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Theses

2012

An investigation into the neural substrates of to determine the key place of in human moral development

Andrew P.L Mullins University of Notre Dame Australia

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Publication Details Mullins, A. P. (2012). An investigation into the neural substrates of virtue to determine the key place of virtues in human moral development (Doctor of Philosophy (PhD)). University of Notre Dame Australia. http://researchonline.nd.edu.au/theses/78

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An investigation into the neural substrates of virtue to determine the key place of virtues in human moral development.

Research thesis submitted for the degree of Doctor of Philosophy.

Andrew P. J. Mullins

School of Philosophy and Theology University of Notre Dame, Australia.

December 2012.

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Table of Contents

Abstract. Declaration of Authorship. Acknowledgements. Abbreviations.

Chapter 1 Page Aristotle and the neuroscientists sit down to talk.

1.1. The structure of this study. ………………………………….……………... 21

1.2. Preamble. The prima facie case. ………………………………………….. 22 1.2.1 Material and non-material constituents to human rationality. 1.2.1.1. We are not disembodied spirits. Evidence of the involvement of the physical in rationality. 1.2.1.2. Nor are we mere bodies. Intimations of the non-material. 1.2.1.3. A prima facie case exists for a profound integration of the physical and rational in the human person. 1.2.2. Prima facie evidence for a neuronal underpinning of virtue.

1.3. Situating this study within contemporary approaches to the philosophy of mind. …………………………………………………………….. 34

1.4. The hylomorphic underpinnings of this study. …………………….. 37 1.4.1 An overview of hylomorphic theory. a) Substance. b) Soul. 1.4.2 Rational psychology. 1.4.3 Restoring the hylomorphic notion of person. 1.4.4 Advantages offered by an hylomorphic notion of person.

1.5. An hylomorphic critique of 20th century currents in philosophy of mind. …………………………………………………………….. 65

1.6. Situating this study in current approaches to ethics. ……………. 70 1.6.1 Some reasons for the growing stature of virtue ethics. 1.6.2 Within virtue ethics there exists a spectrum of significantly contrasting approaches. 1.6.3 This study is situated under the umbrella of eudaimonist virtue ethics.

1.7 An overview of the potential benefits of this study. …………….. 81

1.8 Conclusion. ………………………………………………………………………….. 83

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Chapter 2 Neural mechanisms and systems.

2.1 Overview. ……………………………………………………………………………. 85 2.1.1 Rationale for the structure adopted. 2.1.1.1 Imaging the brain. 2.1.1.2 Prima facie confirmation for the broad direction of this investigation: Projects recognised in the 2009 Templeton Positive Awards. 2.1.1.3 The human act provides insight into the function of virtue. 2.1.1.4 Insights into the basal ganglia. a) Pathological insights into dysfunctional basal ganglia: Huntington disease, Obsessive Compulsive Disorder, Parkinson disease and Encephalitis Lethargica. b) A role for the basal ganglia in volitional habits. 2.1.2 Search information. 2.1.3 This neuroscientific investigation carries some caveats. a) The brain is constantly in a state of change. b) Our appreciation of the complexity of brain networking is at a rudimentary level. c) The basis of the interface between molecular processes and conscious processes, such as learning and memory, is still conjectural. d) Our experimental knowledge of brain function is derived from animal as well as human studies. e) Our capacity to study neuronal activity in real time and with precision is uneven and still developing.

2.2 Plasticity. …………………………………………………………………………….. 107 2.2.1 Historical considerations. 2.2.2 Plasticity is categorised as functional or structural. 2.2.3 The big picture: synaptic strengthening as the net result of structural plasticity. 2.2.4 Developmental plasticity. 2.2.4.1 Sensitive periods in humans. 2.2.4.2 Mechanisms of developmental phases. a) Experience tuned development. b) The role of axons. c) Synapse formation and elimination. 2.2.5 Other synaptic factors. 2.2.5.1 NMDAR and AMPAR mediated plasticities. a) NMDA plasticities. b) Interplay with AMPARs. 2.2.5.2 DA and the GPCR receptor. 2.2.5.3 Neurotrophic factors.

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2.2.5.4 Other chemical signalling systems. a) Ca2+. b) ACh. c) Other chemical signalling. d) Desensitization. 2.2.5.5 Astrocytes at the synapse. 2.2.5.6 Structural considerations. a) Dendritic spines. b) Dendritic processing facilitates plasticity and cortical control. c) The speed of morphological change. d) Plasticity as a consequence of neurogenesis. e) Microtubule assembly. 2.2.6 Use-induced plasticities. 2.2.7 Exercise induced plasticity. 2.2.8 Plasticity induced by transcranial stimulation.

2.3. Neural bases for learning and memory. ………………………………. 144 2.3.1 Learning and memory. a) Learning. b) Memory. 2.3.2 The five major memory regions of the brain. a) Hippocampus. b) Amygdala. c) Striatum. d) Cerebellum. e) Cerebral cortex. 2.3.3 Mechanisms of memory. 2.3.3.1 Declarative memory storage. 2.3.3.2 Non-declarative, or procedural memory, storage. 2.3.3.3 Working memory, a form of short term memory, storage. 2.3.4 LTP as an underpinning mechanism for memory. 2.3.4.1 Further factors for the synaptic facilitation involved in memory. 2.3.4.2 The neuroscience of memory enhancement. 2.3.5 Skill related learning. 2.3.6 Neural systems of imitation and learning. 2.3.6.1 Neural systems for affective feelings. 2.3.6.2 Neuropeptides and mechanisms of trust.

2.4 Habit formation and the role of the basal ganglia. ………………. 165 2.4.1 The basal ganglia and related structures. 2.4.1.1 A key functional division of the basal ganglia. a) The ventral pars. b) The dorsal pars. 2.4.2 An overview of the contribution of the basal ganglia. 2.4.3 Connectivity to the basal ganglia.

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a) Inputs. b) Outputs. 2.4.4 Loops. 2.4.4.1 Frontal lobe-basal gaglia loops provide a key to complex goal directed behaviours. 2.4.5 The role of the basal ganglia in motor learning. a) Motor learning as a form of memory. b) The learning process: from the explicit to the automatized. c) The comparative roles of the cerebellum and the basal ganglia. d) Numerous brain areas are implicated at both acquisition and completion stages of this learning process. e) and procedural memory. 2.4.6 Habit formation, a particular form of learning. 2.4.7 Plasticity in the basal ganglia and related structures. 2.4.7.1 Forms of basal ganglia plasticity. 2.4.7.2 DA: principal mediator of plasticity in the basal ganglia. 2.4.7.3 Insights from plasticity within the hippocampus. 2.4.8 Initial reflections on the possible role of the basal ganglia in virtue acquisition.

2.5 Neural bases for emotional control. ……………………………………. 198 2.5.1 Theories of emotion. 2.5.2 Emotion and memory. 2.5.3 areas associated with emotion. 2.5.3.1 Involvement of cortical areas in emotional regulation. 2.5.4 Plasticity in the mechanisms of emotional regulation. 2.5.5 Development of attentional mechanisms.

2.6 Neural bases for goal directed behaviours and reward activation. ……………………………………………………………………………. 218 2.6.1 Reward pathways and motivation. 2.6.2 Classical conditioning and DA reinforcement. 2.6.3 Goal directed plasticity in the amygdala. 2.6.4 Reinforcement in basal ganglia learning. 2.6.4.1 DA modulation in striatum for goal directed action. 2.6.5 Frontal cortex and reward systems. 2.6.5.1 The role of the OFC. 2.6.5.2 Close alignment between OFC and striatum. 2.6.6 Addiction studies offer insights into the reward systems.

2.7. Cognition and executive function. ………………………………………. 241 2.7.1 The PFC coordinates executive function. 2.7.2 Functional studies of the PFC. 2.7.3 Prefrontal plasticity.

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2.7.3.1 Plasticity studies in the motor cortices.

2.8 Conclusion. …………………………………………………………………………… 248

Chapter 3 Investigating the characteristics of virtue.

3.1 Dr Takashi Nagai (1908-1951). ……………………………………………… 259 3.1.1 Context: August 9, 1945. 3.1.2 Acts of virtue. a) Scenario 1. Treating a prisoner. b) Scenario 2. The Rising Sun. c) Scenario 3. The Pamphlet. d) Scenario 4. Nyokodo. 3.1.3 Virtue and the human act. 3.1.4 Some observations about the characteristics of virtue. 3.1.5 The superiority of virtue ethics in accounting for noble human behaviour.

3.2 The state of virtue. ………………………………………………………………. 282 3.2.1 Virtues dispose the appetites, the source of all human acts, to rationality. 3.2.1.1 The virtues of the sensitive appetite refine our habitual dispositions to pain and pleasure. 3.2.1.2 Virtue is a habit of choosing. 3.2.2 Virtue lies in choice of the mean. 3.2.3 The motivation of virtue. “Virtuous actions are done for the sake of the noble.” 3.2.4 Virtues bring about states of character. 3.2.5 Virtues accord with the natural perfecting of the person. 3.2.5.1 Virtue is pleasant. 3.2.5.2 “Happiness is the reward of virtue.” 3.2.5.3 Virtue brings about human fulfilment.

3.3 Acquisition of virtue. …………………………………………………………… 303 3.3.1 Both “training” and “education” are needed to build virtue. 3.3.2 An emphasis on the early years assists in fostering virtue. 3.3.3 Imitation can assist the formation of virtue. 3.3.4 Virtues are acquired by repeated acts. 3.3.5 Obedience to others empowers for obedience to one’s own reason. 3.3.6 Affection assists in fomenting virtue. 3.3.7 Development of virtue requires formation in what is appropriate in respect to pleasure and pain… emotional

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education. 3.3.8 Laws, culture and expectations assist in fostering virtue. 3.3.9 The development of virtue takes place over time and respects the development of the body. 3.3.10 Practice of virtue requires effortful attention. 3.3.11 Virtue requires intrinsic motivation. 3.3.12 Education specifically in wisdom and beauty is necessary in the formation of virtue. 3.3.13 Overcoming incontinence and unlearning vice.

3.4 Conclusion. ………………………………………………………………………………… 324

Chapter 4 Understanding the cardinal virtues.

4.1 The cardinal virtues. ……………………………………………………………. 326 4.1.1 Arguments opposing the view that fortitude and temperance are ordered biophysical qualities of the human person. 4.1.1.1 Virtue is of the soul. 4.1.1.2 Reason is the “form” of the virtues. 4.1.1.3 Virtue is a “habit of choosing” and therefore all virtue would appear to be a disposition of the rational appetite. 4.1.1.4 Aquinas appears to suggest that the moral virtues are rational. 4.1.1.5 The practical intellect appears to play an intrinsic role in the virtues. 4.1.1.6 The speculative intellect appears to play a role each of the virtues. 4.1.1.7 It would appear impossible to hold the view that acts of individual virtues are not rational. 4.1.1.8 Virtue results from the ordering of material processes by reason. 4.1.1.9 As virtue is “perfection of a power to act”, therefore it “belongs to that which is proper to the soul”. 4.1.1.10 The habits of virtue all enjoy a certain durability. 4.1.2 Arguments supporting the view that fortitude and temperance are ordered biophysical qualities of the human person. 4.1.2.1 Virtues “in the soul” must be understood correctly. 4.1.2.2 Reason as the “form” of virtue correctly understood. 4.1.2.3 A correct understanding of “habit of choosing” is required. 4.1.2.4 The moral virtues of the sensitive appetite are rational “by participation”. 4.1.2.5 The practical intellect is perfected by its own proper habit, prudence.

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4.1.2.6 If in addition to prudence, the speculative intellect plays a role in virtue, the speculative intellect will be perfected by its own proper habits. 4.1.2.7 Rather than of “acts of isolated virtues” it appears more appropriate to think in terms of “moral operations performed with the integral involvement of multiple virtues”. 4.1.2.8 As it is the cause of virtue, reason is necessarily external to moral virtue. 4.1.2.9 Analysis of the human act supports the view that various virtues are present in concert in a single human act. a) Input from the sensitive appetite precedes the rational in the human act. b) The contribution of emotion to rationality also argues for a “moral-operations view” rather than of a view of actions disposed by isolated virtues. c) Furthermore, discrete ascription of a particular action to a single virtue is problematic. Actions in real life manifest the constellation of virtues. 4.1.2.10 The “durability” of the individual virtues differs in meaning according to the specific virtue.

4.2 Biological and functional distinctions between the virtues. … 358 4.2.1 The cardinal virtues have distinctive roles in the operation of virtue. a) The distinction between the virtues properly of the soul and those properly of the body. b) Prudence and justice, as they are associated more directly with the rationality, are more removed from the material substrate. c) Fortitude and temperance have a dual dependency: on the soul and on the corporeal organs. d) Temperance and fortitude are at the heart of a rational response to pain and pleasure and therefore to appetition that is truly human. 4.2.2 Prudence. 4.2.3 Justice. a) A broad understanding of justice. 4.2.4 An understanding of the biophysical dimensions of prudence and justice. a) What is the matter? b) Biological perfections. 4.2.5 Fortitude. 4.2.6 Temperance. 4.2.7 Biophysical aspects of fortitude and temperance. a) Subordination to reason. b) Biological perfections.

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4.3 Unity of the virtues. ……………………………………………………………… 379 4.3.1 The virtues are “integrated distinct elements” united in the person.

4.4 Conclusions. …………………………………………………………………………… 381 4.4.1 Characteristics pertaining to the state of virtue.

Chapter 5 The neural bases of virtue.

5.1 Reflections on the task being attempted. ……………………………. 386 5.1.1 Overview of this chapter.

5.2 Insights from contemporary cross-disciplinary studies in emotion and morality. ………………………………………………………….. 389 5.2.1 Understanding emotion. 5.2.1.1 Joseph LeDoux. 5.2.1.2 Antonio Damasio. 5.2.1.3 Nancy Sherman and Martha Nussbaum. 5.2.2 Research into the neural bases of morality. 5.2.2.1 Gillett’s studies in psychopathology. a) The field of psychopathology. b) Gillett’s work. 5.2.2.2 Gazzaniga and the neural bases of . 5.2.2.3 Neural subdivisions implicated in moral behaviour.

5.3 Neural bases. ………………………………………………………………………… 407 5.3.1 A complex of systems. 5.3.2 The principal brain systems contributing to the development and exercise of virtue. 5.3.3 The neural substrates of virtue. The state of virtue reflects a state of neural and systemic maturity. 5.3.3.1 A “neural excellence” understanding of the neural bases of virtue. 5.3.3.2 The neural bases of virtue.

5.4 A case study: a simplified neural analysis of a virtuous action performed by Takashi Nagai. …………………………………………………. 476

5.5. Conclusion. …………………………………………………………………………….. 484

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Chapter 6 Flourishing.

6.1 This study proposes a necessary role for virtue in human flourishing. …………………………………………………………………………….. 487 6.1.1. Reflecting on the structure of this study. 6.1.2 The task of Chapter 6.

6.2 Eudaimonic conclusions. ………………………………………………………. 491 6.2.1 Flourishing at the biological level. 6.2.1.1 Rational operations in the present state of life are conducted with sensible knowledge and sensitive appetites as their “material cause”. Therefore neural development that forms the biological basis of virtue is a “material cause” for eudaimonia of the person. 6.2.1.2 Rational operations in the present state of life are conducted with the body as their final cause. 6.2.2 The neurobiology of eudaimonia. 6.2.2.1 Affective fulfilment. Discussion of the neural bases for the feelings of contentment associated with virtue. 6.2.2.2 Functional fulfilment. Discussion of neural and cognitive efficiencies. The intrinsic functional benefits of virtue. 6.2.2.3 Teleological fulfilment. The neurobiology of eudaimonia at the teleological level. 6.2.3 Fulfilment through virtue. 6.2.3.1 Samuel Franklin. 6.2.3.2 .

6.3 Neuroscientific conclusions. …………………………………………………. 516 6.3.1 Reflections on the model that has been proposed. 6.3.2 A summary of neuroscientific conclusions. 6.3.2 Associated insights into brain structure and function.

6.4 Philosophical conclusions. …………………………………………………….. 523 6.4.1 Anthropology. Understanding of the human person. Advantages of an hylomorphic approach. 6.4.1.1 Unity of the virtues reveals itself as a biological necessity. 6.4.1.2 Some thoughts on the nature of vice. 6.4.2 The benefits of a non-reductionist, hylomorphic approach to philosophy of mind. 6.4.2.1 The importance of an hylomorphic understanding of brain function and structure. 6.4.2.2 An enriched understanding of causality accords with the anti- reductionist view and permits dialogue between scientific and philosophical explorations. 6.4.2.3 Functionalist approaches to hylomorphism. a) Rationality is functionally integrative.

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b) A response to prevailing physicalism. c) Bridges to contemporary philosophy of mind. d) Conclusions. 6.4.3 Advantages of a virtue based approach to ethics over consequentialist and deontological approaches. 6.4.3.1 A contemporary validation of virtue ethics in the light of neuroscientific evidence. 6.4.3.2 Understandings of neurobiology assist the discipline of virtue ethics. 6.4.4 Fine tuning the hylomorphic approach.

6.5 Pedagogical conclusions. ………………………………………………………. 548 6.5.1 Ineffective moral pedagogies. 6.5.2 Insights from the neuroscience. a) Principles of plasticity. b) Habit formation as learning. c) Pathways of habit formation. d) Principles of memory. e) Principles of imitation. f) Interplay between emotion and learning. g) Healthy body; healthy mind. 6.5.3 Parenting conclusions. 6.5.3.1 Applications to parenting.

6.6 The last word. ……………………………………………………………………….. 565

Figures.

Figure 2.1 Gross anatomy of the human brain. …………………………………. 87 Figure 2.2 Examples of high resolution magnetic resonance images. 89 Figure 2.3 An example of the use of combined PET and MRI imagery 90 to detect areas of increased cortical activity in specified tasks. Figure 2.4 The astonishing capacity for structural change in the brain, 108 for reengineering connectivities and function. Figure 2.5 Signal transmission at the chemical synapse. 115 Figure 2.6 The developing cortex. …………………………………………………….. 119 Figure 2.7 Location of the basal ganglia. 168 Figure 2.8 The main connections of the basal ganglia. 177 Figure 2.9a Areas associated with the limbic system. 202 Figure 2.9b Location of key limbic areas. 202 Figure 2.10 Phineas Gage and the role of the medial PFC in emotion 212 regulation. Figure 2.11 Links between attention and emotion regulation. …………… 218 Figure 3.1 Dr Takashi Nagai. 261 Figure 3.2 Nagai and his children. 261 Figure 3.3 Nyokodo. 270

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Appendices

Appendix 1 A Response to the Claims of Emergent Rationality by Non- reductive Materialism………………………………………………………. 571 Appendix 2 A Brief Introduction to Neuroscientific Concepts and Terminology…………………………………………………………………….. 578 Appendix 3 Technologies…………………………………………….. 587

Tables

Table 1.1 An hylomorphic critique of 20th century currents in philosophy of mind…………………………………………………………… 592 Table 2.1 Search process. ……………………………………………………………….. 614 Table 2.2 Systems for sensitive periods of development. Table 2.3 The numerous systems and mechanisms of plasticity operating at the cellular and molecular levels. Table 2.4 Systems for learning. Table 2.5 Memory systems. Table 2.6 Systems for imitation and empathy…………………………………… 622 Table 2.7 Systems for habit formation. Table 2.8 Systems for emotional management. Table 2.9 Attentional systems. Table 2.10 Instrumental learning in the basal ganglia. Table 2.11 Reward and motivational systems……………………………………. 631 Table 2.12 A summary of the domains of executive function. Table 2.13 Systems for cognition, consideration of consequences, planning, goal election and executive direction. Table 2.14 Systems for effective execution of motor commands. Table 2.15 Summary of mechanisms for neuronal structural plasticity discussed in Chapter 2……………………………………………………… 643 Table 3.1 The Human Act (This table is included in Chapter 3.) Table 4.1 Characteristics pertaining to the state of virtue, as proposed by Aristotle and Aquinas. (This table is included in Chapter 4.) Table 5.1 Neural subdivisions implicated in moral behaviour…………… 648 Table 5.2 A summary of the principal brain systems contributing to the development and exercise of virtue…………………………… 651 Table 5.3 A simplified neural analysis of Scenario 3. (This table is included in Chapter 5.) Table 6.1 A notional distribution of neural bases of eudaimonia. (This table is included in Chapter 6.)

Reference List 655

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Abstract

Virtues, as described by Aristotle and Aquinas, are understood as dispositions of character to behave in habitual, specific, positive ways; virtue is a critical requirement for human flourishing. From the perspective of Aristotelian-Thomistic anthropology which offers an integrated vision of the material and the rational in the human person, I seek to identify the neural bases for the development and exercise of moral virtue. First I review current neuroscientific knowledge of the capacity of the brain to structure according to experience, to facilitate behaviours, to regulate emotional responses and support goal election. Then, having identified characteristics of moral virtue in the light of the distinctions between cardinal virtues, I propose neural substrates by mapping neuroscientific knowledge to these characteristics. I then investigate the relationship between virtue, including its neurobiological features, and human flourishing. This process allows a contemporary and evidence-based corroboration for a model of moral development based on growth in virtue as understood by Aristotle and Aquinas, and a demonstration of a biological aptitude and predisposition for the development of virtue. Conclusions are drawn with respect to science, ethics, and parenting.

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Declaration of Authorship

This thesis is my own work and contains no material which has been accepted for the award of any degree or diploma in any other institution. To the best of my knowledge, the thesis contains no material previously published or written by another person, except where due reference is made in the text of the thesis.

Andrew P.J. Mullins 20 December 2012

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Acknowledgements

Fresh out of school I was visiting our old retired parish priest, Fr Tosi. When I told him I was thinking of teaching, he boomed, “Blessed are those who instruct others in the paths of virtue, they shall shine like bright stars.” Well, in life I have had many stars that have not given up on me. As for any wanderings from the path, they are of my doing alone!

In this project I have many stars to thank. I am deeply grateful to my supervisor Professor Hayden Ramsay. Hayden took this on despite desperately busy commitments. Without Hayden’s reassurance and in this project, it most certainly would not have materialized. To a large degree, any success it may enjoy will be due to his patient, good humoured guidance. I am also greatly endebted to Professor Julie Quinlivan, my neuroscience supervisor, who provided crucial direction in overseeing my neuroscience research, and to Dr Paul Foley of Neuroscience Research Australia, the NeuRA institute. Paul has shown genuine interest in this undertaking and has generously given of his expertise over the past year. His close knowledge of the basal ganglia and dopamine systems in the brain has been invaluably reassuring.

Others must not be forgotten. It was Professor George Mendz who sowed the seeds of further study when it was the last thing on my mind. And the university has been good to me: a grant made possible a short and very productive stay in Oxford this year. My gratitude also to Gay-Louise, Hayden’s secretary, for her untiring, nothing’s-a-problem capacity to run off reams of drafts and cut through Chancellery red tape to schedule appointments with Hayden; and to Adele at the medical library who, magically and with a smile, tracked down countless papers. My employer, the Parents for Education Foundation, allowed me some ongoing time; and Matt Aldous, my deputy at Wollemi College, took on the busiest of terms to allow me leave to tie up the many loose ends for submission. I will long remember the staff and students of the Veech Library in Strathfield for making this home stretch a delightful experience.

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Friends have offered feedback on the drafts of chapters and sections of chapters. I thank especially Dr Robin Choong for his generosity and for stretching my thinking time and again, Dr Thomas Lickona and John Poon for psychological insights, and Dr Michael Cook, for the right word at the right time. I thank also the parents and staff of Redfield and Wollemi for their friendship and encouragement.

At this time I remember Mum, whose eyes never lost their sparkle, Dad whose gift of the telescope represented something far more, and St. Josemaria who put the star chart of the virtues in my hand. I thank Michael and Richard, the best sounding boards in the business, and Rosemary who is, now and always, the brightest of stars in the firmament.

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5-HT 5-hydroxytryptamine (serotonin) ACC anterior cingulated cortex ACh acetylcholine ADHD Attention Deficit Hyperactivity Disorder AMPA α-amino-3-hydroxyl-5-methyl-4-isoxazole-propionate AMPAR AMPA receptor A-O action-outcome aPFC anterior PFC aTL anterior temporal lobes ATP adenosine triphosphate BDNF brain derived neurotrophic factor BG basal ganglia BLA basolateral complex of the amygdala BOLD blood oxygen level dependent cAMP cyclic AMP (AMP is agonist for AMPA channels) CAT computerised axial tomography CEA central nucleus of the amygdala CNS central nervous system CREB cAMP response-element binding protein CRF corticotrophin releasing factor DA dopamine DLPFC dorsolateral prefrontal cortex DLS deep limbic system, also, dorsolateral striatum in rodents DMPFC dorsalmedial PFC DMS dorsomedial striatum in rodents DSCAM Downs Syndrome cell adhesion molecule DT-MRI Diffusion tensor magnetic resonance imaging EC endocannabinoid EEG elF2x a particular translation initiation factor Epi epinephrine EPSP excitatory postsynaptic potential ERP event related potentials FAPs fixed action patterns FGF fibroblast growth factor FGF-2 fibroblast growth factor 2

18 fMRI functional magnetic resonance imaging GABA -Aminobutyric Acid GP globus pallidus GPi internal segment GPe external segment GPCR G-protein-coupled-receptor HFS high frequency afferent stimulation HSV1 herpes simplex virus type 1 IGF insulin-like growth factor ILN intralaminar nuclei of the thalamus IMM medial mesopalium IT inferotemporal cortex ITM intermediate term memory LA lateral amygdala LC locus ceruleus LGG low grade glioma LIP lateral intraparietal cortex LOFC lateral OFC LTD long term depression LTM long term memory LTP long term potentiation M1 primary motor cortex MA motor adaptation MEG magnetoelectroencephalography MRI magnetic resonance imaging MRS magnetic resonance spectroscopy MSL motor sequence learning MSN medium spiny neuron MTL medial temporal lobe mPFC medial PFC (in some literature, the rodent equivalent to human DLPFC) NAc nucleus accumbens nAChR nicotinic receptor NE norepinephrine NMDA N-methyl-D-aspartate NMJ neuromuscular junction NMP neuromotor prosthesis NMR-MOUSE nuclear magnetic resonance mobile universal surface explorer NOIR near optical infra red

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NTF nerve growth factor OCD obsessive compulsive disorder OFC orbitofrontal cortex PAS paired associative stimulation PCD programmed cell death PET positron emission tomography PKA and PKC particular protein kinases PNS peripheral nervous system PPC posterior parietal cortex preNMDARs presynaptic NMDARs PSP post synaptic potential pSTS post central gyrus rTMS repetitive transcranial magnetic stimulation SACD social and character development programs SMA supplementary motor area S-O stimulus-outcome SN substantia nigra SNpc substantia nigra pars compacta SNr substantia nigra pars reticulata STDP spike timing dependent plasticities S-R stimulus response S-S stimulus-stimulus STN subthalamic nuclei STS posterior superior temporal sulcus TAN tonically active neuron tDCS transcranial direct current stimulation tLTD spike timing-dependent LTD TMS transcranial magnetic stimulation VBM voxel based morphometry VGCC voltage gated calcium channel VL ventrolateral nucleus of the thalamus VLPFC ventrolateral PFC VMPFC ventromedial PFC VP ventral palladium VTA ventral tegmental area

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