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

Our behaviour is guided by the ability to predict future events. The predictive saccade paradigm has been shown to be a valuable tool that uses eye movements to measure the control of predictive behaviour. In this task, subjects follow a visual target that alternates or “steps” between two fixed locations at either predictable or unpredictable inter-stimulus time intervals (ISIs). Response times can be measured by subtracting the time of saccade initiation from the time of target appearance. When the ISI is predictable, saccadic reaction times (SRTs) become predictive (SRT <100ms) within 3-4 target steps, but when the ISI is unpredictable, the SRTs remain reactive to target appearance (SRT >100ms). The goal of our study was to investigate neural mechanisms controlling prediction by contrasting areas in the brain that were more active for predictive (PRED) versus reactive (REACT) saccades in young healthy adults using functional magnetic resonance imaging (fMRI). fMRI analysis revealed two distinct neural networks more recruited for REACT and PRED tasks. We observed greater activation for the REACT task compared to the PRED task in oculomotor network areas including the frontal, supplementary, parietal eye fields, dorsolateral prefrontal cortex, thalamus, and putamen. These structures are all involved with the control of saccades. We also observed greater activation for the PRED task compared to the REACT task in default network areas, including the medial prefrontal cortex, posterior cingulate cortex, inferior parietal lobule, and hippocampus. These structures are known to be involved with passive thinking when subjects are not focused on their external environments. We also observed greater activation for the PRED task in the cerebellum (crus I), which may serve as the internal clock that drives the regular rhythmic behaviour observed for predictive saccades. In summary, our findings suggest brain activation in the PRED task reflects automated and motor-timed responses, while that for the REACT task reflects externally-driven responses. Therefore, the predictive saccade task is an excellent tool for measuring prediction involving fast internally-guided responses. / Thesis (Master, Neuroscience Studies) -- Queen's University, 2011-08-12 10:21:37.744

Thesis (Ph.D.). / Written for the Dept. of Physiology. Title from title page of PDF (viewed 2009/06/10). Includes bibliographical references.

La recherche présentée ici constitue l’une des applications du projet de recherche « InterHumain », et son objectif principal est double : développer les connaissances dont on dispose actuellement sur le fonctionnement de l’interaction en général ; et, décrire avec plus de précision les capacités interactionnelles des schizophrènes dans le but de contribuer à l’élaboration de techniques de prise en charge adaptées à leurs troubles cognitifs. Afin de concevoir un modèle de l’interaction plurimodal, nous avons analysé des interactions en face à face entre un expérimentateur (psychologue) et un schizophrène. Cela nous a amené à comparer certaines propriétés des compétences interactionnelles des sujets « normaux » à celles des sujets schizophrènes afin d’identifier les capacités et les difficultés qu’ils manifestent au niveau du langage et des mouvements oculaires. Nous centrons les analyses dans ce manuscrit sur deux aspects de l’interaction : la séquentialité du discours et les saccades oculaires. Ce modèle sera éprouvé au moyen du système faceLAB5. Il s’agit d’un système d’enregistrement et de suivi des mouvements oculaires (« eye-tracking ») que nous avons doublé afin d’obtenir des données sur les deux interlocuteurs en situation d’interaction. De plus, les résultats seront mis en perspective avec les données issues de bilans neuropsychologiques afin, notamment, de déterminer le rôle que pourrait jouer le lobe frontal dans le maintien de l’interaction. Les principaux résultats sont en accord avec ceux issus de nombreuses études antérieures utilisant un seul système d’ « eye-tracking » antérieures, notamment que les patients schizophrènes produisent plus de mouvements de saccades oculaires que les participants témoins. Mais le dispositif, tel que nous l’avons conçu, permet également l’analyse du fonctionnement de l’interlocuteur donc de l’expérimentateur. Celui-ci manifeste une baisse de sa production de saccades oculaires lorsqu’il est en interaction avec les patients schizophrènes. De plus, les résultats aux tests neuropsychologiques montrent que malgré certaines déficiences déjà identifiées chez les patients schizophrènes, certaines capacités cognitives semblent préservées. Cette observation pourraient être le point d’ancrage d’une restauration des capacités cognitives actuellement en déclins voir déficitaires chez certains patients schizophrènes, par la mise au point de thérapies spécifiques et adaptées. Cela leur permettrait d’exploiter au mieux toutes leurs capacités cognitives, soit au quotidien, soit en vue d’une insertion sociale et professionnelle à long terme. / The research presented here is one of the application of the research project “inter-human” and it has a twofold objective: to develop the knowledge currently available on the functioning of the interaction in general; and to describe more accurately interactional skills of schizophrenic patients in order to contribute to the development of care techniques appropriate to their cognitive disorders. In order to design a model of multimodal interaction, we analyzed face to face interactions between an experimenter (psychologist) and a schizophrenic patient. This led us to compare some properties of interactional skills of “normal” subjects to those of schizophrenic patients in order to identify the capabilities and challenges they manifest via language and eye movements. In this thesis, we focus the analyzes on two aspects of interaction: sequentiality of speech and saccadic eye movements. This model will be tested using the faceLAB5 system. This system is a recording and monitoring system of eye movements ("eye-tracking") that we doubled to get data on both parties in a situation of interaction. In addition, the results will be put into perspective with data taken from neuropsychological evaluations, in order to identify the possible role of the frontal lobe in maintaining the interaction. The main results are consistent with those from many other studies that used an earlier single “eye-tracking” system, in particular that schizophrenic patients produce more saccadic eye movements than “normal” participants. But designed as it was, the device also allows the analysis of the behavior of the interlocutor, that is to say the experimenter. This shows a decrease in production of saccadic eye movements from the experimenter when interacting with schizophrenic patients. In addition, the results on neuropsychological tests show that despite some disabilities already identified in schizophrenic patients, some cognitive abilities seem to be preserved.This observation could be the cornerstone of the restoration of cognitive capacities that are declining or are deficient in some schizophrenia patients, by the development of specific and appropriate therapies. This would allow them to exploit all their cognitive abilities, either daily or in a view to a social and professional long term integration.

Understanding of visual scenes takes place within very brief episodes known as fixations. To explore the extent of the scene, the eye shifts between fixation locations at intervals of roughly 300 ms. Currently, it is a matter of open inquiry as to what factors influence the timing of these movements. This thesis focuses on understanding the mechanisms that govern the rapid adjustment of fixation and saccade timings when novel stimulus information is encountered during a fixation. In part I, I use an experimental technique known as the fixation-contingent scene quality paradigm to control the quality of incoming visual scene information. This approach is used to assess how fixation timing adapts to moment-by-moment changes in the quality level of the stimulus. I find that quality changes tend to result in an increase in fixation durations and this occurs whether the quality is increased or decreased. Using distributional analytic techniques, I argue that these results reflect the combined influence of a rapid surprise related process and a slower acting encoding related influence. In part II, I study how fixation durations are influenced by the underlying saccade programming mechanisms. An important assumption within the eye-movement control literature is that there exists a threshold called the point-of-no-return. Once this point has been reached, a saccade may no longer be modified or cancelled. I adapt a classic psychophysical technique known as the double-step procedure to study the point-of-no-return within scene viewing tasks. I also provide a measurement of the saccadic dead time, the last point in time that a saccade may be modified. In Part III, a formal model of fixation durations in high-level tasks is presented. I build on recent modelling work and develop a formal account for the early-surprise late-encoding modulation account of fixation durations in scene viewing tasks. The model is tested against data observed in Part I of the thesis. I demonstrate that the model does a very good job of predicting these distributions with relatively few assumptions. In summary, I use experimental techniques in combination with computational modelling to reveal how a composite of low-level (saccade programming) and high-level (information processing) considerations can, and must, be taken into consideration when understanding eye-movement control behaviour in scene viewing tasks.

Les saccades verticales sont importantes pour l'exploration du monde visuel 3D. Ces mouvements complexes nécessitent le contrôle de la distribution d'innervation aux six muscles extraoculaires de chaque ¿il. Peu d'études existent chez quelques adultes. Elles montrent une asymétrie haut/bas : latence plus courte pour les saccades vers le haut, convergence pendant les saccades vers le bas et divergence pendant les saccades vers le haut. Une controverse persiste concernant l'origine centrale versus musculaire de la vergence. Cette thèse apporte des données de référence sur le développement des saccades verticales en interaction avec la vergence chez des enfants de 6 à 17 ans. Le résultat marquant est une convergence pendant toutes les saccades verticales qui diminue avec l'âge pour les saccades vers le haut, tendant vers la divergence de l'adulte. Ainsi, les asymétries haut/bas évoluent avec l'âge ; nos résultas plaident en faveur d'une synergie continue saccade-vergence qui soutiendrait un biais perceptif (champ visuel haut perçu comme plus lointain en profondeur que le champ visuel bas). Cette thèse étudie aussi le contrôle postural, focalisant sur l'interaction vision-oculomotricité-posture des enfants de 6 à 17 ans sains et des enfants avec strabisme. Les résultats montrent un effet stabilisateur de la vergence sur la posture, l'existence d'un espace privilégié pour la stabilité posturale (lointain pour strabismes divergents, mais proche pour strabismes convergents et enfants sains) et un bénéfice de la vision bi-oculaire rudimentaire présente chez des strabiques. Cette thèse ouvre des pistes multiples de recherche fondamentale en clinique. / Vertical saccades eye movements are very important for exploration of the 3-D space. There are complex movements, requiring control of the distribution of innervation to the six extraocular muscles of each eye. Few studies exist in some adults subjects. These showed up/down anisotropies: shorter latency for upward saccades, convergence during downward and divergence during upward saccades. A controversy remains about the origin, central versus muscular, of the vergence. This thesis provides referential developmental data of vertical saccades in interaction with vergence in children from 6 to 17 years. The striking result is a convergence of the eyes during all vertical saccades that decrease with age for upward saccades, tending towards divergence like adults. Thus up/down asymmetries changed with age, in line with a continuous saccade-vergence synergy that supports a perceptual bias (upper visual field further away in depth than lower visual field). This thesis studied also postural control, focusing on the vision-oculomotricity-posture interaction in children from 6 to 17 years old, and in children with strabismus. Our results showed a stabilizing effect of vergence on posture, the existence of a favorite space for postural stability in strabismic children (near for convergent and healthy children and far for divergent strabismus) and a benefit of bi-ocular visual stimulation. This thesis opens multiple avenues for fundamental research in clinic.

When the eyes fixate at a point in a visual scene, small saccades rapidly shift the image on the retina. The effect of these microsaccades on the latency of subsequent large-scale saccades may be twofold. First, microsaccades are associated with an enhancement of visual perception. Their occurrence during saccade target perception should, thus, decrease saccade latencies. On the other hand, microsaccades likely indicate activity in fixation-related oculomotor neurons. These represent competitors to saccade-related cells in the interplay of gaze holding and shifting. Consequently, an increase in saccade latencies after microsaccades would be expected. Here, we present evidence for both aspects of microsaccadic impact on saccade latency. In a delayed response task, participants made saccades to visible or memorized targets. First, microsaccade occurrence up to 50 ms before target disappearance correlated with 18 ms (or 8%) faster saccades to memorized targets. Second, if microsaccades occurred shortly (i.e., < 150 ms) before a saccade was required, saccadic reaction times in visual and memory trials were increased by about 40 ms (or 16%). Hence, microsaccades can have opposite consequences for saccade latencies, pointing at a differential role of these fixational eye movements in preparation of motor programs.

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.

Bakalauro baigiamajame darbe tiriamas įcentrinių ir išcentrinių žmogaus akies sakadų tikslumas. Eksperimento metu, kompiuterio ekrane, horizontalia kryptimi per penkias fiksuotas padėtis, šuoline trajektorija judėjo žalias (0,25 laipsnio diametro) taškas. Tiriamųjų paprašyta žvilgsniu sekti šį tašką. Apdorojus gautus tyrimo duomenis pastebėta, kad tos pačios amplitudės įcentrinės sakados yra tikslesnės, nei išcentrinės. / In this Bachelor thesis accuracy of the centripetal and centrifugal human eye saccades was studied. During the experiment, on the computer screen, in horizontal direction within five fixed positions, the hopping green spot (diameter of 0,25 degrees) was moving. Subjects were instructed to follow this green spot with gaze. Processing the research data reported that the centripetal saccades is more accurate than the centrifugal of the same amplitude.

In order to grasp an object in their visual field, humans orient their visual axis to targets of interest. While scanning their environment, humans perform multiple saccades (rapid eye movements that correct for a position error between eye and target) to align their visual axis with objects of interest. Humans are also able to track objects that move in their environment by means of smooth pursuit eye movements (slow eye movements that correct for any velocity error between eye and target, i.e. for any retinal slip). The appearance of a moving stimulus in the environment elicits smooth pursuit eye movements with a latency of around 100ms. Accordingly, the smooth pursuit system accounts for a change in the trajectory of a moving target with a similar delay. Due to this delay, the oculomotor system needs to develop strategies to avoid the build up of position error during tracking of a moving target. To do so, the oculomotor system uses prediction to try and anticipate the future target trajectory. However, this strategy is limited to conditions where target trajectory is predictable. Otherwise, primates have to combine pursuit and saccades in visual tracking of unpredictable moving targets to avoid large position error. This thesis focuses on both the prediction mechanisms and the interactions between saccades and pursuit. In order to investigate prediction mechanisms, we asked human subjects to pursue a moving target when it was transiently occluded. During occlusions, subjects continued to pursue the invisible target. This thesis demonstrates that this predictive pursuit response is based on a dynamic internal representation of target motion, i.e. a representation that evolves with time. This internal representation could be either built up by repetition of the same target motion or extrapolated on the basis of the pre-occlusion target motion. In addition, it is shown that during occlusions, saccades are adjusted in order to account for the large variability of the smooth pursuit response. As a consequence, it shows that the smooth pursuit command is used by internal models in order to predict future smooth pursuit response. These results demonstrate that both prediction and internal models are necessary to track the invisible and the visible.