THE NEUROBIOLOGICAL FOUNDATIONS OF ALTRUISM by Dharol Tankersley Department of Philosophy Duke University Date:_______________________ Approved: ___________________________ Alex Rosenberg, Supervisor ___________________________ Scott Huettel ___________________________ Karen Neander ___________________________ Michael Platt ___________________________ Dale Purves Dissertation submitted in partial fulfillment of the requirements for the degree of Ph.D. of Philosophy in the Department of Philosophy in the Graduate School of Duke University 2008 ABSTRACT THE NEUROBIOLOGICAL FOUNDATIONS OF ALTRUISM by Dharol Tankersley Department of Philosophy Duke University Date:_______________________ Approved: __________________________ Alex Rosenberg, Supervisor ___________________________ Scott Huettel ___________________________ Karen Neander ___________________________ Michael Platt ___________________________ Dale Purves An abstract of a dissertation submitted in partial fulfillment of the requirements for the degree of Ph.D. in the Department of Philosophy in the Graduate School of Duke University 2008 Copyright by Dharol Tankersley 2008 Abstract This project advocates an urgent role for neurobiological evidence and models in the study of altruism. I argue for two claims: that neurobiological evidence should be used to constrain candidate scientific accounts of altruistic behavior, and that neurobiological techniques can be used to elucidate component mechanisms of altruistic behavior. Chapter 1 reviews the historical progression of theories of altruism, and the empirical observations that motivated their development. A distinction is drawn between evolutionary altruism ‐‐ any self‐sacrificial, fitness‐reducing behavior, and psychological altruism ‐‐ self‐sacrificial behaviors that are caused by psychological states like desire and motivation. Three theories of psychological altruism are described, and it is argued that the crucial difference between these theories is their conceptions of the role of affect in motivation, and how the processes of affect and motivation contribute to psychological altruism. Chapter 2 describes dominant theories of motivation and the neurobiological and psychological mechanisms that support motivated behavior. Although the evidence is not conclusive, I argue that our best scientific models and neurobiological evidence support affective models of psychological altruism, and that other models are at best incomplete and possibly implausible in light of neurobiological considerations. iv Chapter 3 introduces mind reading approaches to altruism, which argue that the capacity for altruistic motivations depends upon the capacity to represent the psychological states or circumstances of others. I conclude that altruism requires at a minimum the ability to attribute affective experiences to others. Further, I argue that the representations produced by mind reading processes provide a means for distinguishing between self‐regarding and altruistic motivations. In contrast with the dominant philosophical theory of psychological altruism, the mind reading model I propose is compatible with the affective theory of motivation depicted in Chapter 2. My own empirical work is described as an example of how neurobiological techniques can reveal the differential role of neural systems in producing self‐regarding and altruistic behavior. Chapter 4 departs from the mechanistic approach to altruism discussed in the previous chapters, and presents an overview of how the fields of philosophy, psychology, psychobiology and genetics, have investigated altruism as a stable characteristic or personality trait. Recent technological advances make this a promising approach for investigating the psychological and neurobiological systems supporting altruistic behavior. v Contents Abstract.........................................................................................................................................iv List of Tables.................................................................................................................................ix List of Figures................................................................................................................................x Chapter 1: Models of Altruism....................................................................................................1 1.1 Evolutionary Altruism: Self‐sacrificial behavior in a Darwinian world.........3 1.2 Psychological Altruism: Motivated helping........................................................5 1.2.1 Directionality of desires: Egoism versus AltruismP..........................................9 1.2.2 Evolutionary arguments for Psychological AltruismP....................................15 1.3 Motivational mechanisms: Affective versus propositional approaches to altruismP................................................................................................................19 Chapter 2: Models of Motivation..............................................................................................29 2.1 Behaviorist models of motivation: The case for affective altruismP...............30 2.2 Neurobiology of motivation: Mechanistic monism..........................................35 2.2.1 Reward Event Theory..........................................................................................38 2.2.2 Partitioning motivation: ‘wanting’ versus ‘liking’..........................................39 2.3 Theoretical implications of the neurobiology of motivation...........................43 Chapter 3: AltruismP and Theory of Mind..............................................................................49 3.1 Mind reading and altruismP................................................................................50 3.1.1 Emotional contagion: AltruismP without mind reading.................................51 3.1.2 AltruismP with mind reading: What gets represented?..................................56 3.2 Distinguishing self‐ and other‐regarding motivations.....................................60 vi 3.3 Neurocognitive systems of perception and altruismP .....................................63 3.4 From mind reading to altruisticP motivation...................................................79 3.5 Conclusions............................................................................................................85 Chapter 4: The AltruisticP Personality.....................................................................................86 4.1 Virtuous tendencies..............................................................................................86 4.2 Empathic and altruisticP dispositions.................................................................88 4.3 Biological bases of resilience and affiliation......................................................91 Concluding Thoughts.................................................................................................................97 Appendix 01: FMRI Data Acquisition and Analysis............................................................100 01.1 Cognition and neuronal metabolism................................................................100 01.2 Data acquisition...................................................................................................102 01.3 Data analysis........................................................................................................105 01.3.1 Preprocessing..................................................................................................106 01.3.2 Statistical analysis...........................................................................................110 01.4 Strengths and limitations of BOLD fMRI.........................................................113 Appendix 02: Experimental Methods and Results...............................................................119 02.1 Experiment 1: AltruismP predicts increased neural response to agency.....119 02.1.1 Methods............................................................................................................119 02.1.2 Results..............................................................................................................123 02.2 Experiment 2: Relationship between altruismP and rSTC activation not mediated by other psychometric variables.....................................................130 02.2.1 Methods............................................................................................................130 vii 02.2.2 Results..............................................................................................................132 02.3 Experiment 3: Perceptual ability and altruismP predict activation of rSTC137 02.3.1 Methods............................................................................................................137 02.3.2 Results..............................................................................................................138 02.4 Experiment 4: Contribution of simple and complex perceptual processes to altruismP..............................................................................................................142 02.4.1 Methods............................................................................................................142 02.4.2 Results..............................................................................................................146 Appendix 03: Personal Altruism Level (PAL) Scale............................................................149 Acknowledgements..................................................................................................................152 References..................................................................................................................................154 Biography...................................................................................................................................168 viii List of Tables Table 02.1: Regions Active for Watching > Playing in Experiment 1................................126 Table 02.2: Brain Regions Predicting AltruismP in Experiment 1.......................................127 Table 02.3: Active Regions for Watching > Playing Contrast in Experiment 2................129 Table 02.4: Brain Regions Predicting AltruismP in Experiment 2.......................................136 Table 02.5 Active Regions for Social > Nonsocial Contrast in Experiment 3....................142 ix List of Figures Figure 3.1: Distinguishing Self‐ and Other‐Regarding Motivation......................................61 Figure 3.2: AltruismP is Predicted by rSTC Activation.........................................................66 Figure 3.3: Enhanced rSTC Activation to the Detection of Agency.....................................69 Figure 3.4: Contingency Discrimination Ability Predicts AltruismP...................................72 Figure 3.5: Activation in rSTC Predicts AltruismP and Contingency Discrimination.......74 Figure 3.6: Trend Between AltruismP and Discrimination....................................................76 Figure 3.7: Visual Attention and Social Perception Recruit Distinct rSTC Regions..........78 x
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