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Electrophysiological Brain Dynamics of Visuospatial Planning PONTIFICIA UNIVERSIDAD CATÓLICA DE CHILE Doctorado en Neurociencias Electrophysiological Brain Dynamics of Visuospatial Planning by Marcos Eduardo Ernesto Domic Siede Doctoral Thesis Submitted to the Neuroscience Program in Partial Fulfillment of the Requirements for the Degree of Ph. D. in Neurosciences at the Pontificia Universidad Católica de Chile November 2020 Advisor : Dr. Tomás Ossandón Valdés Co-advisor : Dr. Marcela Perrone-Bertolotti President of the committee : Dr. Nicolás Crossley Karmelic External evaluator : Dr. Pedro Maldonado Arbogast External evaluator : Dr. Daniel Rojas-Líbano PONTIFICIA UNIVERSIDAD CATÓLICA DE CHILE Doctorado en Neurociencias El Comité de Tesis, constituido por los Profesores abajo firmantes, aprueba la Defensa de la Tesis Doctoral titulada: Electrophysiological Brain Dynamics of Visuospatial Planning Aprobación Defensa: MARCOS EDUARDO ERNESTO DOMIC SIEDE Calificándose el trabajo realizado, el manuscrito sometido y la defensa oral, con nota …..... (……………..) _______________________________ ____________________________ Dr. Mauricio Cuello Dr. Luis Ibáñez A. Director de Investigación y Doctorado Decano Escuela de Medicina Facultad de Medicina Pontificia Universidad Católica de Chile Pontificia Universidad Católica de Chile ______________________________ ______________________________ Dra. Claudia Sáez Dr. Tomás Ossandón Valdés Sub-Directora Director de Tesis Dirección de Investigación y Doctorado Facultad de Medicina Escuela de Medicina Pontificia Universidad Católica de Chile Pontificia Universidad Católica de Chile ______________________________ ____________________________ Dr. Nicolás Crossley Karmelic Dra. Marcela Perrone-Bertolotti Presidente comité de tesis Co-Directora de Tesis Escuela de Medicina Université Grenoble Alpes Pontificia Universidad Católica de Chile ______________________________ _______________________________ Dr. Francisco Aboitiz D. Dr. Pedro Maldonado Jefe Programa Doctorado en Neurociencias Profesor Evaluador Externo Centro Interdisciplinario de Neurociencias Facultad de Medicina Facultad de Medicina Universidad de Chile Pontificia Universidad Católica de Chile _______________________________ Dr. Daniel Rojas-Líbano Profesor Evaluador Externo Facultad de Psicología Universidad Diego Portales Santiago, November 2020 “El cerebro no solo es un órgano capaz de conservar o reproducir nuestras pasadas experiencias, sino que también es un órgano combinador, creador, capaz de reelaborar y crear con elementos de experiencias pasadas nuevas normas y planteamientos.” —Lev Vygotsky Agradecimientos Nunca había entendido tan bien como ahora las veces que escuché desde muchos compañeros, colegas y amigos, frases que hacen referencia a que la tesis de doctorado se transforma en una extensión de uno mismo. Luego de este largo recorrido, horas de extenso trabajo, varios momentos de alegría, pero bastantes también de frustración, hasta finalmente concluir una de las etapas más importantes de mi carrera profesional y académica, es que esta idea hace tanto sentido. En este largo recorrido son muchas las personas a las que quisiera agradecer, desde mis directores de tesis del Magister en Neurociencias de la Universidad de Chile que vieron en mi un candidato a realizar este recorrido y me alentaron a postular al Programa de Doctorado en Neurociencias de esta Universidad: el Dr. José Luis Valdés, Dr. Francisco Aboitiz, y por supuesto, quien es mi actual director de esta tesis de doctorado, el Dr. Tomás Ossandón, el cual además me ha brindado tantas oportunidades valiosas para desarrollarme y ser el científico que actualmente soy. Además, quisiera agradecer todo el aporte, y horas de trabajo dedicado por quien es mi co-tutora de tesis, la Dra. Marcela Perrone-Bertolotti, quien amable y alegremente me recibió en el Laboratoire de Psychologie et NeuroCognition de la Université Grenoble-Alpes durante mi pasantía doctoral, y ha revisado minuciosamente cada uno de mis avances, borradores, y escritos que le envío, contribuyendo enormemente a lo que este trabajo sea lo que es hoy. Quisiera agradecer además al equipo que conforma mi laboratorio: Neurodinámica de la Cognición, quienes han dedicado largas horas de atención y discusión a mis presentaciones o incluso leído algunos de mis borradores del artículo de investigación, y enriquecieron este trabajo con muy valiosos aportes, entre ellos quisiera mencionar a Vicente Medel, Joaquín Valdés, Brice Follet, Gonzalo Boncompte Josefina Ihnen, Daniela Santander, David Arriagada, y Catalina Fabar. Además, quisiera agradecer al Dr. Pablo Billeke, quien me aceptó en su laboratorio en la Universidad del Desarrollo, para realizar una unidad de investigación, y contribuyó importantemente en el desarrollo del diseño del paradigma conductual de mi tesis de doctorado. Quiero agradecer también al colaborador de colaboradores, a mi amigo y colega, quien es un profesional muy completo e hizo una contribución enorme a este trabajo, el mismísimo Martín Irani, a quien cariñosamente apodé como el dios programador. Una mención muy especial de agradecimiento para mis colegas y amigos, especialmente a Sergio Osorio e Ishani Thakkar, a quien les di más de un dolor de cabeza al corregirme el inglés de varios de mis escritos, incluyendo el de esta tesis, además de sus excelentes retroalimentaciones científicas sobre mi trabajo. Es inconcebible dejar de mencionar a mis fieles amigos y colegas, pero por sobre todo amigos: Rafael Gatica, Felipe Oyarzún, Andrea Sánchez (quién me sacó de tantos apuros por culpa de MATLAB de manera tan desinteresada), Sol Torruella, Rilda León, y Alexandra García, quienes hicieron de este camino, a veces tan duro, mucho más llevadero. También estoy muy agradecido del apoyo financiero otorgado por el programa de Becas de Doctorado Nacional, año 2015 de la Agencia Nacional de Investigación y Desarrollo ex- CONICYT 21150295, los fondos otorgados por FONDECYT regular 1180932, y el Institut Universitaire de France (IUF). Finalmente, quiero agradecer a todos mis amigos y familia en Santiago, a mi padre Jorge Domic, y a los que están en Antofagasta, muy especialmente y sobre todo a mi madre Gloria Siede, y a mi mama Adriana Astudillo que vive en mi memoria, por su apoyo constante e incondicional, a quienes les dedico con mucho amor este trabajo. TABLE OF CONTENT 1. INTRODUCTION ............................................................................................................ 5 1.1. Cognitive Control and Executive Functions .................................................... 5 1.2. Neuroanatomical Basis of Cognitive Control .................................................. 6 1.3. Cognitive Psychology of Planning .................................................................. 9 1.3.1. Successive Refinement Models............................................................... 9 1.3.2. Opportunistic Models ............................................................................... 9 1.3.3. Integrating Successive Refinement and Opportunistic Models .............. 10 1.3.4. Simple motor planning behavior ............................................................ 11 1.3.5. Cognitive Planning ................................................................................. 12 1.4. Behavioral Planning Paradigms ................................................................... 14 1.4.1. Tower of London Task ........................................................................... 15 1.4.2. Porteus Maze Task ................................................................................ 17 1.4.3. Zoo Map Task........................................................................................ 18 1.4.4. Ecological behavioral paradigms ........................................................... 20 1.4.5. A Novel Planning Task .......................................................................... 22 1.5. Neural Correlates of Planning ...................................................................... 22 1.5.1. Functional Contributions of the Anterior Cingulate Cortex and the Mid- cingulate Cortex in Cognitive Control ................................................................. 26 1.5.2. The Frontopolar Cortex and Planning .................................................... 29 1.6. Frontal Theta as a Marker of Cognitive Control ........................................... 35 1.6.1. Frontal Midline Theta Activity ................................................................ 36 1.6.2. Theta Phase Connectivity ...................................................................... 40 1.7. Summary of the Theoretical Background ..................................................... 42 2. HYPOTHESIS AND OBJECTIVES ............................................................................... 45 2.1. Hypothesis ................................................................................................... 45 2.2. General Objective ........................................................................................ 45 2.3. Experimental predictions: ............................................................................. 46 3. STUDY 1: Theta Activity from Frontopolar Cortex, mid-Cingulate Cortex and Anterior Cingulate Cortex Shows Different Role in Cognitive Planning Performance. ........................ 48 3.1. INTRODUCTION ......................................................................................... 50 3.2. MATERIALS AND METHODS ..................................................................... 52 3.2.1.
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