TY - THES A1 - Ohl, Sven T1 - Small eye movements during fixation : the case of postsaccadic fixation and preparatory influences T1 - Kleine Augenbewegungen während der Fixation: Der Fall der postsakkadischen Fixation und vorbereitender Einflüsse N2 - 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. N2 - 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. KW - Augenbewegungen KW - Mikrosakkaden KW - Sekundärsakkaden KW - Korrektursakkaden KW - Landepositionsfehler KW - eye movements KW - microsaccades KW - secondary saccades KW - corrective saccades KW - saccadic error Y1 - 2013 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus-69862 ER - TY - CHAP A1 - Ohl, Sven A1 - Brandt, S. A1 - Kliegl, Reinhold T1 - Immediate preparatory influences on microsaccades before saccade onset to endogenously vs. exogenously defined targets T2 - Perception Y1 - 2013 SN - 0301-0066 SN - 1468-4233 VL - 42 IS - 4 SP - 37 EP - 38 PB - Sage Publ. CY - London ER - TY - JOUR A1 - Ohl, Sven A1 - Brandt, Stephan A. A1 - Kliegl, Reinhold T1 - Secondary (micro-)saccades the influence of primary saccade end point and target eccentricity on the process of postsaccadic fixation JF - Vision research : an international journal for functional aspects of vision. N2 - We examine how the size of saccadic under-/overshoot and target eccentricity influence the latency, amplitude and orientation of secondary (micro-)saccades. In our experiment, a target appeared at an eccentricity of either 6 degrees or 14 degrees of visual angle. Subjects were instructed to direct their gaze as quickly as possible to the target and hold fixation at the new location until the end of the trial. Typically, increasing saccadic error is associated with faster and larger secondary saccades. We show that secondary saccades at distant in contrast to close targets have in a specific error range a shorter latency, larger amplitude, and follow more often the direction of the primary saccade. Finally, we demonstrate that an undershooting primary saccade is followed almost exclusively by secondary saccades into the same direction while overshooting primary saccades are followed by secondary saccades into both directions. This supports the notion that under- and overshooting imply different consequences for postsaccadic oculomotor processing. Results are discussed using a model, introduced by Rolfs, Kliegl, and Engbert (2008), to account for the generation of microsaccades. We argue that the dynamic interplay of target eccentricity and the magnitude of the saccadic under-/overshoot can be explained by a different strength of activation in the two hemispheres of the saccadic motor map in this model. KW - Secondary saccade KW - Microsaccade KW - Saccadic error KW - Error-correction KW - Target eccentricity Y1 - 2011 U6 - https://doi.org/10.1016/j.visres.2011.09.005 SN - 0042-6989 SN - 1878-5646 VL - 51 IS - 23-24 SP - 2340 EP - 2347 PB - Elsevier CY - Oxford ER - TY - CHAP A1 - Ohl, Sven A1 - Brandt, S. A1 - Kliegl, Reinhold T1 - Post-saccadic location judgments after presentation of multiple target-like objects T2 - Perception Y1 - 2012 SN - 0301-0066 SN - 1468-4233 VL - 41 IS - 1 SP - 171 EP - 171 PB - Sage Publ. CY - London ER - TY - JOUR A1 - Ohl, Sven A1 - Brandt, Stephan A. A1 - Kliegl, Reinhold T1 - The generation of secondary saccades without postsaccadic visual feedback JF - Journal of vision N2 - Primary saccades are often followed by small secondary saccades, which are generally thought to reduce the distance between the saccade endpoint and target location. Accumulated evidence demonstrates that secondary saccades are subject to various influences, among which retinal feedback during postsaccadic fixation constitutes only one important signal. Recently, we reported that target eccentricity and an orientation bias influence the generation of secondary saccades. In the present study, we examine secondary saccades in the absence of postsaccadic visual feedback. Although extraretinal signals (e.g., efference copy) have received widespread attention in eye-movement studies, it is still unclear whether an extraretinal error signal contributes to the programming of secondary saccades. We have observed that secondary saccade latency and amplitude depend on primary saccade error despite the absence of postsaccadic visual feedback. Strong evidence for an extraretinal error signal influencing secondary saccade programming is given by the observation that secondary saccades are more likely to be oriented in a direction opposite to the primary saccade as primary saccade error shifts from target undershoot to overshoot. We further show how the functional relationship between primary saccade landing position and secondary saccade characteristics varies as a function of target eccentricity. We propose that initial target eccentricity and an extraretinal error signal codetermine the postsaccadic activity distribution in the saccadic motor map when no visual feedback is available. KW - monocular deprivation KW - binocular combination KW - sensory balance Y1 - 2013 U6 - https://doi.org/10.1167/13.5.11 SN - 1534-7362 VL - 13 IS - 5 PB - Association for Research in Vision and Opthalmology CY - Rockville ER - TY - JOUR A1 - Rolfs, Martin A1 - Ohl, Sven T1 - Visual suppression in the superior colliculus around the time of microsaccades JF - Journal of neurophysiology N2 - Miniature eye movements jitter the retinal image unceasingly, raising the question of how perceptual continuity is achieved during visual fixation. Recent work discovered suppression of visual bursts in the superior colliculus around the time of microsaccades, tiny jerks of the eyes that support visual perception while gaze is fixed. This finding suggests that corollary discharge, supporting visual stability when rapid eye movements drastically shift the retinal image, may also exist for the smallest saccades. Y1 - 2011 U6 - https://doi.org/10.1152/jn.00862.2010 SN - 0022-3077 VL - 105 IS - 1 SP - 1 EP - 3 PB - American Chemical Society CY - Bethesda ER - TY - JOUR A1 - Ohl, Sven A1 - Kliegl, Reinhold T1 - Revealing the time course of signals influencing the generation of JF - Vision research : an international journal for functional aspects of vision. N2 - Saccadic eye movements are frequently followed by smaller secondary saccades which are generally assumed to correct for the error in primary saccade landing position. However, secondary saccades can also occur after accurate primary saccades and they are often as small as microsaccades, therefore raising the need to further scrutinize the processes involved in secondary saccade generation. Following up a previous study, we analyzed secondary saccades using rate analysis which allows us to quantify experimental effects as shifts in distributions, therefore going beyond comparisons of mean differences. We use Aalen’s additive hazards model to delineate the time course of key influences on the secondary saccade rate. In addition to the established effect of primary saccade error, we observed a time-varying influence of under- vs. overshooting – with a higher risk of generating secondary saccades following undershoots. Moreover, increasing target eccentricity influenced the programming of secondary saccades, therefore demonstrating that error-unrelated variables co-determine secondary saccade programs. Our results provide new insights into the generative mechanisms of small saccades during postsaccadic fixation that need to be accounted for by secondary saccade models. KW - Eye movements KW - Corrective saccades KW - Secondary saccades KW - Rate analysis KW - Survival analysis Y1 - 2016 U6 - https://doi.org/10.1016/j.visres.2016.06.007 SN - 0042-6989 SN - 1878-5646 VL - 124 SP - 52 EP - 58 PB - Elsevier CY - Oxford ER - TY - JOUR A1 - Ohl, Sven A1 - Wohltat, Christian A1 - Kliegl, Reinhold A1 - Pollatos, Olga A1 - Engbert, Ralf T1 - Microsaccades Are Coupled to Heartbeat JF - The journal of neuroscience N2 - During visual fixation, the eye generates microsaccades and slower components of fixational eye movements that are part of the visual processing strategy in humans. Here, we show that ongoing heartbeat is coupled to temporal rate variations in the generation of microsaccades. Using coregistration of eye recording and ECG in humans, we tested the hypothesis that microsaccade onsets are coupled to the relative phase of the R-R intervals in heartbeats. We observed significantly more microsaccades during the early phase after the R peak in the ECG. This form of coupling between heartbeat and eye movements was substantiated by the additional finding of a coupling between heart phase and motion activity in slow fixational eye movements; i.e., retinal image slip caused by physiological drift. Our findings therefore demonstrate a coupling of the oculomotor system and ongoing heartbeat, which provides further evidence for bodily influences on visuomotor functioning. KW - eye movements KW - heartbeat KW - microsaccades Y1 - 2016 U6 - https://doi.org/10.1523/JNEUROSCI.2211-15.2016 SN - 0270-6474 VL - 36 SP - 1237 EP - 1241 PB - Society for Neuroscience CY - Washington ER -