Uncertainty and Decision-Making in the Human Brain
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UNCERTAINTY AND DECISION-MAKING IN THE HUMAN BRAIN Iris M. D. Vilares Dissertation presented to obtain the Ph.D degree in Biology | Neuroscience Instituto de Tecnologia Química e Biológica | Universidade Nova de Lisboa Research work coordinated by: Oeiras, December, 2013 ii To my family iii iv APOIO FINANCEIRO/FINANCIAL SUPPORT Apoio financeiro da FCT e do FSE no âmbito do Quadro Comunitário de Apoio, bolsa de doutoramento n° SFRH/BD/33272/2007. O ultimo ano foi financiado pelo National Institute of Health, bolsa n° 2P01NS044393. Financial support for this thesis was provided by FCT and FSE through the Quadro Comunitário de Apoio, doctoral fellowship n° SFRH/BD/33272/2007. The last year was provided by the National Institute of Health, grant n° 2P01NS044393. v vi ACKNOWLEDGEMENTS Many people have helped me through this path, and I am immensely grateful to all of them. I would like to thank: My supervisor, Dr. Konrad Kording. Thank you for being an amazing supervisor, for always being there to help me. You developed in me a love for science and always believed in my scientific skills, sometimes even more than myself. Thank you for your guidance and support at all times. For always feeling welcomed to go to you for any question, be it scientific or not. Thank you for an incredible mentorship. You are my scientific role model. My internal supervisor, Dr. Rui Costa. Thank you for your enthusiasm and support at all times. Thank you for all the help you gave me in taking care of bureaucracy, especially this past month, when almost every day I was sending you an email. Your constant smile and optimism are a source of positive energy. The International Neuroscience Doctoral Program, or INDP (previous Programa Gulbenkian Champalimaud em Neurociências) for giving me the possibility of doing my PhD, and allowing me to develop my research at Northwestern University. I and my colleagues were the “first batch” of this amazing program, and it has been a very interesting and rewarding experience. We had one year of full-time classes at the Instituto Gulbenkian de Ciencia (IGC), in which we could interact and learn with successful scientists from all over the world; and then we had the luxury to have 4 years of funding to go and do our research anywhere. Thank you to Dr. Zachary Mainen for embarking with us in this journey and being the first director of INDP. Thank you as well for your support and love for science. Thank you to the Fundação Champalimaud, the IGC and the Fundação para vii a Ciência e Tecnologia (FCT), for funding INDP and giving institutional and monetary support. A special thanks to IGC for kindly being our host institution when the Champalimaud Foundation was still being built; to Fundação Champalimaud for institutional support and organizing retreats througout the years where I could discuss and present my work; and to FCT for providing me financial support (grant n° SFRH/BD/33272/2007). My thesis committee members, Dr. Jorge Carneiro and Dr. Rosalina Fonseca. Thank you for your questions and for enabling me to look at my results through different perspectives. Thank you as well for your support, friendliness and friendship throughout the years. The doctors at Northwestern Memorial hospital and the Rehabilitation Institute of Chicago, for showing me how rewarding and stimulating it can be to work with patients, for helping me recruit patients and for promoting the combination of science and motor rehabilitation. I would specifically like to thank Dr. Tania Simuni, Dr. Santiago Toledo and Dr. Deborah Gaebler. The people that were always there to help me with paperwork, reimbursement, visa issues or whatever else came along. Namely, I would like to thank: Manuela Cordeiro, Ana Portocarrero, Teresa Carona, Alexandra Piedade, Lance Adams, Robin Millard, Paul Hixon, Kathryin Niedner, and, of course, Debbie Znajda Kitterman. Hugo Fernandes, both colleague and friend. We started our PhD together and, if everything goes well, we will also finish it at the same time. Thank you for being there through the good and bad times. Thank you for basically helping me with all my projects, either through discussions, suggestions or even help making the figures. Thank you for giving me the viii extra bit of motivation that helped me to go across the ocean for my PhD. I don’t think this thesis would have existed without you. Mark Albert, ex-colleague and still good friend. Thank you for making a great lab environment. Thank you for the discussions we had, the help you gave me with my projects, and the useful feedback you gave to my thesis. Thank you for always being there for me, both to celebrate with me when things were going well, and as a shoulder to rely on when things were difficult, both inside and outside the lab. Thank you for your positivism, energy, and for being such a great and trustworthy friend. The other past and present members of the Kording lab, including Ben Walker, Daniel Acuña, Max Berniker, Steve Antos, Pat Lawlor, Ted Cybulski, Josh Glaser, Rebecca Moore, Paul Wanda and Kunlin Wei. A special thanks to Greg Dam, who first taught me how to use Matlab; to Ian Stevenson, whose capacity to write code at a super-natural speed still amazes me; and to Pavan Ramkumar: thank you for your support in writing the thesis (including making me company in coffee shops!), your friendliness and enthusiasm, and your friendship, that has been a source of positive energy in these last months. In general, thank you to all the members of the Kording lab for making the lab a friendly and scientifically stimulating place to work. My colleagues from the INDP program, including Patricio Simoes, Patricia Correia, Rodrigo Abreu, Maria InêsVicente, Margarida Agrochão, Pedro Ferreira and Isabel Henriques. A special thanks to José Fernandes, who has helped me a lot in these last months, advising me on labs, thesis writing and paperwork; and to Mariana Cardoso, for being such a good friend and always always being there. ix The whole range of amazing people that I have met in Chicago. Specifically, I would like to thank: Inês Domingues, who gave me food, shelter and friends when I arrived to Chicago; Lisa Morselli, Fanny Delebecque; Sandra Maximiano and Brenda Lopez Silva, who gave me their friendship from the very beginning and still do, even if I’m sometimes absent for a long time; Kalina Michalska and Jonas Olofsson, for interesting discussions, help with fMRI and friendship; James Howard: thank you for teaching me fMRI, developing with me one of my studies, and for being a good friend; Joanna Wu: When I first met you, you were the “significance maker” of our study. From then on, you have been an invaluable friend and also teacher (cooking, crochet, pottery, etc.); Neha Mehta: we met over a poor dying bee at the bus after the NUIN retreat, and since then you have been a great friend and a constant supply of positive energy and support.; Thanks also to David Spjut, for doing the piece of artwork that ended up being the cover of my thesis, even though he didn’t personally know me; and thanks to Jos Zwanikken, for thinking about it and asking his friend David to do the artwork, and for good discussions, moments, and support. Thanks to Mário Neto, for introducing me to an amazing set of people, for advising me on thesis-writing, and for friendship: I am very glad that you ended up staying much longer in Chicago!; Special thanks to Laetitia Chauve: thank you for your company at coffee shops when writing my thesis; the great moments and trips we spent together; for your constant support and friendship; and in general for always being there; Thanks to Irina Franco, for the constant supply of positive energy, support and friendship; Thanks to Kai Orton: for the comments on my thesis, the uplifting kai-hugs, and for your support and friendliness; Thanks to Bahar Sharafi and Marije De Bruin, for your enthusiasm, support and friendship here at RIC. In general, thank you to all my friends here at Chicago, for amazing moments, celebrations, conversations and friendship that made living in Chicago a great and unforgettable experience. x My friends in Portugal and other parts of the world. I would like to thank: Raquel Antunes: thank you for the comments on my thesis, your support even if oceans apart (glad they invented gchat;)) and in general thanks for being an amazing “mana”; Candida Shinn, for your strength, care and kindness; and Barbara Mauricio, for always being so nice and greeting me with such enthusiasm when I come to Portugal. Thanks to Paula Silva and Anabela Maia, for help in the thesis and a friendship that started when we were small children and still goes on. Thanks to my master thesis colleagues and friends, for making math a much more exciting subject and for being always welcoming: Telma Correia, Catarina Miranda, Teresa Sampaio, Laura Wise, Zé Leal and Ricardo São João. Thanks also to Barbara Vreede and Nuno Sepúlveda, for helping me with my thesis and for friendship. A special thanks also my non-scientist friends, namely Marta Alarcão, Teresa Pessoa, Ana Rita Correia, Ana David, Sara Brasileiro, Alexandre Sousa and Bernardo Santos: thank you for your love and support even when I went to live so far away and wasn’t so present anymore. Thank you all for your amazing friendship. Finally, I would like to thank my family: without you, nothing of this would ever be possible. I would like to thank: my cousin and “irmona” Alexandra Vilares, for our conversations that developed in me a passion for psychology and friendship; Oma, Margaret Wardenaar-Donga, Rachel Wardenaar, Piet Donga and Stefan Donga, for wanting to fly all the way from the Netherlands just to come to my PhD defense and support me; Pedro, Isaltina and Eduardo Vilares, Annelies and Oscar Donga, Vincent Pietersma, Stijn and Rene Wardenaar.