Kognitive Interpretationen mehrdeutiger visueller Reize

Unser Gehirn muss zu jeder Zeit relevante Signale von irrelevanten Informationen trennen. Dazu müssen diese als spezifische Einheiten erkannt und klassifiziert werden. Mehrdeutigkeit ist ein wesentlicher Aspekt dieses Verarbeitungsprozesses und kann durch verrauschte Eingangssignale und durch den Aufbau unserer sensorischer Systeme entstehen. Beispielsweise können Reize mehrdeutig sein, wenn sie verrauscht oder unvollständig sind oder nur kurzzeitig wahrgenommen werden. Unter solchen Bedingungen werden Wahrnehmung und Klassifikation eines Reizes deutlich erschwert. Bereits vorhandene kognitive Repräsentationen werden somit möglicherweise nicht aktiviert. Folglich müssen Rückschlüsse über die Reize aufgrund von Kontext und Erfahrung gezogen werden. Ein und derselbe Reiz kann jedoch unterschiedlich repräsentiert und im sensorischen System kodiert werden. Da nur eine Repräsentation die Basis zukünftigen Handelns bilden kann, entsteht eine Art Konkurrenz innerhalb der Wahrnehmung. Derartige Wahrnehmungsphänomene, die mit der Mehrdeutigkeit von Reizen in Verbindung stehen, bilden den Mittelpunkt der vorliegenden Dissertation. Wenn einem physikalisch konstanten Reiz mehrere Interpretationen zugeordnet werden, entsteht ein Wechsel zwischen diesen Einordnungen, den man wahrnimmt und Rivalität ("rivalry") nennt. In dieser Dissertation werden diverse neue Erkenntnisse zu diesem grundlegenden Phänomen der sensorischen Verarbeitung beschrieben. So wird gezeigt, dass Übergänge zwischen drei wahrgenommenen Interpretationen – ein vergleichsweise selten untersuchtes Phänomen, da Rivalität meist mit zweideutigen Reizen untersucht wird – vorhersehbaren Mustern folgen (Kapitel 2). Darüber hinaus zeigt sich, dass derartige Übergänge spezifische Eigenschaften aufweisen, welche die Geschwindigkeit und die Richtung ihrer räumlichen Ausbreitung im visuellen Feld bestimmen (Kapitel 3). Diese Eigenschaften der Mehrdeutigkeit werden weiterhin stark von Aufmerksamkeit und anderen, introspektiven Prozessen beeinflusst. Um die der Rivalität in der Wahrnehmung tatsächlich zugrundeliegenden Prozesse und die damit verbundenen Änderungen des Bewusstseins von derartigen subjektiven Prozessen abzugrenzen, müssen letztere kontrolliert oder sogar vollständig umgangen werden. Ein objektives Maß der Rivalität in der Wahrnehmung wird zur Lösung dieser Aufgabe vorgeschlagen und bietet eine wertvolle Alternative zu introspektivem Berichten über den Wahrnehmungszustand (Kapitel 4). Übergänge in der Wahrnehmung entstehen entlang einer bestimmten Merkmalsdimension des Reizes, wie beispielsweise der Orientierung des berühmten Neckerwürfels. Zudem kann auch eine Änderung in der Merkmalsdimension der Luminanz eine unterschiedliche Interpretation des Reizes hervorrufen. Es wird gezeigt, dass die Pupille kleiner wird, wenn eine Interpretation mit hoher Luminanz die Wahrnehmung übernimmt, und umgekehrt, dass die Pupille größer wird, wenn eine Interpretation mit niedriger Luminanz die Wahrnehmung übernimmt. Folglich kann die Pupille als ein zuverlässiges und objektives Maß für Änderungen in der Wahrnehmung verwendet werden. Durch die Verwendung solcher objektiven Maße konnten neue Eigenschaften der Übergänge in der Wahrnehmung aufgezeigt werden, welche die Theorie unterstützen, dass Introspektion die der Verarbeitung mehrdeutiger Situationen zugrundeliegenden Prozesse merklich beeinflussen kann. Als Nächstes wurden mehrdeutiger Reize im Zusammenhang mit der Wahrnehmung von Objekten eingesetzt (Kapitel 5). Am Beispiel der Kippfigur des "bewegten Diamanten" wird dabei die Bedeutung von mehrdeutigen Reizen veranschaulicht. Beim bewegten Diamanten werden zwei Interpretationen wahrgenommen, die sich entlang der Dimension der Objektkohärenz abwechseln. Das bedeutet, dass die Wahrnehmung zwischen einem einzelnen zusammenhängenden Objekt (Diamant) und mehreren unzusammenhängenden Komponenten kippt. Es wird gezeigt, dass die Interpretation des Reizes als ein einziges kohärentes Objekt, verglichen mit der Interpretation als mehrere Komponenten, zu einer Erhöhung der visuellen Empfindlichkeit innerhalb des Objektes führt. Diese Ergebnisse sind ein Beleg dafür, wie die Aktivierung einer Interpretation eines Reizes als Einzelobjekt (im Vergleich zur Komponentenwahrnehmung) dazu führt, dass die Aufmerksamkeit top-down zu den relevanten Bereichen des Gesichtsfeldes gelenkt wird. Es wird weiter untersucht, welche Eigenschaften des Reizes zu einer bottom-up Aktivierung der Interpretation solcher Objekte beitragen (Kapitel 6). Die Mehrdeutigkeit von Objekten kann erfolgreich aufgehoben werden, indem man einen starken Kontrast in Luminanz oder Farbe zwischen dem Objekt und dem Hintergrund erzeugt. Auch die Größe und die Form haben einen großen Einfluss auf die Detektion und Identifikation von Objekten. Des Weiteren sind die Eigenschaften eines Objektes nicht nur bestimmend für die Erfolgsquote bei der Objekterkennung, sondern ebenso bedeutend für die Speicherung der Repräsentation im Gedächtnis, beispielsweise von neu wahrgenommenen Objekten. Das Klassifizieren von Objekten durch die Versuchsperson wird ebenfalls durch Mehrdeutigkeit beeinflusst. So kann ein Objekt der Versuchsperson einerseits als neu erscheinen, obwohl es bereits bekannt war, weil es beispielsweise der Versuchsperson schon einmal gezeigt worden ist. Andererseits kann auch ein eigentlich unbekanntes Objekt der Versuchsperson dennoch vertraut vorkommen. In dieser Arbeit wird gezeigt, dass solche subjektiven Effekte einen Einfluss auf die Pupillengröße haben (Kapitel 7). Außerdem verkleinert sich die Pupille der Versuchspersonen beim Betrachten neuer Bilder stärker als bei bekannten. Ein ähnlicher Effekt wird gefunden, wenn das Bild vorher erfolgreich im Gedächtnis gespeichert wurde. Daher ist es wahrscheinlich, dass die Pupille die Verfestigung von neuen Objekten im Gedächtnis widerspiegelt. Abschließend wird untersucht, ob sich kognitive Prozesse, wie Entscheidungsfindung – ein wichtiger Prozess, falls mehreren Optionen zur Verfügung stehen und Mehrdeutigkeit aufgehoben werden soll – auch in der Pupille widerspiegeln (Kapitel 8). Es wird zunächst bestätigt, dass die Pupillen sich erweitern, nachdem man eine Entscheidung getroffen hat. Neu wird gezeigt, dass diese Pupillenausdehnungen erfolgreich von anderen Personen erkannt und verwendet werden können, um ein interaktives Spiel gegen die erste Person (den "Gegner") zu gewinnen. Insgesamt wird in dieser Dissertation untersucht, wie mehrdeutige Reize die Wahrnehmung beeinflussen und wie Mehrdeutigkeit verwendet werden kann, um Prozesse des Gehirns zu studieren. Es hat sich gezeigt, dass Mehrdeutigkeit vorhersehbaren Mustern folgt, sie objektiv mit Reflexen gemessen werden kann, und Einblicke in neuronale Prozesse wie Aufmerksamkeit, Objektwahrnehmung und Entscheidungsmechanismen liefern kann. Diese Ergebnisse zeigen, dass Mehrdeutigkeit eine zentrale Eigenschaft sensorischer Systeme ist, und Lebewesen in die Lage versetzt, mit ihrer Umwelt flexibel zu interagieren. Mehrdeutigkeit macht das Verhalten vielfältiger, ermöglicht es dem Gehirn, mit der Welt auf verschiedenen Wegen zu interagieren, und ist die Basis der Dynamik von Wahrnehmung, Interpretation und Entscheidung.Brains can sense and distinguish signals from background noise in physical environments, and recognize and classify them as distinct entities. Ambiguity is an inherent part of this process. It is a cognitive property that is generated by the noisy character of the signals, and by the design of the sensory systems that process them. Stimuli can be ambiguous if they are noisy, incomplete, or only briefly sensed. Such conditions may make stimuli indistinguishable from others and thereby difficult to classify as single entities by our sensory systems. In these cases, stimuli fail to activate a representation that may have been previously stored in the system. Deduction, through context and experience, is consequently needed to reach a decision on what is exactly sensed. Deduction can, however, also be subject to ambiguity as stimuli and their properties may receive multiple representations in the sensory system. In such cases, these multiple representations compete for perceptual dominance, that is, for becoming the single entity taken by the system as a reference point for subsequent behavior. These types of ambiguity and several phenomena that relate to them are at the center of this dissertation. Perceptual rivalry, the phenomenal experience of alternating percepts over time, is an example of how the brain may give multiple interpretations to a stimulus that is physically constant. Rivalry is a very typical and general sensory process and this thesis demonstrates some newly discovered properties of its dynamics. It was found that alternations between three perceptual interpretations – a relatively rare condition as rivalry generally occurs between two percepts – follow predictable courses (Chapter 2). Furthermore, such alternations had several properties that determine their speed and direction of spatial spread (suppression waves) in the visual field (Chapter 3). These properties of ambiguity were further strongly affected by attention and other introspective processes. To demarcate the true underlying process of perceptual rivalry and the accompanied changes in awareness, these subjective processes need to be either circumvented or controlled for. An objective measure of perceptual rivalry was proposed that resolved this issue and provided a good alternative for introspective report of ambiguous states (Chapter 4). Changes in percepts occur along a specific feature domain such as depth orientation for the famous Necker cube. Alternatively, luminance may also be a rivalry feature and one percept may appear brighter as the other rivaling percept. It was demonstrated that the pupil gets smaller when a percept with high luminance becomes dominant, and vice versa, gets bigger when a percept with low luminance gets dominant during perceptual rivalry. As such, the pupil can serve as a reliable objective indicator of changes in visual awareness. By using such reflexes during rivalry, several new properties of alternations were discovered and it was again confirmed that introspection can confound the true processes involved in ambiguity. Next, the usefulness of ambiguous stimuli was explored in the context of objects as entities (Chapter 5). Some ambiguous stimuli can induce two percepts that alternate along the feature domain of object coherency, that is, whether a single coherent object or multiple incoherent objects are seen. In other words, an ambiguous stimulus can induce two cognitive interpretations of either seeing an entity or not. It was reported that being aware of a single coherent object results in the increase in visual sensitivity for the areas that constitute the object. These results are evidence of how the activation of a representation of a single and unique object can guide and allocate attentional resources to relevant areas in the visual field in a top-down way. It was further explored which features help to bottom-up access such object representations (Chapter 6). Ambiguity of objects can be successfully resolved by adding strong contrasts between the object and its background in luminance and color. The size and variability of the object's shape was also found to be an important factor for its successful detection and identification. Furthermore, the characteristics of objects do not only determine the rate of success in a recognition task, but are equally important for the storage of their representations in memory if, for instance, the object is novel to the observer. The subjective experience of a novel object is also subject to ambiguity and objects may appear novel to the observer although they are familiar (i.e., previously shown to the observer), or vice versa, they appear familiar to the observer although they are actually novel. It was here shown that such subjective effects are reflected in the pupil (Chapter 7). In addition, if novel images were presented to observers, their pupils constricted stronger as compared to if familiar images were presented. Similarly, if novel stimuli were shown to observers, pupillary constrictions were stronger if these stimuli were successfully stored in memory as compared to those later forgotten. As such, the pupil reflected the cognitive process of novelty encoding. Finally, it was tested whether other cognitive processes, such as decision-making – an important process when multiple options are available and ambiguity has to be resolved with a conscious decision – were also reflected in changes of pupil size (Chapter 8). It was confirmed that the pupil tends to dilate after an observer has made a decision. These dilations can successfully be detected between individuals and further used to gain the upper hand during an interactive game. In sum, this thesis has explored how ambiguous signals affect perception and how ambiguity inside perceptual systems can be used to study processes of the brain. It is found that ambiguity follows predictable courses, can be objectively assessed with reflexes, and can provide insights into other neuronal mechanisms such as attention, object representations, and decision-making. These findings demonstrate that ambiguity is a core property of the sensory systems that enable living beings to interact with their surroundings. Ambiguity adds variation to behavior, allows the brain to flexibly interact with the world, and lies at the bottom of the dynamics of sense, interpretations, and behavioral decisions

Pupil size signals novelty and predicts later retrieval success for declarative memories of natural scenes

Declarative memories of personal experiences are a key factor in defining oneself as an individual, which becomes particularly evident when this capability is impaired. Assessing the physiological mechanisms of human declarative memory is typically restricted to patients with specific lesions and requires invasive brain access or functional imaging. We investigated whether the pupil, an accessible physiological measure, can be utilized to probe memories for complex natural visual scenes. During memory encoding, scenes that were later remembered elicited a stronger pupil constriction compared to scenes that were later forgotten. Thus, pupil size predicts success or failure of memory formation. In contrast, novel scenes elicited stronger pupil constriction than familiar scenes during retrieval. When viewing previously memorized scenes, those that were forgotten (misjudged as novel) still elicited stronger pupil constrictions than those correctly judged as familiar. Furthermore, pupil constriction was influenced more strongly if images were judged with high confidence. Thus, we propose that pupil constriction can serve as a marker of novelty. Since stimulus novelty modulates the efficacy of memory formation, our pupil measurements during learning indicate that the later forgotten images were perceived as less novel than the later remembered pictures. Taken together, our data provide evidence that pupil constriction is a physiological correlate of a neural novelty signal during formation and retrieval of declarative memories for complex, natural scenes

Pupil size signals novelty and predicts later retrieval success for declarative memories of natural scenes

Declarative memories of personal experiences are a key factor in defining oneself as an individual, which becomes particularly evident when this capability is impaired. Assessing the physiological mechanisms of human declarative memory is typically restricted to patients with specific lesions and requires invasive brain access or functional imaging. We investigated whether the pupil, an accessible physiological measure, can be utilized to probe memories for complex natural visual scenes. During memory encoding, scenes that were later remembered elicited a stronger pupil constriction compared to scenes that were later forgotten. Thus, pupil size predicts success or failure of memory formation. In contrast, novel scenes elicited stronger pupil constriction than familiar scenes during retrieval. When viewing previously memorized scenes, those that were forgotten (misjudged as novel) still elicited stronger pupil constrictions than those correctly judged as familiar. Furthermore, pupil constriction was influenced more strongly if images were judged with high confidence. Thus, we propose that pupil constriction can serve as a marker of novelty. Since stimulus novelty modulates the efficacy of memory formation, our pupil measurements during learning indicate that the later forgotten images were perceived as less novel than the later remembered pictures. Taken together, our data provide evidence that pupil constriction is a physiological correlate of a neural novelty signal during formation and retrieval of declarative memories for complex, natural scenes

Gaze-contingent flicker pupil perimetry detects scotomas in patients with cerebral visual impairments or glaucoma

The pupillary light reflex is weaker for stimuli presented inside as compared to outside absolute scotomas. Pupillograph perimetry could thus be an objective measure of impaired visual processing. However, the diagnostic accuracy in detecting scotomas has remained unclear. We quantitatively investigated the accuracy of a novel form of pupil perimetry. The new perimetry method, termed gaze-contingent flicker pupil perimetry, consists of the repetitive on, and off flickering of a bright disk (2 hz; 320 cd/m; 4° diameter) on a gray background (160 cd/m) for 4 seconds per stimulus location. The disk evokes continuous pupil oscillations at the same rate as its flicker frequency, and the oscillatory power of the pupil reflects visual sensitivity. We monocularly presented the disk at a total of 80 locations in the central visual field (max. 15°). The location of the flickering disk moved along with gaze to reduce confounds of eye movements (gaze-contingent paradigm). The test lasted ~5 min per eye and was performed on 7 patients with cerebral visual impairment (CVI), 8 patients with primary open angle glaucoma (age >45), and 14 healthy, age/gender-matched controls. For all patients, pupil oscillation power (FFT based response amplitude to flicker) was significantly weaker when the flickering disk was presented in the impaired as compared to the intact visual field (CVI: 12%, AUC = 0.73; glaucoma: 9%, AUC = 0.63). Differences in power values between impaired and intact visual fields of patients were larger than differences in power values at corresponding locations in the visual fields of the healthy control group (CVI: AUC = 0.95; glaucoma: AUC = 0.87). Pupil sensitivity maps highlighted large field scotomas and indicated the type of visual field defect (VFD) as initially diagnosed with standard automated perimetry (SAP) fairly accurately in CVI patients but less accurately in glaucoma patients. We provide the first quantitative and objective evidence of flicker pupil perimetry's potential in detecting CVI-and glaucoma-induced VFDs. Gaze-contingent flicker pupil perimetry is a useful form of objective perimetry and results suggest it can be used to assess large VFDs with young CVI patients whom are unable to perform SAP

Pupil Mimicry is the Result of Brightness Perception of the Iris and Pupil

Recent scientific investigations suggest that people automatically mimic each other’s pupil sizes during interaction. However, instead of being a social mimicry effect, it could also be the result of brightness perception. When observers look at individuals with dilated pupils, little of the brighter iris is visible, leading to the perception of a relatively low-illuminated eye region. In the current study we tested whether pupil mimicry remains present when pupils and irises are equalized for luminance values across pupil sizes. We tested several stimulus sets, including faces with static pupils that varied in size across images and dynamic pupils that changed in size over time in videos. Results showed that for traditional, not-luminance-equalized videos, participants’ pupil sizes adapted to the observed pupils, showing a pattern that is roughly in line with pupil mimicry. However, no such pupil response in line with mimicry was seen for static images (regardless of whether they were equalized for luminance) nor for luminance-equalized videos. These findings suggest that only salient, dynamic stimuli attract enough attention to the luminance in the eye region to evoke a pupillary response. However, although such responses suggest pupil mimicry, the underlying factor is the change in brightness within the eye as a function of pupil size rather than social mimicry

Predicting Product Preferences on Retailers' Web Shops through Measurement of Gaze and Pupil Size Dynamics

Previous studies used gaze behavior to predict product preference in value-based decision-making, based on gaze angle variables such as dwell time, fixation duration and the first fixated product. While the application for online retail seems obvious, research with realistic web shop stimuli has been lacking so far. Here, we studied the decision process for 60 Dutch web shops of a variety of retailers, by measuring eye movements and pupil size during the viewing of web shop images. The outcomes of an ordinal linear regression model showed that a combination of gaze angle variables accurately predicted product choice, with the total dwell time being the most predictive gaze dynamic. Although pupillometric analysis showed a positive relationship between pupil dilation and product preference, adding pupil size to the model only slightly improved the prediction accuracy. The current study holds the potential to substantially improve retargeting mechanisms in online marketing based on consumers' gaze information. Also, gaze-based product preference proves to be a valuable metric in pre-testing product introductions for market research and prevent product launches from failure

Effects of Natural Scene Inversion on Visual-evoked Brain Potentials and Pupillary Responses: A Matter of Effortful Processing of Unfamiliar Configurations

The inversion of a picture of a face hampers the accuracy and speed at which observers can perceptually process it. Event-related potentials and pupillary responses, successfully used as biomarkers of face inversion in the past, suggest that the perception of visual features, that are organized in an unfamiliar manner, recruits demanding additional processes. However, it remains unclear whether such inversion effects generalize beyond face stimuli and whether indeed more mental effort is needed to process inverted images. Here we aimed to study the effects of natural scene inversion on visual evoked potentials and pupil dilations. We simultaneously measured responses of 47 human participants to presentations of images showing upright or inverted natural scenes. For inverted scenes, we observed relatively stronger occipito-temporo-parietal N1 peak amplitudes and larger pupil dilations (on top of an initial orienting response) than for upright scenes. This study revealed neural and physiological markers of natural scene inversion that are in line with inversion effects of other stimulus types and demonstrates the robustness and generalizability of the phenomenon that unfamiliar configurations of visual content require increased processing effort

Reaction time coupling in a joint stimulus-response task: A matter of functional actions or likable agents?

Shaping one owns actions by observing others’ actions is driven by the deep-rooted mechanism of perception-action coupling. It typically occurs automatically, expressed as for example the unintentional synchronization of reaction times in interactive games. Theories on perception-action coupling highlight its benefits such as the joint coordination of actions to cooperatively perform tasks properly, the learning of novel actions from others, and the bonding with likable others. However, such functional aspects and how they shape perception-action coupling have never been compared quantitatively. Here we tested a total of hundred-fifteen participants that played a stimulus-response task while, in parallel, they observed videos of agents that played the exact same task several milliseconds in advance. We compared to what degree the reaction times of actions of agents, who varied their behavior in terms of functionality and likability in preceding prisoner dilemma games and quizzes, shape the reaction times of human test participants. To manipulate functionality and likability, we varied the predictability of cooperative behavior and correctness of actions of agents, respectively, resulting in likable (cooperative), dislikable (uncooperative), functional (correct actions), and dysfunctional (incorrect actions) agents. The results of three experiments showed that the participants’ reaction times correlated most with the reaction times of agents that expressed functional behavior. However, the likability of agents had no effects on reaction time correlations. These findings suggest that, at least in the current computer task, participants are more likely to adopt the timing of actions from people that perform correct actions than from people that they like

Maintaining fixation by children in a virtual reality version of pupil perimetry

The assessment of visual field sensitivities in young children continues to be a challenge. Children often do not sit still, fail to fixate stimuli for longer durations, and have limited verbal capacity to report visibility. We investigated the use of a head-mounted VR display, gaze-contingent flicker pupil perimetry (gcFPP), and three fixation stimulus conditions to determine best practices for optimal fixation and pupil response quality. A total of twenty children (3-11y) passively fixated a dot, counted the repeated appearance of an animated character, and watched an animated movie in separate trials of 80s each. We presented large flickering patches at different eccentricities and angles in the periphery to evoke pupillary oscillations (20 locations, 4s per location). The results showed that gaze precision and accuracy did not differ significantly across the fixation conditions but pupil amplitudes were strongest for the dot and count task. We recommend the use of the fixation counting task for pupil perimetry because children enjoyed it the most and it achieved strongest pupil responses. The VR set-up appears to be an ideal apparatus for children to allow free range of movement, an engaging visual task, and reliable eye measurements

The pupil near response is short lasting and intact in virtual reality head mounted displays

The pupil of the eye constricts when moving focus from an object further away to an object closer by. This is called the pupil near response, which typically occurs together with accommodation and vergence responses. When immersed in virtual reality mediated through a head-mounted display, this triad is disrupted by the vergence-accommodation conflict. However, it is not yet clear if the disruption also affects the pupil near response. Two experiments were performed to assess this. The first experiment had participants follow a target that first appeared at a far position and then moved to either a near position (far-to-near; FN) or to another far position (far-to-far; FF). The second experiment had participants follow a target that jumped between five positions, which was repeated at several distances. Experiment 1 showed a greater pupil constriction amplitude for FN trials, compared to FF trials, suggesting that the pupil near response is intact in head-mounted display mediated virtual reality. Experiment 2 did not find that average pupil dilation differed when fixating targets at different distances, suggesting that the pupil near response is transient and does not result in sustained pupil size changes