COLOR AND FORM SIGNALS IN THE HUMAN BRAIN

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1 DISS. ETH NO COLOR AND FORM SIGNALS IN THE HUMAN BRAIN A dissertation submitted to ETH ZURICH for the degree of Doctor of Sciences presented by ILIAS RENTZEPERIS M.Sc. in Information Networking, Carnegie Mellon University born on September, 13, 1980 citizen of Greece accepted on the recommendation of Prof. Dr. Kevan A.C. Martin, examiner P.D. Dr. Daniel Kiper, co-examiner Dr. Andrey Nikolaev, co-examiner 2012

2 Abstract Everything that surrounds us and forms our reality does not appear as disjoint pieces of information. On the contrary, we incorporate different attributes of a visual scene such as motion, color and form into a unitary, coherent percept. Influential early studies on visual processing have suggested that our experience of the world as a whole is generated by modular units in the brain that process specific information. This proposal has been challenged by studies that indicated that visual cortical mechanisms process several features simultaneously. We are far from knowing the way our brain binds all these different features into a unified representation of reality even though experimental evidence has given rise to several propositions. The work presented here examines how our visual brain processes color and form information. Human visual cortex contains mechanisms that pool local orientation information over large areas of visual space to support percepts of global form. In a psychophysical study we used Glass patterns and the phenomenon of adaptation to determine whether form perception is mediated by mechanisms that are color and/or luminance selective. Glass patterns are made by superimposing two identical random dot fields after geometrically transforming one of them. The sparse local cues elicit the perception of global patterns. Depending on the geometrical rule used, different global form percepts can arise. Subjects were adapted to either a radial or concentric Glass pattern of a given color or luminance polarity. We measured the effect of adaptation on subsequent detection of Glass patterns with the same or different visual attributes. Our results show that adapting to a concentric or radial pattern significantly elevates thresholds for the subsequent detection of patterns of the same form, irrespective of their color or luminance polarity. We conclude that Glass pattern perception is mediated by perceptual mechanisms that are color and luminance invariant. Even though our psychophysical results strongly suggest that form adaptation is color invariant we cannot say with certainty at which stage of visual processing this happens. To address this problem and determine the time course and interaction of color and form adaptation processes we analyzed the event-related potentials (ERP) evoked by the presentation of colored Glass patterns in an adaptation paradigm similar to the one in our psychophysical study. We found that the form adaptation effect evoked by the ERP waveforms manifests itself late, 220 ms after stimulus onset, right after the N1 component, and is also color invariant. We further analyzed the data from the ERP experiment to study the relationship between adaptation and response selectivities. We consider response selectivity to be I

3 the brain activity in response to a specific property of a stimulus. We found that response selectivities to form and, to some extent, color of the patterns were reflected in the C1 and N1 ERP components. Thus adaptation and response selectivities to form were manifested in non-overlapping intervals. Since adaptation to form makes form perception harder, we hypothesize that it could induce the employment of feedback mechanisms by the visual brain and operate at a different time scale than uninhibited form processing. In further psychophysical studies we examined the contrast detection and orientation discrimination sensitivities of peripherally located red-green, blue-yellow and luminance Gabor patterns. Analysis of the contrast detection experiments revealed increased sensitivity to red-green Gabor patterns that were oriented radially to the fixation point. This effect was less prominent for luminance Gabors and nonexistent for blue-yellow ones. Furthermore we found distinct sensitivity distributions across the visual field for the three different Gabor types. Our results indicate that at the level of contrast detection of local orientation cues the two color systems and the luminance one are processed by functionally distinct mechanisms. II

4 Zusammenfassung Nichts was uns umgibt und unsere Realität formt erscheint uns als separierte Bruchstücke von Information. Vielmehr integrieren wir verschiedene Attribute einer visuellen Szene, wie zum Beispiel Bewegung, Farbe und Form, zu einer einheitlichen und kohärenten Empfindung. Die ersten massgebenden Studien über visuelle Verarbeitung deuteten darauf hin, dass unser Erleben der äusseren Welt aus bausteinförmigen Einheiten des Gehirns, die spezifische Informationen verarbeiten, erzeugt wird. Dieser Vorschlag ist von Studien angefochten worden, die darauf hinweisen, dass visuelle kortikale Mechanismen mehrere Eigenschaften simultan verarbeiten. Wir sind weit entfernt vom Verständnis wie unser Gehirn alle diese Eigenschaften zu einer vereinheitlichten Representation der Realität zusammenfügt, obwohl experimentelle Evidenz zu mehreren Vorschlägen geführt hat. Die hier vorgelegte Arbeit untersucht wie unser visuelles Gehirn Informationen zu Farbe und Form verarbeitet. Der menschliche visuelle Kortex beinhaltet Mechanismen die lokale Information zu Orientation über grosse Regionen des visuellen Bereichs vereinigt, um die Wahrnehmung globaler Form zu stützen. Wir haben Glass Muster und das Phenomen der Adaptierung in einer psychophysikalischen Untersuchung verwendet, um zu bestimmen ob die Formwahrnehmung durch Mechanismen, die selektiv für Farbe und/oder Luminanz sind, vermittelt wird. Glass Muster werden erzeugt indem zwei identische zufällige Punktfelder überlagert werden, nachdem eines davon geometrisch transformiert wurde. Diese wenigen lokale Signale lösen die Wahrnehmung globaler Muster aus. Abhängend von den benützten geometrischen Regeln können verschiedene globale Formwahrnehmungen aufkommen. Die Testpersonen wurden entweder an ein radiales oder ein konzentrisches Glass Muster gegebener Farbe oder Luminanzpolarität adaptiert. Wir haben den Einfluss der Adaptierung auf die anschliessende Erkennung von Glass Mustern derselben oder anderer visueller Attribute gemessen. Unsere Resultate zeigen dass die Adaptierung an ein konzentrisches oder radiales Muster die Schwellenwerte für die anschliessende Mustererkennung derselben Form anheben, unabhängig von deren Farbe oder Luminanzpolarität. Wir schlussfolgern, dass die Perzeption von Glass Mustern durch Wahrnehmungsmechanismen vermittelt wird, die form- und luminanzinvariant sind. Obwohl unsere psychophysischen Resultate stark darauf hinweisen, dass Formadaption ein farbunabhängiger Vorgang ist, können wir nicht mit Sicherheit eingrenzen in welchem Stadium der visuellen Verarbeitung die Formadaption stattfindet. Um diese Problemstellung zu beantworten und um den zeitlichen Verlauf der Farb- und Form-Adaptionsprozesse zu ermitteln analysierten wir die III

5 sogenannten Event-related-potentials (ERP), welche durch die Präsentation von farbigen Glasrastern, im Zuge eines unserer Adaptationsstudie ähnlichen Adaptionsparadigmas, hervorgerufen werden. Wir fanden heraus, dass sich der Formadaptationseffekt, welcher durch einen ERP Schwingungsverlauf hervorgerufen wird, als late, 220 ms nach dem Stimulusbeginn, unmittelbar nach der N1 Komponente, manifestiert und farbunabhängig ist. Weitere Analysen der Daten des ERP Experiments, befassten sich mit der Beziehung zwischen Adaptation und Response-Selektivität (Response-Trennschärfe). Wir betrachten die Response-Selektivität (Response-Trennschärfe) als eine Hirnaktivität hervorgerufen durch eine spezifische Stimuluseigenschaft. Unsere Untersuchungen ergaben, das Response-Selektivitäten für die Form und, in einem gewissen Masse, auch für die Farbe der Raster, in den C1 und N1 ERP- Komponenten reflektiert werden. Demnach manifestieren sich Adaption an und Response-Selektivität für Formwahrnehmung in sich nicht überschneidenden Intervallen. Nachdem Formadaptation die Formerkennung erschwert nehmen wir an, dass sie zur Anwendung von Feedbackmechanismen des visuellen Hirns führt und in einem Zeitrahmen operiert, welcher nicht von der Verarbeitung inhibiert wird. In weiteren psychophysikalischen Versuchen untersuchten wir die Sensibilität von Kontrasterkennung und Orientierungsunterscheidung unter peripher gelegenen rot-grün, blau-gelb und luminanten Gabor Mustern. Kontrasterkennungs-Experimente zeigten erhöhte Empfindlichkeit gegenüber rot-grün Gabor Mustern mit radialer Orientierung zum Fixationspunkt. Dieser Effekt war weniger markant für luminante Gabor Muster und nicht vorhanden bei blau-gelber Anordnung. Desweiteren fanden wir für die drei unterschiedlichen Gabor Typen eindeutige Empfindlichkeitsverteilungen im visuellen Feld. Unsere Ergebnisse deuten darauf hin, dass auf der Ebene von Kontrasterkennung lokale Orientierung das 2- Farben-System koodiniert und Luminanz von funktionell unterschiedlichen Mechanismen gesteuert wird. IV

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