Download Emotion Explained

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
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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
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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'
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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?
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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
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The
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4.6
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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
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4.11
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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
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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
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Thirst
6.1 Introduction
6.2 Cellular stimuli for drinking
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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
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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
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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
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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
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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
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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
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Glossary
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References
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Index
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