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Characteristics and Neuronal Correlates of Superior Memory Performance

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Characteristics and Neuronal Correlates of Superior Memory Performance Characteristics and Neuronal Correlates of Superior Memory Performance Inaugural-Dissertation zur Erlangung des Doktorgrades der Philosophie an der Ludwig-Maximilians-Universität München vorgelegt von Boris Nikolai Konrad aus Bochum München, Oktober 2013 1 Referentin: Prof. Dr. Kristina Hennig-Fast Koreferent: Prof. Dr. Patrick Khader Tag der mündlichen Prüfung: 24.01.2014 2 3 Acknowledgment It was several years ago that I first talked to Martin Dresler about the possibility of studying superior memorizers in order to learn more about their skill and memory in general. I was still a physics and computer science student at the time and had no idea that large parts of what we discussed would eventually end up in my PhD thesis. I would hereby like to express my deepest gratitude to everyone who supported, assisted and motivated me during the last approximately four years that it took me since starting at the Max Planck Institute of Psychiatry to finish this thesis, and I also extend my gratitude to everyone who contributed to the studies. In particular I would like to thank Michael Czisch for giving me the opportunity to undertake this work within his research group, Kristina Hennig-Fast for supervising my work and providing many helpful comments and ideas during the progress, Patrick Khader for co- supervising this thesis and Axel Steiger for assisting with the sleep-related parts of the study and having me in his research group in the last few months. Martin Dresler first had the idea to study memory athletes, encouraged me to join him and supported me on each step to realizing the various studies that eventuated from that initial idea. Thank you, Martin! I also would like to thank Lisa Genzel and Victor Spoormaker for their endless helpfulness and willingness to comment on all parts of my work, answering any and all questions that I had. This is also true for each and every person in the RGs Czisch and Steiger, including the technicians in both groups who provided a lot of help with equipment, measurements and sleep scoring. Thanks to all of you for your many helpful comments and encouraging words in group meetings, journal clubs and beyond. Thank you also to everyone who commented on prior versions of this thesis. 4 Hannah Nickl conducted the learning sessions and sleep parts of Study 3 and I would like to thank her for her support and her patience. Laurentius Halimsetiawan performed analysis of the sleep data in Study 1. I would also like to thank all of the subjects who joined me and were patient and flexible when needed. Over 150 in total (counting all pilots and controls) participated; some of them even came all the way from Sweden and Norway despite full schedules as full-time doctors or lawyers! Thank you! Besides work-related support I am also grateful for the support I received from many friends and of course my family, in particular my parents. Thanks to all of you for always being there when I needed someone to talk to. Finally I like to thank the financial sponsors: The Friedrich-Ebert-Stiftung (FES) who funded me with a PhD student scholarship and the Max Planck Institute of Psychiatry, who covered all of the study-related costs. 5 Zusammenfassung „Jedermann klagt über sein Gedächtnis, niemand über seinen Verstand.“ (François de La Rochefoucauld, ca. 1664) Viele Menschen fürchten sich vor Gedächtnisverlust im Alter und sind auch im Alltag mit ihrem Gedächtnis nicht zufrieden. Im Gegensatz dazu gibt es einzelne Personen die mit außergewöhnlich guten Gedächtnisleistungen imponieren. Sie zeigen ihre Fähigkeiten in Fernsehshows oder bei Gedächtnismeisterschaften und werden dafür oft bewundert. Schon im Mittelalter sind Gedächtniskünstler aufgetreten (Yates, 1966) und Wissenschaftlern (Valentine & Wilding, 1997) aufgefallen. Ihre Leistungen basieren zumeist auf Mnemotechnik genannten Gedächtnistechniken (Maguire, Valentine, Wilding, & Kapur, 2003), die schon seit der Antike bekannt sind (Hrees, 1986). Dennoch ist der Umfang der wissenschaftlichen Forschung über Gedächtnisverbesserung und Mnemotechniken im Vergleich zu Studien über Gedächtniserkrankungen noch stark unterrepräsentiert (Worthen & Hunt, 2011). Im Jahr 1990 wurde die erste Gedächtnisweltmeisterschaft ausgetragen. Seitdem hat sich der Gedächtnissport etabliert und es finden inzwischen weltweit entsprechende Meisterschaften statt (Wilding & Valentine, 1994). Erstmalig wurden vor rund zehn Jahren einige Gedächtnissportler auch mit Hilfe von bildgebenden Verfahren (funktionelle Magnetresonanztomographie, fMRT) untersucht (Maguire, Valentine, et al., 2003). Seitdem haben sich Teilnehmerzahlen und Rekorde (etwa das Merken der Reihenfolge eines gemischten Kartenspiels in 21,19 Sekunden) im Gedächtnissport rasant entwickelt (Konrad, 2013). In der ersten Studie wurden insgesamt 28 Gedächtnissportler in einer Reihe von Teilstudien untersucht. Dabei konnte gezeigt werden, dass anders als noch von Maguire und Kollegen berichtet, die Gedächtnissportler überdurchschnittlich intelligent waren. Insbesondere fielen ihre Leistungen in einer Aufgabe zur Messung der Verarbeitungsgeschwindigkeit auf. Selbst im Vergleich zu einer Kontrollgruppe gleicher Intelligenz waren die Gedächtnissportler besser und zudem korrelierten ihre Bestleistungen in der schnellsten Disziplin im Gedächtnissport (Speed Cards) mit der 6 Verarbeitungsgeschwindigkeit. Mit Hilfe zweier Gedächtnisaufgaben (Directed Forgetting und False Memories) konnte gezeigt werden, dass die Gedächtnissportler nur dann außerordentliche Leistungen zeigen, wenn sie auf ihre Techniken zurückgreifen. -Sofern dies geschieht, können sie sich nicht nur mehr merken, sondern haben auch einen genaueren Zugriff auf Gedächtnisinhalte und sind weniger anfällig für Gedächtnisfehler. Im Rahmen einer fMRT-Untersuchung wurde die Wiedergabe von Folgen von Binärziffern untersucht, die entweder einige Tage zuvor oder direkt vor der Wiedergabe auswendig gelernt wurden. Hierbei zeigte sich, dass selbst bei gerade erst mit Hilfe der Mnemotechniken gelernten Ziffernfolgen keine Aktivierung von mit Arbeitsgedächtnis-Gedächtnisprozessen assoziierten fronalten Gehirnarealen feststellbar ist. Im Gegensatz stehen Folgen kurzer Ziffern, die mit dem Arbeitsgedächtnis gelernt wurden, sowie andere Studien, die zeigen, dass bei Verwendung von Gruppierungsstrategien im Arbeitsgedächtnis die frontale Aktivierung noch zunimmt (Bor, Duncan, Wiseman, & Owen, 2003). Diese Ergebnisse unterstützen das Modell des Langzeitarbeitsgedächtnisses (Ericsson & Kintsch, 1995), welches postuliert, dass Experten in ihrem Themengebiet direkt auf Netzwerke im Langzeitgedächtnis zurückgreifen können, wenn sie dazu passende neue Inhalte lernen. Dies geschieht im gleichen Tempo wie sonst nur beim Arbeitsgedächtnis. Dazu passt auch, dass bei Arbeitsgedächtnisaufgaben selbst keine Unterschiede zwischen Gedächtnissportlern und Kontrollgruppen zu finden waren. Obwohl die Gedächtnissportler ausnahmslos alle angegeben haben, dass ihre außergewöhnlichen Leistungen auf den Mnemotechniken und intensivem Training basieren, kann allein auf diesen Daten nicht herausgefunden werden, welche etwaige Fähigkeiten möglicherweise nötig sind, um solche Leistungen zu erreichen. Deshalb wurden in der zweiten Studie normale Probanden in Gedächtnistechniken trainiert. Dabei standen die Routenmethode (Roediger, 1980) und das Major-System (Patton, 1986) im Vordergrund, welche sich beide in zahlreichen Studien bereits als wirksam erwiesen haben (Worthen & Hunt 2011). Allerdings gab es bisher nur wenige Studien, die Gedächtnistraining über einen längeren Zeitraum begleitet haben (Ericsson, Chase, & Faloon, 1980; Higbee, 1997; Kliegl, 7 Smith, Heckhausen, & Baltes, 1987). Diese blieben zudem auf die Untersuchung von Einzelpersonen begrenzt. Die vorliegende Studie war so die erste, welche ein intensives Gedächtnistraining mit einer Gruppe von Probanden begleitet hat. 20 Probanden haben an einem zweitägigen Kurs zu Gedächtnistechniken teilgenommen und anschließend über sechs bis acht Wochen mittels einer Online-Plattform von zu Hause aus trainiert. Insgesamt 13 der Probanden erfüllten das Trainingssoll von mindestens 20 Stunden. Dabei zeigten sich deutliche Trainingsverbesserungen in den Gedächtnisaufgaben. So konnten die Teilnehmer in einer Zahlenmerkaufgabe ihre Leistung mehr als verdoppeln und auch im Wörtermerken deutliche Verbesserungen erzielen. Dabei waren die erzielten Verbesserungen unabhängig von der Intelligenz oder der Leistungsfähigkeit zu Beginn. Darüber hinaus verbesserten sich die Trainingsprobanden auch in der Transferaufgabe zur Verarbeitungsgeschwindigkeit signifikant, im Gegensatz zur Kontrollgruppe, die nicht trainiert hat. Interessanterweise stieg die Selbsteinschätzung der eigenen Gedächtnisleistung trotz großen messbaren Erfolgs nur gering. Dies ist ein Indiz dafür, warum sich in Umfragen zeigt, dass viele trotz Kenntnis von Gedächtnismethoden diese nicht anwenden (Soler, María JoseRulz, 1996). In der dritten Studie wurden Probanden nur relativ kurz (eine Stunde) in der Routenmethode instruiert und sollten diese anschließend benutzen, um sich 50 Begriffe in der richtigen Reihenfolge einzuprägen. Danach hielten sie für eine Stunde Mittagsschlaf oder blieben wach. Aus der Forschung ist bekannt, dass die Gedächtnisleistung von Schlaf profitiert (Rasch & Born, 2013) und dies konnte auch für mittels der Routenmethode gelernte Begriffe gezeigt werden. Wenn im Schlaf zusätzliche Töne eingespielt wurden, die mit den gelernten Begriffen assoziiert worden waren (Rudoy, Voss, Westerberg,
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