When the eye rotates, switching from one fixation point to another, the perception of motion is strongly suppressed and rarely perceived. During these quick ‘saccadic’ eye movements, other aspects of visual perception become suppressed or compressed as well, with certain effects being stronger or weaker along the plane of the saccade - such differences can help identify the underlying neuronal pathways, since some exhibit directional tuning (e.g. neurons projecting from primate V1 to middle temporal area (MT)), and others do not (e.g. relay neurons linking the superior colliculus to area MT). A briefly presented motion probe was placed at a number of points relative to saccade to plot sensitivity to motion along different planes and directions. The results suggest that saccadic motion is suppressed before the eye begins to move, and is applied evenly across planes and directions.

When the eye rotates, switching from one fixation point to another, the perception of motion is strongly suppressed and rarely perceived. During these quick ‘saccadic’ eye movements, other aspects of visual perception become suppressed or compressed as well, with certain effects being stronger or weaker along the plane of the saccade - such differences can help identify the underlying neuronal pathways, since some exhibit directional tuning (e.g. neurons projecting from primate V1 to middle temporal area (MT)), and others do not (e.g. relay neurons linking the superior colliculus to area MT). A briefly presented motion probe was placed at a number of points relative to saccade to plot sensitivity to motion along different planes and directions. The results suggest that saccadic motion is suppressed before the eye begins to move, and is applied evenly across planes and directions.

Considerable evidence indicates that visual sensitivity is reduced during saccadic eye movement. A central question has been whether saccadic suppression results from a non-visual central signal, or whether the obligate image motion that accompanies saccades is itself sufficient to mask vision. In the first of a series of experiments described here, the visual and non-visual effects of saccades were distinguished by measuring contrast sensitivity to luminance modulated low spatial frequency gratings, at 17 cd·m¯² and 0.17 cd·m¯², in saccade conditions and in conditions in which saccade-like image motion was produced by the rotation of a mirror but when observers’ eyes were kept still. The time course of suppression was examined by making measurements from well before image motion began until well after it had ended. A tenfold decrease in contrast sensitivity was found for luminance-modulated gratings with saccades, but little suppression was found with simulated saccades. Adding high contrast noise to the visual display increased the magnitude and the duration of the suppression during simulated saccades but had little effect on suppression produced by real saccades. At lower luminance, suppression was found to be reduced, and its course shallower than at higher luminance. Simulated saccades produced shallower suppression over a longer time course at both higher and lower luminance. In a second experiment the time course of contrast sensitivity to chromatically modulated gratings, at 17 cd·m¯², was examined. No suppression was found; rather there was some evidence of an enhancement of sensitivity, both before and after saccades, relative to fixation conditions. Differences in the effects of real and simulated saccades in the magnitude and time course of sensitivity loss with luminance modulated gratings suggest that saccadic suppression has an extraretinal component that acts on the magnocellular system; the pattern of enhancement found in the later experiment suggests a selective favouring of the parvocellular system both immediately prior to and immediately after saccades. The possibility that the degree of enhancement in sensitivity varies across the visual field was examined using spatially localized stimuli (either high spatial frequency chromatically modulated gratings or letter combinations). Sensitivity was found to decrease at the initial fixation point during the 75 ms prior to saccadic onset and simultaneously to improve at the saccadic target. In the immediate post-saccadic period, sensitivity at the saccadic target was found to exceed that which had been manifest at the initial fixation point prior to saccades, suggesting that post-saccadic enhancement may improve the temporal contrast between one fixation and the next. The final experiments investigated the possibility that our sense of continuity across saccades (as opposed to stability) is influenced by saccade-induced errors in locating events in time. The results of these experiments suggest that saccades can result in errors in judging (a) the time at which external events occur relative to saccadic onset, (b) the temporal order of visual events, and (c) the magnitude of temporal intervals. It is concluded that apparent time is generally foreshortened prior to saccades. This might be due to selective suppression of magnocellular activity and might function to hide saccades and their effects from our awareness. A speculative synthesis is presented based on the idea that recurrent feedback between the neocortical and cortical structures on the one hand, and the thalamic nuclei on the other, has special importance for perception around the time of saccades

The understanding of the subjective experience of a visually stable world despite the occurrence of an observer's eye movements has been the focus of extensive research for over 20 years. These studies have revealed fundamental mechanisms such as anticipatory receptive field (RF) shifts and the saccadic suppression of stimulus displacements, yet there currently exists no single explanatory framework for these observations. We show that a previously presented neuro-computational model of peri-saccadic mislocalization accounts for the phenomenon of predictive remapping and for the observation of saccadic suppression of displacement (SSD). This converging evidence allows us to identify the potential ingredients of perceptual stability that generalize beyond different data sets in a formal physiology-based model. In particular we propose that predictive remapping stabilizes the visual world across saccades by introducing a feedback loop and, as an emergent result, small displacements of stimuli are not noticed by the visual system. The model provides a link from neural dynamics, to neural mechanism and finally to behavior, and thus offers a testable comprehensive framework of visual stability.

Das visuelle System erreicht enorme Verarbeitungsmengen, wenn wir unsere Augen auf ein Objekt richten. Mehrere Prozesse sind aktiv bevor unser Blick das neue Objekt erreicht. Diese Arbeit erforscht die räumlichen und zeitlichen Eigenschaften drei solcher Prozesse: 1. aufmerksamkeitsbedingte Steigerung der neuronalen Aktivität und sakkadische Suppression; 2. aufmerksamkeitsbasierte Auswahl des Zielreizes bei einer visuellen Suchaufgabe; 3. zeitliche Entwicklung der Detektiongenauigkeit bei der Objekt-Substitutionsmaskierung. Wir untersuchten diese Prozesse mit einer Kombination aus humaner Elektroenzephalografie (EEG), eye tracking und psychophysischen Verhaltensmessungen. Zuerst untersuchten wir, wie die neuronale Repräsentation eines Reizes von seiner zeitlichen Nähe zur Sakkade geprägt wird. Wir zeigten, dass direkt vor der Sakkade erscheinende Reize am meisten durch Aufmerksamkeit und Suppression geprägt sind. In Studie 2 wurde die Sichtbarkeit des Reizes mit der Objekt-Substitutionsmaskierung verringert, und wir analysierten das Verhältnis zwischen sakkadischen Reaktionszeiten und ihrer Genauigkeit. Dazu erfassten wir neuronale Marker der Aufmerksamkeitslenkung zum Zielreiz und eine subjektive Bewertung seiner Wahrnehmbarkeit. Wir stellten fest, dass schnelle Sakkaden der Maskierung entgingen und Genauigkeit sowie subjektive Wahrnehmbarkeit erhöhten. Dies zeigt, dass bereits in frühen Verarbeitungsstadien eine bewusste und korrekte Wahrnehmung des Reizes entstehen kann. Wir replizierten diesen Befund für manuelle Antworten, um eine Verfälschung der Ergebnisse durch sakkadenspezifische Prozesse auszuschließen. Neben ihrer theoretischen Bedeutung liefern diese Studien einen methodischen Beitrag zum Forschungsgebiet der EEG-Augenbewegung: Entfernung sakkadischer Artefakte aus dem EEG bzw. Erstellung eines künstlichen Vergleichsdatensatzes. Die Arbeit stellt mehrere Ansätze zur Untersuchung der Dynamik visueller Wahrnehmung sowie Lösungen für zukünftige Studien dar. / The visual system achieves a tremendous amount of processing as soon as we set eyes on a new object. Numerous processes are active already before eyes reach the object. This thesis explores the spatio-temporal properties of three such processes: attentional enhancement and saccadic suppression that accompany saccades to target; attentional selection of target in a visual search task; the timecourse of target detection accuracy under object-substitution masking. We monitored these events using a combination of human electrophysiology (EEG), eye tracking and behavioral psychophysics. We first studied how the neural representation of a visual stimulus is affected by its temporal proximity to saccade onset. We show that stimuli immediately preceding a saccade show strongest effects of attentional enhancement and saccadic suppression. Second, using object-substitution masking to reduce visibility, we analyzed the relationship between saccadic reaction times and response accuracy. We also collected subjective visibility ratings and observed neural markers of attentional selection, such as the negative, posterior-contralateral deflection at 200 ms (N2pc). We found that fast saccades escaped the effects of masking, resulted in higher response accuracy and higher awareness ratings. This indicates that early visual processing can trigger awareness and correct behavior. Finally, we replicated this finding with manual responses. Discovering a similar accuracy timecourse in a different modality ruled out saccade-specific mechanisms, such as saccadic suppression and retinal shift, as a potential confound. Next to their theoretical impact, all studies make a methodological contribution to EEG-eye movement research, such as removal of large-scale saccadic artifacts from EEG data and composition of matched surrogate data. In sum, this work uses multiple approaches to describe the dynamics of visual perisaccadic perception and offers solutions for future studies in this field.

Sakkadische Blickbewegungen sind die häufigsten und schnellsten aller menschlichen Bewegungen und führen zur wiederholtem und rapiden Verschiebung von Objektprojektionen über die Retina. Entgegen der verbreiteten Annahme der Suppression untersucht diese Arbeit Ausmaß und Funktion intrasakkadischer visueller Wahrnehmung. Studie I beschreibt eine individuell gefertigte LED-Installation zur ausschließlich intrasakkadischen Präsentation von Text und Bildern, während Studie II einen Algorithmus zur Detektion von Sakkaden vorstellt, welcher blickkontingente Stimulusmanipulationen mithilfe eines DLP Projektionssystems mit einer Bildwiederholungsrate von 1440 Hz ermöglicht. Studien III und IV untersuchten ob visuelle Bewegungsspuren (sog. motion streaks), welche durch die schnelle Bewegung von Objekten über die Retina erzeugt werden, Korrespondenz zwischen Objekten über Sakkaden hinweg herstellen könnten. Diese Bewegungsspuren erlaubten Versuchsteilnehmern nicht nur einen präsakkadischen Stimulus aus zwei identischen postsakkadischen Stimuli zu identifizieren, während diese Fähigkeit von der Deutlichkeit der Bewegungsspur abhing, sondern auch Korrektursakkaden zu einem ursprünglichen präsakkadischen Stimulus zu erleichtern, falls dieser während der Sakkade versetzt wurde. Studie V untersuchte die subjektive Wahrnehmung und Lokalisierung von intrasakkadischen Bewegungsspuren, indem Teilnehmer gezeichnete Berichte angaben. Die Modellierung letzterer ergab, dass retinale Positionssignale mit einer zeitlich gedämpften mentalen Repräsentation von Augenposition kombiniert wurden, um eine Lokalisation in weltzentrierten Koordinaten zu ermöglichen. Diese Ergebnisse legen nahe, dass intrasakkadische visuelle Signale einen Einfluss auf transsakkadische perzeptuelle und motorische Prozesse haben könnten. Letztlich werden die mögliche Funktionen intrasakkadischer Wahrnehmung, sowie Möglichkeiten für zukünftige wissenschaftliche Untersuchungen, diskutiert. / Rapid eye movements, so-called saccades, are the fastest and most frequent human movements and cause projections of objects in the world to constantly shift across the retina at high velocities, thereby producing large amounts of motion blur. In contrast to accounts of saccadic suppression, this work explores the extent and potential functional role of intra-saccadic perception. As saccades are fast and brief events, technical challenges were addressed. Study I describes a custom LED-based anorthoscopic presentation setup capable of displaying text and images strictly during saccades. In study II, a novel online saccade detection algorithm enabled rapid, gaze-contingent display changes using a DLP projection system running at 1440 fps. Studies III and IV investigated whether intra-saccadic motion streaks, i.e., blurred traces routinely induced by stimuli moving at saccadic speeds, could serve as cues to establishing object correspondence across saccades. Motion streaks not only enabled perceptual matching of pre- and post-saccadic object locations, while performance depended strongly on streak efficiency, but also facilitated gaze correction in response to intra-saccadic target displacements, that was previously found to be mainly driven by objects’ surface features. Finally, study V explored the subjective appearance and localization of intra-saccadic motion streaks, tasking observers to reproduce their trajectories. Computational modeling of resulting response patterns suggested that retinal positions over time were combined with a damped eye position signal to readily localize intra-saccadic input in world-centered coordinates. Taken together, these results invite the intriguing hypothesis that intra-saccadic visual signals are not discarded from processing and might affect trans-saccadic perceptual and motor processes. The potential role of intra-saccadic perception for active vision, as well as directions for future research, are discussed.

The understanding of the subjective experience of a visually stable world despite the occurrence of an observer's eye movements has been the focus of extensive research for over 20 years. These studies have revealed fundamental mechanisms such as anticipatory receptive field (RF) shifts and the saccadic suppression of stimulus displacements, yet there currently exists no single explanatory framework for these observations. We show that a previously presented neuro-computational model of peri-saccadic mislocalization accounts for the phenomenon of predictive remapping and for the observation of saccadic suppression of displacement (SSD). This converging evidence allows us to identify the potential ingredients of perceptual stability that generalize beyond different data sets in a formal physiology-based model. In particular we propose that predictive remapping stabilizes the visual world across saccades by introducing a feedback loop and, as an emergent result, small displacements of stimuli are not noticed by the visual system. The model provides a link from neural dynamics, to neural mechanism and finally to behavior, and thus offers a testable comprehensive framework of visual stability.