Cognitive Interpretations of Ambiguous Visual Stimuli

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.