Emotion Explained

by Rolls, Edmund T.

• ISBN13: 9780198570035
• ISBN10: 0198570031
• Format: Hardcover
• Copyright: 2005-10-20
• Publisher: Oxford University Press

Summary

What produces emotions? Why do we have emotions? How do we have emotions?Why do emotional states feel like something? This book seeks explanations ofemotion by considering these questions.Emotion continues to be a topic of enormous scientific interest. This new book,a successor to 'The Brain and Emotion', (OUP, 1998), describes the nature,functions, and brain mechanisms that underlie both emotion and motivation.'Emotion Explained' goes beyond examining brain mechanisms of emotion, byproposing a theory of what emotions are, and an evolutionary, Darwinian, theoryof the adaptive value of emotion. It also shows that there is a clearrelationship between motivation and emotion. The book also examines howcognitive states can modulate emotions, and in turn, how emotions can influencecognitive states. It considers the role of sexual selection in the evolution ofaffective behaviour. It also examines emotion and decision making, with links tothe burgeoning field of neuroeconomics. The book is also unique in consideringemotion at several levels - the neurophysiological, neuroimaging,neuropsychological, behavioural, and computational neuroscience levels.

Author Biography

Edmund T. Rolls is Professor of Experimental Psychology at the University of Oxford, and a Fellow and Tutor of Corpus Christi College, Oxford. He read preclinical medicine at the University of Cambridge, and now performs research in neuroscience at Oxford. His research links neurophysiological and computational neuroscience approaches to human functional neuroimaging and neuropsychological studies in order to provide a fundamental basis for understanding human brain function and its disorders. He is author of The Brain and Emotion (1999, Oxford University Press), with A.Treves of Neural Networks and Brain Function (1998, Oxford University Press), and with G.Deco of Computational Neuroscience of Vision (2002, Oxford University Press).

Table of Contents

1 Introduction: the issues	1	(9)
1.1 Introduction	1	(1)
1.2 Rewards and punishers	2	(3)
1.3 The approaches taken to emotion and motivation	5	(2)
1.4 The plan of the book	7	(3)
2 The nature of emotion	10	(31)
2.1 Introduction	10	(1)
2.2 A theory of emotion	11	(2)
2.3 Different emotions	13	(8)
2.4 Refinements of the theory of emotion	21	(4)
2.5 The classification of emotion	25	(1)
2.6 Other theories of emotion	26	(6)
2.6.1 The James–Lange and other bodily theories	26	(4)
2.6.2 Appraisal theory	30	(1)
2.6.3 Dimensional and categorical theories of emotion	31	(1)
2.6.4 Other approaches to emotion	31	(1)
2.7 Individual differences in emotion, personality, and emotional intelligence	32	(3)
2.8 Cognition and Emotion	35	(1)
2.9 Emotion, motivation, reward, and mood	36	(1)
2.10 The concept of emotion	37	(1)
2.11 Advantages of the approach to emotion described here (Rolls' theory of emotion)	38	(3)
3 The functions of emotion: reward, punishment, and emotion in brain design	41	(22)
3.1 Introduction	41	(2)
3.2 Brain design and the functions of emotion	43	(6)
3.2.1 Taxes, rewards, and punishers: gene-specified goals for actions, and the flexibility of actions	43	(4)
3.2.2 Explicit systems, language, and reinforcement	47	(1)
3.2.3 Special-purpose design by an external agent vs evolution by natural selection	48	(1)
3.3 Selection of behaviour: cost–benefit 'analysis'	49	(2)
3.4 Further functions of emotion	51	(8)
3.4.1 Autonomic and endocrine responses	51	(1)
3.4.2 Flexibility of behavioural responses	52	(1)
3.4.3 Emotional states are motivating	53	(1)
3.4.4 Communication	54	(3)
3.4.5 Social attachment	57	(1)
3.4.6 Separate functions for each different primary reinforcer	57	(1)
3.4.7 The mood state can influence the cognitive evaluation of moods or memories	58	(1)
3.4.8 Facilitation of memory storage	58	(1)
3.4.9 Emotional and mood states are persistent, and help to produce persistent motivation	59	(1)
3.4.10 Emotions may trigger memory recall and influence cognitive processing	59	(1)
3.5 The functions of emotion in an evolutionary, Darwinian, context	59	(2)
3.6 The functions of motivation in an evolutionary, Darwinian, context	61	(1)
3.7 Are all goals for action gene-specified?	62	(1)
4 The brain mechanisms underlying emotion	63	(158)
4.1 Introduction	63	(1)
4.2 Overview	63	(3)
4.3 Representations of primary reinforcers	66	(5)
4.3.1 Taste	67	(1)
4.3.2 Smell	67	(1)
4.3.3 Pleasant and painful touch	67	(2)
4.3.4 Visual stimuli	69	(2)
4.4 Representing potential secondary reinforcers	71	(20)
4.4.1 The requirements of the representation	71	(3)
4.4.2 High capacity	74	(1)
4.4.3 Objects, and not their reward and punishment associations, are represented in the inferior temporal visual cortex	75	(2)
4.4.4 Object representations	77	(1)
4.4.5 Invariant representations of faces and objects in the inferior temporal visual cortex	78	(11)
4.4.6 Face expression, gesture and view represented in a population of neurons in the cortex in the superior temporal sulcus	89	(1)
4.4.7 The brain mechanisms that build the appropriate view-invariant representations of objects required for learning emotional responses to objects, including faces	89	(2)
4.5 The orbitofrontal cortex	91	(58)
4.5.1 Historical background	91	(1)
4.5.2 Topology	92	(1)
4.5.3 Connections	93	(2)
4.5.4 Effects of damage to the orbitofrontal cortex	95	(2)
4.5.5 Neurophysiology and functional neuroimaging of the orbitofrontal cortex	97	(34)
4.5.6 The human orbitofrontal cortex	131	(9)
4.5.7 A neurophysiological and computational basis for stimulus–reinforcer association learning and reversal in the orbitofrontal cortex	140	(7)
4.5.8 Executive functions of the orbitofrontal cortex	147	(2)
4.6 The amygdala	149	(30)
4.6.1 Associative processes involved in emotion-related learning	149	(6)
4.6.2 Connections of the amygdala	155	(2)
4.6.3 Effects of amygdala lesions	157	(7)
4.6.4 Neuronal activity in the primate amygdala to reinforcing stimuli	164	(6)
4.6.5 Responses of these amygdala neurons to novel stimuli that are reinforcing	170	(2)
4.6.6 Neuronal responses in the amygdala to faces	172	(2)
4.6.7 Evidence from humans	174	(4)
4.6.8 Amygdala summary	178	(1)
4.7 The cingulate cortex	179	(8)
4.7.1 Perigenual cingulate cortex and affect	181	(4)
4.7.2 Mid-cingulate cortex, the cingulate motor area, and action–outcome learning	185	(2)
4.8 Human brain imaging investigations of mood and depression	187	(1)
4.9 Output pathways for emotional responses	188	(6)
4.9.1 The autonomic and endocrine systems	188	(1)
4.9.2 Motor systems for implicit responses, including the basal ganglia	189	(1)
4.9.3 Output systems for explicit responses to emotional stimuli	190	(1)
4.9.4 Basal forebrain and hypothalamus	191	(1)
4.9.5 Basal forebrain cholinergic neurons	191	(3)
4.9.6 Noradrenergic neurons	194	(1)
4.10 Effects of emotion on cognitive processing and memory	194	(6)
4.11 Laterality effects in human emotional processing	200	(2)
4.12 Summary	202	(3)
4.13 Colour plates	205	(16)
5 Hunger	221	(53)
5.1 Introduction	221	(1)
5.2 Peripheral signals for hunger and satiety	221	(3)
5.3 The control signals for hunger and satiety	224	(9)
5.3.1 Sensory-specific satiety	224	(6)
5.3.2 Gastric distension	230	(1)
5.3.3 Duodenal chemosensors	230	(1)
5.3.4 Glucostatic hypothesis	230	(1)
5.3.5 Body fat regulation – leptin or OB protein	231	(1)
5.3.6 Conditioned appetite and satiety	232	(1)
5.4 The brain control of eating and reward	233	(38)
5.4.1 The hypothalamus	233	(10)
5.4.2 Brain mechanisms for the reward produced by the taste of food	243	(10)
5.4.3 Convergence between taste and olfactory processing to represent flavour	253	(1)
5.4.4 Brain mechanisms for the reward produced by the odour of food	254	(5)
5.4.5 The responses of orbitofrontal cortex taste and olfactory neurons to the sight of food	259	(1)
5.4.6 Functions of the amygdala and temporal cortex in feeding	259	(4)
5.4.7 Functions of the orbitofrontal cortex in feeding	263	(3)
5.4.8 Functions of the striatum in feeding	266	(5)
5.5 Obesity, bulimia, and anorexia	271	(2)
5.6 Conclusions on reward, affective responses to food, and the control of appetite	273	(1)
6 Thirst	274	(14)
6.1 Introduction	274	(1)
6.2 Cellular stimuli for drinking	275	(1)
6.3 Extracellular thirst stimuli	276	(3)
6.3.1 Extracellular stimuli for thirst	276	(2)
6.3.2 Role of the kidney in extracellular thirst: the reninangiotensin system	278	(1)
6.3.3 Cardiac receptors for thirst	279	(1)
6.4 Control of normal drinking	279	(3)
6.5 Reward and satiety signals for drinking	282	(4)
6.6 Summary	286	(2)
7 Brain-stimulation reward	288	(20)
7.1 Introduction	288	(1)
7.2 The nature of the reward produced	288	(4)
7.3 The location of brain-stimulation reward sites in the brain	292	(1)
7.4 The effects of brain lesions on intracranial self-stimulation	293	(1)
7.5 The neurophysiology of reward	294	(6)
7.5.1 Lateral hypothalamus and substantia innominata	294	(2)
7.5.2 Orbitofrontal cortex	296	(2)
7.5.3 Amygdala	298	(1)
7.5.4 Nucleus accumbens	299	(1)
7.5.5 Central gray of the midbrain	299	(1)
7.6 Some of the properties of brain-stimulation reward	300	(4)
7.6.1 Lack of satiety with brain-stimulation reward	300	(2)
7.6.2 Rapid extinction	302	(1)
7.6.3 Priming	302	(2)
7.7 Stimulus-bound motivational behaviour	304	(1)
7.8 Conclusions	305	(1)
7.9 Apostasis	306	(2)
8 Pharmacology of emotion, reward, and addiction; the basal ganglia	308	(50)
8.1 Introduction	308	(3)
8.2 The noradrenergic hypothesis	311	(1)
8.3 Dopamine and reward	312	(9)
8.3.1 Dopamine and electrical self-stimulation of the brain	312	(2)
8.3.2 Self-administration of dopaminergic substances, and addiction	314	(2)
8.3.3 Behaviours associated with the release of dopamine	316	(2)
8.3.4 The activity of dopaminergic neurons and reward	318	(3)
8.4 The basal ganglia	321	(31)
8.4.1 Systems-level architecture of the basal ganglia	322	(1)
8.4.2 Effects of basal ganglia damage	323	(2)
8.4.3 Neuronal activity in the striatum	325	(14)
8.4.4 What computations are performed by the basal ganglia?	339	(1)
8.4.5 How do the basal ganglia perform their computations?	340	(9)
8.4.6 Synthesis on the role of dopamine in reward and addiction	349	(1)
8.4.7 Synthesis: emotion, dopamine, reward, punishment, and action selection in the basal ganglia	350	(2)
8.5 Opiate reward systems, analgesia, and food reward	352	(1)
8.6 Pharmacology of depression in relation to brain systems involved in emotion	353	(1)
8.7 Pharmacology of anxiety in relation to brain systems involved in emotion	354	(1)
8.8 Cannabinoids	355	(1)
8.9 Overview of behavioural selection and output systems involved in emotion	355	(3)
9 Sexual behaviour, reward, and brain function; sexual selection of behaviour	358	(42)
9.1 Introduction	358	(2)
9.2 Mate selection, attractiveness, and love	360	(7)
9.2.1 Female preferences	361	(2)
9.2.2 Male preferences	363	(3)
9.2.3 Pair-bonding, and Love	366	(1)
9.3 Parental attachment, care, and parent–offspring conflict	367	(1)
9.4 Sperm competition and its consequences for sexual behaviour	368	(7)
9.5 Concealed ovulation and its consequences for sexual behaviour	375	(1)
9.6 Sexual selection of sexual and non-sexual behaviour	376	(5)
9.6.1 Sexual selection and natural selection	376	(3)
9.6.2 Non-sexual characteristics may be sexually selected for courtship	379	(2)
9.7 Individual differences in sexual rewards	381	(6)
9.7.1 Overview	381	(3)
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?	384	(2)
9.7.3 How being tuned to different types of reward could help to produce individual differences in sexual behaviour	386	(1)
9.8 The neural reward mechanisms that might mediate some aspects of sexual behaviour	387	(8)
9.9 Neural basis of sexual behaviour	395	(3)
9.10 Conclusion	398	(2)
10 Emotional feelings and consciousness: a theory of consciousness	400	(26)
10.1 Introduction	400	(1)
10.2 A theory of consciousness	401	(10)
10.3 Dual routes to action	411	(7)
10.4 Content and meaning in representations	418	(2)
10.5 Discussion	420	(3)
10.6 Conclusions and comparisons	423	(3)
11 Conclusions, and broader issues	426	(28)
11.1 Conclusions	426	(5)
11.2 Decision-making	431	(14)
11.2.1 Selection of mainly autonomic responses, and their classical conditioning	431	(1)
11.2.2 Selection of approach or withdrawal, and their classical conditioning	431	(1)
11.2.3 Selection of fixed stimulus–response habits	432	(1)
11.2.4 Selection of arbitrary behaviours to obtain goals, action–outcome learning, and emotional learning	432	(1)
11.2.5 The roles of the prefrontal cortex in decision-making and attention	433	(7)
11.2.6 Neuroeconomics, reward value, and expected utility	440	(4)
11.2.7 Selection of actions by explicit rational thought	444	(1)
11.3 Emotion and ethics	445	(4)
11.4 Emotion and literature	449	(3)
11.5 Close	452	(2)
A Neural networks and emotion-related learning	454	(47)
A.1 Neurons in the brain, the representation of information, and neuronal learning mechanisms	454	(12)
A.1.1 Introduction	454	(1)
A.1.2 Neurons in the brain, and their representation in neuronal networks	454	(2)
A.1.3 A formalism for approaching the operation of single neurons in a network	456	(2)
A.1.4 Synaptic modification	458	(1)
A.1.5 Long-Term Potentiation and Long-Term Depression	459	(5)
A.1.6 Distributed representations	464	(2)
A.2 Pattern association memory	466	(17)
A.2.1 Architecture and operation	466	(3)
A.2.2 A simple model	469	(3)
A.2.3 The vector interpretation	472	(1)
A.2.4 Properties	472	(3)
A.2.5 Prototype extraction, extraction of central tendency, and noise reduction	475	(1)
A.2.6 Speed	475	(1)
A.2.7 Local learning rule	476	(6)
A.2.8 Implications of different types of coding for storage in pattern associators	482	(1)
A.3 Autoassociation memory: attractor networks	483	(8)
A.3.1 Architecture and operation	483	(2)
A.3.2 Introduction to the analysis of the operation of autoassociation networks	485	(1)
A.3.3 Properties	486	(5)
A.4 Coupled attractor networks	491	(2)
A.5 Reinforcement learning	493	(8)
A.5.1 Associative reward–penalty algorithm of Barto and Sutton	494	(2)
A.5.2 Error correction or delta rule learning, and classical conditioning	496	(1)
A.5.3 Temporal Difference (TD) learning	497	(4)
B Reward reversal in the orbitofrontal cortex – a model	501	(24)
B.1 Introduction	501	(2)
B.2 The model of stimulus–reinforcer association reversal	503	(8)
B.2.1 The network	504	(3)
B.2.2 Reward reversal: the operation of the rule module neurons	507	(2)
B.2.3 The neurons in the model	509	(1)
B.2.4 The synapses in the model	510	(1)
B.3 Operation of the reward reversal model	511	(4)
B.4 A model of reversal of a conditional object-response task by the dorsolateral prefrontal cortex	515	(2)
B.5 Evaluation of the models	517	(4)
B.6 Integrate-and-Fire model equations and parameters	521	(1)
B.7 Simulation of fMRI signals: haemodynamic convolution of synaptic activity	522	(3)
C Glossary	525	(3)
References	528	(73)
Index	601