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The Neural and Cognitive Foundations of Human Planning

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The Neural and Cognitive Foundations of Human Planning The Neural and Cognitive Foundations of Human Planning Inaugural-Dissertation zur Erlangung des Doktorgrades der Wirtschafts- und Verhaltenswissenschaftlichen Fakultät der Albert-Ludwigs-Universität Freiburg im Breisgau Vorgelegt von Kai Nitschke geboren am 07.11.1988 in Erfurt 03. April 2017 Freiburg im Breisgau Gutachter Prof. Dr. Heinrichs Prof. Dr. Tuschen-Caffier Dekan Prof. Dr. Renkl Tag der Promotion 28. September 2017 0 List of Contents Page Acknowledgments..................................... iv Abstract...........................................v Zusammenfassung.................................... vii 1 Theoretical and Methodological Background1 1.1 Planning and Executive Functions.......................1 1.2 The Neural Foundation of Planning.......................5 1.2.1 Insights from Lesion Studies.......................5 1.2.2 A Literature Research on Lesion Data.................6 1.2.3 The Influence of Structuredness.................... 12 1.2.4 Overviews on Hemispheric Lateralization.............. 14 1.2.5 The Present Approach on Examining Neural Lateralization... 15 1.3 The Cognitive Foundation of Planning..................... 16 1.3.1 A Sequential Model of Planning.................... 16 1.3.2 The Tower of London and its Structural Parameters........ 18 1.3.3 The Segmentation of Planning..................... 21 1.3.3.1 Cognitive Architectures for Modeling Sub-processes.. 21 1.3.3.2 Dissociating Components of Planning on Tower Tasks. 25 1.3.3.3 Insights from Eye Movement Studies........... 27 1.3.4 The Present Approach on Cognitive Processes of Planning.... 34 1.4 Methodological Background........................... 35 1.4.1 Activation Likelihood Estimation.................... 35 1.4.2 Eye Movement Measurement...................... 36 1.4.3 Pupillometry................................ 37 1.5 Aims and Objectives of the Thesis........................ 39 i 2 A Meta-Analysis on the Neural Basis of Planning: Activation Likelihood Estimation of Functional Brain Imaging Results in the Tower of London Task 41 2.1 Theoretical Background.............................. 41 2.2 Methods........................................ 43 2.2.1 Study Selection............................... 43 2.2.2 Meta-Analyses on Overall Planning and Planning Complexity.. 45 2.2.3 ALE Approach................................ 47 2.3 Results......................................... 48 2.3.1 The Functional Anatomy of Planning Processes........... 48 2.3.1.1 Meta-analysis of neural activation patterns for overall planning............................. 48 2.3.1.2 Meta-analysis of neural activation patterns for planning complexity............................ 48 2.3.1 The Anatomical Localization of Planning Processes in the dlPFC..................................... 51 2.3.2 The Hemispheric Lateralization of Planning Processes in the PFC and beyond................................. 52 2.3.2.1 ALE contrast analysis..................... 52 2.3.2.2 Exploration of Systematic Differences between Studies 52 2.3.2.3 Whole-brain lateralization analyses............ 55 2.4 Discussion...................................... 55 2.4.1 The Localization of Planning Processes within the Mid-dlPFC.. 56 2.4.2 The Role of the Left and Right Mid-dlPFC in Planning...... 57 2.4.3 The Functional Anatomy of Planning Processes beyond (Mid)- dlPFC..................................... 62 2.4.4 Conclusion.................................. 63 2.5 Additional Data................................... 64 3 Dissociating task-demand-specific differences in cognitive processing during planning and problem solving: A validation approach using pupillometry 71 3.1 Introduction..................................... 71 3.2 Methods........................................ 75 3.2.1 Sample Description............................ 75 3.2.2 The Tower of London Task and Problem Parameters........ 75 ii 3.2.3 Experimental Groups........................... 76 3.2.4 Procedure, Apparatuses, and Processing of Data.......... 77 3.2.5 Statistical Analysis............................. 81 3.3 Results......................................... 81 3.3.1 Main Analysis................................ 81 3.3.2 Post-hoc Analyses............................. 83 3.3.3 Control Analysis.............................. 85 3.4 Discussion...................................... 86 3.4.1 General effects on pupil dilation.................... 87 3.4.2 Parameter-specific effects of Tower Configuration on pupil dilation and processes of internalization ............... 88 3.4.3 Parameter-specific effects of Search Depth on pupil dilation and processes of core planning ........................ 90 3.4.4 Limitations................................. 92 3.4.5 Conclusion.................................. 92 4 General Discussion 95 4.1 Summary of the Studies.............................. 95 4.2 The Neural Lateralization of Planning..................... 97 4.2.1 Different Perspectives on the Neural Functional Lateralization. 97 4.2.2 An Approach on a Unified Neural Theory on Planning...... 103 4.2.3 Limitations of the Meta-analysis on Neural Correlates of Planning................................... 106 4.3 Pupillometry and the Cognitive Framework of Planning.......... 109 4.3.1 Integration of Present Results...................... 109 4.3.2 The Potentials and Limitations of Pupillometry as an Indicator for Cognitive Processes.......................... 111 4.3.3 A Critique of a Sequential Model of Planning............ 114 4.4 Outlook on Linking the Cognitive and Neural Foundations of Planning 116 4.5 Conclusion...................................... 121 5 Bibliography 123 6 Appendix 145 Curriculum Vitae..................................... 146 Publication record and scientific work........................ 149 iii ACKNOWLEDGMENTS Acknowledgments This thesis would not have been possible without the support of numerous people. I am most greatly indebted to Dr. Christoph Kaller for supervising my PhD thesis from the early days of planning to the final manuscript. Thank you for the opportunity to participate in your research and your support in all stages of this dissertation. Thank you to Prof. Dr. Markus Heinrichs and Prof. Dr. Cornelius Weiller; without your support this thesis would not have been possible. I am very grateful for your professional supervision and the opportunity to be able to use structures of both the university clinic as well as the university. I am greatly indebted to my colleague Dr. Lena Köstering all the discussions we had about various topics and all advice that you gave me during the PhD process. Thank you to Dr. Elisa Scheller for your support, especially in the final stages. I thank my colleagues Konrad Schumacher and Charlotte Schmidt for an enjoyable and relaxing work atmosphere. You provided a great work environment full of discussions, support, and company in my daily work life. Finally, I would like to thank Dr. Julia Asbrand for her tremendous encouragement that helped me overcome numerous obstacles in the entire course of this thesis. Freiburg im Breisgau, 03. April 2017 Kai Nitschke iv Abstract Executive functions (EF) in terms of conscious control of behavior are highly relevant for adequate functioning in everyday life. Planning is a prototypical EF numerously studied in cognitive psychology and neuroscience. However, the specific neural and cognitive correlates are still subject to debate. In detail, there is an ongoing discussion about the neural lateralization of planning in the human brain as well as about the temporal separability of different phases of planning. These two issues were addressed in the presented thesis. Concerning the issue of neural lateralization, a recently published review article on the neural involvement during human planning based on selected brain lesion studies postulated a rather strict hemispheric lateralization of different types of planning tasks. Planning in well-structured tasks such as the Tower of London (ToL) was postulated to be processed solely by the prefrontal cortex (PFC) of the left but not the right hemisphere. This proposal was however contrasted by a qualitative overview of functional neuroimaging studies showing that the right PFC was activated in a large number of functional neuroimaging studies using the well-structured ToL. To substantiate either conclusion, in the first study of this thesis a quantitative meta- analysis was conducted based on the well-established activation likelihood estimation methodology. A thorough literature search on all available ToL studies yielded 29 neuroimaging publications suitable for such an analysis. The analysis statistically confirmed a bilateral involvement of left and right PFC during well-structured planning. Furthermore, an extensive literature search was conducted on brain lesion studies that using well-structured planning tasks also yielded no indication for a unilaterality of well-structured planning. Concerning the issue of temporally separable phases of planning, previous analy- ses of eye-movement patterns during the ToL determined distinct phases of planning. In a recent eye-movement study manipulation of specific structural parameters of ToL problems allowed to isolate and segment of two temporally distinct phases of cognitive processing, an initial phase serving to create a mental representation of the problem (representation creation) and a subsequent phase serving to generate v ABSTRACT the sequence of solution steps, i.e. the actual planning phase (sequence generation). However, the eye tracking methodology entails several
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