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Functional expression of recombinant N-methyl-D-aspartate (NMDA) receptors in eukaryotic cell lines
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Die vorliegende Dissertation berichtet über eine Serie von Verhaltens- und funktionellen Bildgebungsstudien zu experimentalpsychologischen Paradigmata, die eine räumliche Analyse und Koordinatentransformation von Material der visuellen Wahrnehmung oder Vorstellung beinhalten. Nach einer Einführung in die Prinzipien und Techniken der funktionellen Kernspintomographie, der hier benutzten Methode für die Messung von Gehirnaktivität, werden die Versuche einer Replikation des berühmten Stratton'schen Umkehrbrillen-Experiments dargestellt (Kapitel 1). Unsere vier Probanden zeigten zwar eine zügige Anpassung der visuomotorischen Funktionen an die neue visuelle Umwelt, berichteten aber, anders als Stratton, nicht, daß sie die Welt nach einigen Tagen mit der Umkehrbrille wieder normal sähen. Diese Persistenz des umgekehrten Bildes wurde durch eine psychphysische Testbatterie bestätigt. Des weiteren ergaben die funktionellen Kernspinmessungen, daß sich die kortikale retinotope Organisation im Verlaufe des Experiments nicht geändert hat. Da sich also Strattons Haupteffekt, das Aufrechtsehen durch die Umkehrbrille nach einwöchiger Adaptation, nicht replizieren ließ, werden andere Möglichkeiten der Interpretation der verschiedenen Umkehrexperimente der letzten hundert Jahre vorgeschlagen. Dieses Ergebnis einer funktionellen Anpassung ohne größere Änderungen der visuellen Wahrnehmung (und ohne Veränderungen der Repräsentation der Netzhautareale in der Sehrinde) führte zu der Hypothese, daß die erforderlichen Transformationen auf einer höheren Stufe der kortikalen Hierarchie der visuellen Verarbeitung erfolgen. Zur Testung dieser Hypothese wurde eine funktionelle Kernspinstudie des Umkehrlesens durchgeführt (Kapitel 2). Hierbei lasen die Probanden Wörter und Sätze in Spiegelschrift oder auf dem Kopf. Der neuronale Mechanismus der räumlichen Transformationen, die zur Bewältigung dieser Aufgabe nötig sind, konnte in bestimmten Regionen des Parietallappens, die zwischen den Probanden sehr konstant waren, lokalisiert werden. Weiterhin fand sich eine Koaktivierung okzipitootemporaler Objekterkennungs-Areale. Die Spezifizität der parietalen Aktivierungsfoci wurde durch ein Kontrollexperiment bestätigt, in welchem das kortikale System für räumliche Transformationen von den Netzwerken der allgemeinen visuellen Aufmerksamkeit und der Augenbewegungskontrolle unterschieden werden konnte. In einem weiteren Experiment wurden die räumlichen Funktionen des Parietallappens unter dem Vorzeichen der visuellen Vorstellung untersucht. Als Paradigma wurde der "mental clock" - Test verwendet, bei welchem die Probanden die Winkel der Zeiger zweier Uhren vergleichen müssen, deren Zeiten nur akustisch vorgegeben werden (Kapitel 3). Diese Aufgabe erfordert die Generierung eines entsprechenden Vorstellungsbildes und dessen räumliche Analyse, stellt also sowohl ein kontrolliertes Vorstellungs-Paradigma als auch einen Test räumlicher Funktionen dar, der nicht auf visuell präsentiertem Material beruht. Das parietale Aktivierungsmuster, das der Analyse der Winkel dieser vorgestellten Uhren zugeschrieben werden konnte, entsprach weitgehend demjenigen, das mit der räumlichen Transformation von Buchstaben verbunden war. Es handelt sich also wahrscheinlich um ein kortikales System für räumliche Analyse und Koordinatentransformationen, das nicht auf eine visuelle Stimulation angewiesen ist, sondern auch bei bloßer visueller Vorstellung aktiv werden kann. Die vorgelegten Resultate werden im Kontext neuerer neuropsychologischer Befunde zu Defiziten räumlicher Analyse und Vorstellung bei Läsionen des Parietallappens diskutiert (Kapitel 4). Auch die methodologischen Probleme der kognitiven Subtraktion, die in unseren Studien teilweise benutzt wurde, werden behandelt. Dabei wird erläutert, inwiefern diese für die Beurteilung der vorgelegten Studien nur von untergeordneter Bedeutung sind. Nichtsdestoweniger schlagen wir Modifikationen der experimentellen Paradigmata im Sinne des parametrischen Designs und des "event-related functional magnetic resonance imaging" vor, die bei zukünftigen Studien einen vollständigen Verzicht auf die kognitive Subtraktion ermöglichen dürften. Schließlich wird die Bedeutung der vorgelegten Ergebnisse für die Erforschung der Anpassungsfähigkeit des menschlichen Gehirns und des Verhältnisses von Vorstellung und visueller Wahrnehmung dargelegt.