these factors on artifact-free GBA? To investigate this, we conducted separate eye tracking and EEG experiments using identical designs. Participants classified line drawings as objects or non-objects. To introduce low-level differences, contours were defined along different directions in cardinal color space: S-cone-isolating, intermediate isoluminant, or a full-color stimulus, the latter containing an additional achromatic component. Prior to the classification task, object discrimination thresholds were measured and stimuli were scaled to matching suprathreshold levels for each participant. In both experiments, behavioral performance was best for full-color stimuli and worst for S-cone isolating stimuli. Saccade rates 200–700 ms after stimulus onset were modulated independently by low and high-level factors, being higher for full-color stimuli than for S-cone isolating stimuli and higher for objects. Low-amplitude evoked GBA and total GBA were observed in very few conditions, showing that paradigms with isoluminant stimuli may not be ideal for eliciting such responses. We conclude that cortical loops involved in the processing of objects are preferentially excited by stimuli that contain achromatic information. Their activation can lead to relatively early exploratory eye movements even for foveally-presented stimuli.