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Emotion Explained Edmund T. Rolls Professor of Experimental Psychology, University of Oxford and Fellow and Tutor in Psychology, Corpus Christi College, Oxford OXPORD UNIVERSITY PRESS Contents 1 Introduction: the issues 1.1 Introduction 1.2 Rewards and punishers 1.3 The approaches taken to emotion and motivation 1.4 The plan of the book 2 The 2.1 2.2 2.3 2.4 2.5 2.6 nature of emotion Introduction A theory of emotion Different emotions Refinements of the theory of emotion The classification of emotion Other theories of emotion 2.6.1 The James-Lange and other bodily theories 2.6.2 Appraisal theory 2.6.3 Dimensional and categorical theories of emotion 2.6.4 Other approaches to emotion 2.7 Individual differences in emotion, personality, and emotional intelligence 2.8 Cognition and Emotion 2.9 Emotion, motivation, reward, and mood 2.10 The concept of emotion 2.11 Advantages of the approach to emotion described here (Rolls' theory of emotion) 3 The functions of emotion: reward, punishment, and emotion in brain design 3.1 Introduction 3.2 Brain design and the functions of emotion 3.2.1 Taxes, rewards, and punishers: gene-specified goals for actions, and the flexibility of actions 3.2.2 Explicit systems, language, and reinforcement 3.2.3 Special-purpose design by an external agent vs evolution by natural selection 3.3 Selection of behaviour: cost-benefit 'analysis' 1 1 2 5 7 10 10 11 13 21 25 26 26 30 31 31 32 35 36 37 38 41 41 43 43 47 48 49 x I Contents 3.4 Further functions of emotion 3.4.1 Autonomie and endocrine responses 3.4.2 Flexibility of behavioural responses 3.4.3 Emotional states are motivating 3.4.4 Communication 3.4.5 Social attachment 3.4.6 Separate functions for each different primary reinforcer 3.4.7 The mood state can influence the cognitive evaluation of moods or memories 3.4.8 Facilitation of memory storage 3.4.9 Emotional and mood states are persistent, and help to produce persistent motivation 3.4.10 Emotions may trigger memory recall and influence cognitive processing The functions of emotion in an evolutionary, Darwinian, context The functions of motivation in an evolutionary, Darwinian, context Are all goals for action gene-specified? 51 51 52 53 54 57 brain mechanisms underlying emotion Introduction Overview Representations of primary reinforcers 4.3.1 Taste 4.3.2 Smell 4.3.3 Pleasant and painful touch 4.3.4 Visual stimuli 4.4 Representing potential secondary reinforcers 4.4.1 The requirements of the representation 4.4.2 High capacity 4.4.3 Objects, and not their reward and punishment associations, are represented in the inferior temporal visual cortex 4.4.4 Object representations 4.4.5 Invariant representations of faces and objects in the inferior temporal visual cortex 4.4.6 Face expression, gesture and view represented in a population of neurons in the cortex in the superior 63 63 63 66 67 67 67 69 71 71 74 3.5 3.6 3.7 The 4.1 4.2 4.3 57 58 58 59 59 59 61 62 75 77 78 Contents | xi 4.5 4.6 4.7 4.8 4.9 temporal sulcus 4.4.7 The brain mechanisms that build the appropriate view-invariant representations of objects required for learning emotional responses to objects, including faces The orbitofrontal cortex 4.5.1 Historical background 4.5.2 Topology 4.5.3 Connections 4.5.4 Effects of damage to the orbitofrontal cortex 4.5.5 Neurophysiology and functional neuroimaging ofthe orbitofrontal cortex 4.5.6 The human orbitofrontal cortex 4.5.7 A neurophysiological and computational basis for stimulus-reinforcer association learning and reversal in the orbitofrontal cortex 4.5.8 Executive functions of the orbitofrontal cortex The amygdala 4.6.1 Associative processes involved in emotion-related learning 4.6.2 Connections of the amygdala 4.6.3 Effects of amygdala lesions 4.6.4 Neuronal activity in the primate amygdala to reinforcing stimuli 4.6.5 Responses of these amygdala neurons to novel stimuli that are reinforcing 4.6.6 Neuronal responses in the amygdala to faces 4.6.7 Evidence from humans 4.6.8 Amygdala summary The cingulate cortex 4.7.1 Perigenual cingulate cortex and affect 4.7.2 Mid-cingulate cortex, the cingulate motor area, and action-outcome learning Human brain imaging investigations of mood and depression Output pathways for emotional responses 4.9.1 The autonomie and endocrine systems 4.9.2 Motor systems for implicit responses, including the basal ganglia 89 89 91 91 92 93 95 97 131 140 147 149 149 155 157 164 170 172 174 178 179 181 185 187 188 188 189 xii I Contents 4.10 4.11 4.12 4.13 4.9.3 Output systems for explicit responses to emotional stimuli 4.9.4 Basal forebrain and hypothalamus 4.9.5 Basal forebrain cholinergic neurons 4.9.6 Noradrenergic neurons Effects of emotion on cognitive processing and memory Laterality effects in human emotional processing Summary Colour plates 190 191 191 194 194 200 202 205 Hunger 5.1 Introduction 5.2 Peripheral signals for hunger and satiety 5.3 The control signals for hunger and satiety 5.3.1 Sensory-specific satiety 5.3.2 Gastric distension 5.3.3 Duodenal chemosensors 5.3.4 Glucostatic hypothesis 5.3.5 Body fat regulation - leptin or OB protein 5.3.6 Conditioned appetite and satiety 5.4 The brain control of eating and reward 5.4.1 The hypothalamus 5.4.2 Brain mechanisms for the reward produced by the taste of food 5.4.3 Convergence between taste and olfactory processing to represent flavour 5.4.4 Brain mechanisms for the reward produced by the odour of food 5.4.5 The responses of orbitofrontal cortex taste and olfactory neurons to the sight of food 5.4.6 Functions of the amygdala and temporal cortex in feeding 5.4.7 Functions of the orbitofrontal cortex in feeding 5.4.8 Functions of the striatum in feeding 5.5 Obesity, bulimia, and anorexia 5.6 Conclusions on reward, affective responses to food, and the control of appetite 221 221 221 224 224 230 230 230 231 232 233 233 Thirst 6.1 Introduction 6.2 Cellular stimuli for drinking 274 274 275 243 253 254 259 259 263 266 271 273 Contents | xiii 6.3 6.4 6.5 6.6 7 Extracellular thirst stimuli 6.3.1 Extracellular stimuli for thirst 6.3.2 Role of the kidney in extracellular thirst: the reninangiotensin system 6.3.3 Cardiac receptors for thirst Control of normal drinking Reward and satiety signals for drinking Summary Brain-stimulation reward 7.1 Introduction 7.2 The nature of the reward produced 7.3 The location of brain-stimulation reward sites in the brain 7.4 The effects of brain lesions on intracranial self-stimulation 7.5 The neurophysiology of reward 7.5.1 Lateral hypothalamus and substantia innominata 7.5.2 Orbitofrontal cortex 7.5.3 Amygdala 7.5.4 Nucleus accumbens 7.5.5 Central gray of the midbrain 7.6 Some of the properties of brain-stimulation reward 7.6.1 Lack of satiety with brain-stimulation reward 7.6.2 Rapid extinction 7.6.3 Priming 7.7 Stimulus-bound motivational behaviour 7.8 Conclusions 7.9 Apostasis 8 Pharmacology of emotion, reward, and addiction; the basal ganglia 8.1 Introduction 8.2 The noradrenergic hypothesis 8.3 Dopamine and reward 8.3.1 Dopamine and electrical self-stimulation of the brain 8.3.2 Self-administration of dopaminergic substances, and addiction 8.3.3 Behaviours associated with the release of dopamine 8.3.4 The activity of dopaminergic neurons and reward 8.4 The basal ganglia 8.4.1 Systems-level architecture of the basal ganglia 8.4.2 Effects of basal ganglia damage 276 276 278 279 279 282 286 288 288 288 292 293 294 294 296 298 299 299 300 300 302 302 304 305 306 308 308 311 312 312 314 316 318 321 322 323 xiv I Contents 8.5 8.6 8.7 8.8 8.9 8.4.3 Neuronal activity in the striatum 8.4.4 What computations are performed by the basal ganglia? 8.4.5 How do the basal ganglia perform their computations? 8.4.6 Synthesis on the role of dopamine in reward and addiction 8.4.7 Synthesis: emotion, dopamine, reward, punishment, and action selection in the basal ganglia Opiate reward systems, analgesia, and food reward Pharmacology of depression in relation to brain systems involved in emotion Pharmacology of anxiety in relation to brain systems involved in emotion Cannabinoids Overview of behavioural selection and output systems involved in emotion 9 Sexual behaviour, reward, and brain function; sexual selection of behaviour 9.1 Introduction 9.2 Mate selection, attractiveness, and love 9.2.1 Female preferences 9.2.2 Male preferences 9.2.3 Pair-bonding, and Love 9.3 Parental attachment, care, and parent-offspring conflict 9.4 Sperm competition and its consequences for sexual behaviour 9.5 Concealed ovulation and its consequences for sexual behaviour 9.6 Sexual selection of sexual and non-sexual behaviour 9.6.1 Sexual selection and natural selection 9.6.2 Non-sexual characteristics may be sexually selected for courtship 9.7 Individual differences in sexual rewards 9.7.1 Overview 9.7.2 How might different types of behaviour be produced by natural selection altering the relative reward value of different stimuli in different individuals? 9.7.3 How being tuned to different types of reward could 325 339 340 349 350 352 353 354 355 355 358 358 360 361 363 366 367 368 375 376 376 379 381 381 384 Contents | xv help to produce individual differences in sexual behaviour 9.8 The neural reward mechanisms that might mediate some aspects of sexual behaviour 9.9 Neural basis of sexual behaviour 9.10 Conclusion 10 Emotional feelings and consciousness: a theory of consciousness 10.1 Introduction 10.2 A theory of consciousness 10.3 Dual routes to action 10.4 Content and meaning in representations 10.5 Discussion 10.6 Conclusions and comparisons 11 Conclusions, and broader issues 11.1 Conclusions 11.2 Decision-making 11.2.1 Selection of mainly autonomic responses, and their classical conditioning 11.2.2 Selection of approach or withdrawal, and their classical conditioning 11.2.3 Selection of fixed stimulus-response habits 11.2.4 Selection of arbitrary behaviours to obtain goals, action-outcome learning, and emotional learning 11.2.5 The roles of the prefrontal cortex in decision-making and attention 11.2.6 Neuroeconomics, reward value, and expected utility 11.2.7 Selection of actions by explicit rational thought 11.3 Emotion and ethics 11.4 Emotion and literature A 386 387 395 398 400 400 401 411 418 420 423 426 426 431 431 431 432 432 433 440 444 445 449 11.5 Close 452 Neural networks and emotion-related learning A.1 Neurons in the brain, the representation of information, and neuronal learning mechanisms A.1.1 Introduction A.1.2 Neurons in the brain, and their representation in neuronal networks 454 454 454 454 xvi I Contents A.2 A.3 A.4 A.5 B A. 1.3 A formalism for approaching the operation of single neurons in a network A. 1.4 Synaptic modification A. 1.5 Long-Term Potentiation and Long-Term Depression A. 1.6 Distributed representations Pattern association memory A.2.1 Architecture and operation A.2.2 A simple model A.2.3. The vector interpretation A.2.4 Properties A.2.5 Prototype extraction, extraction of central tendency, and noise reduction A.2.6 Speed A.2.7 Local learning rule A.2.8 Implications of different types of coding for storage in pattern associators Autoassociation memory: attractor networks A.3.1 Architecture and operation A.3.2 Introduction to the analysis of the operation of autoassociation networks A.3.3 Properties Coupled attractor networks Reinforcement learning A.5.1 Associative reward-penalty algorithm of Barto and Sutton A.5.2 Error correction or delta rule learning, and classical conditioning A.5.3 Temporal Difference (TD) learning Reward reversal in the orbitofrontal cortex - a model B.1 Introduction B.2 The model of stimulus-reinforcer association reversal B.2.1 The network B.2.2 Reward reversai: the operation of the rule module neurons B.2.3 The neurons in the model B.2.4 The synapses in the model B.3 Operation of the reward reversal model B.4 A model of reversal of a conditional object-response task by the dorsolateral prefrontal cortex 456 458 459 464 466 466 469 472 472 475 475 476 482 483 483 485 486 491 493 494 496 497 501 501 503 504 507 509 510 511 515 Contents I xvii C B.5 Evaluation ofthe models B.6 Integrate-and-Fire model equations and parameters B.7 Simulation of fMRI signals: haemodynamic convolution of synaptic activity 517 521 Glossary 525 522 References 528 Index 601