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1 Introduction: the issues |
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1 | (12) |
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1 | (1) |
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1.2 Rewards and punishers |
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2 | (2) |
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1.3 The approaches taken to emotion and motivation |
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4 | (6) |
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10 | (3) |
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13 | (32) |
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13 | (1) |
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14 | (3) |
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17 | (7) |
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2.4 Refinements of the theory of emotion |
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24 | (4) |
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2.5 The classification of emotion |
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28 | (1) |
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2.6 Other theories of emotion |
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29 | (7) |
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2.6.1 The James-Lange and other bodily theories |
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30 | (3) |
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33 | (2) |
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2.6.3 Dimensional and categorical theories of emotion |
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35 | (1) |
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2.6.4 Other approaches to emotion |
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35 | (1) |
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2.7 Individual differences in emotion, personality, and emotional intelligence |
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36 | (3) |
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2.8 Cognition and emotion |
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39 | (1) |
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2.9 Emotion, motivation, reward, and mood |
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40 | (1) |
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2.10 The concept of emotion |
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41 | (1) |
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2.11 Advantages of the approach to emotion described here (Rolls' theory of emotion) |
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42 | (3) |
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3 The functions of emotion: reward, punishment, and emotion in brain design |
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45 | (22) |
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45 | (2) |
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3.2 Brain design and the functions of emotion |
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47 | (6) |
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3.2.1 Taxes, rewards, and punishers: gene-specified goals for actions, and the flexibility of actions |
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47 | (4) |
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3.2.2 Explicit systems, language, and reinforcement |
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51 | (1) |
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3.2.3 Special-purpose design by an external agent vs evolution by natural selection |
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51 | (2) |
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3.3 Selection of behaviour: cost-benefit `analysis' of net value |
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53 | (2) |
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3.4 Further functions of emotion |
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55 | (8) |
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3.4.1 Autonomic and endocrine responses |
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55 | (1) |
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3.4.2 Flexibility of behavioural responses |
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56 | (1) |
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3.4.3 Emotional states are motivating |
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57 | (1) |
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58 | (2) |
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60 | (1) |
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3.4.6 Separate functions for each different primary reinforcer |
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61 | (1) |
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3.4.7 The mood state can influence the cognitive evaluation of moods or memories |
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62 | (1) |
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3.4.8 Facilitation of memory storage |
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62 | (1) |
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3.4.9 Emotional and mood states are persistent, and help to produce persistent motivation |
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62 | (1) |
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3.4.10 Emotions may trigger memory recall and influence cognitive processing |
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62 | (1) |
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3.5 The functions of emotion in an evolutionary, Darwinian, context |
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63 | (2) |
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3.6 The functions of motivation in an evolutionary, Darwinian, context |
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65 | (1) |
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3.7 Are all goals for action gene-specified? |
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65 | (2) |
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4 The brain mechanisms underlying emotion |
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67 | (157) |
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67 | (1) |
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67 | (4) |
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4.3 Representations of primary reinforcers, i.e. of unlearned value |
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71 | (5) |
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71 | (1) |
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71 | (1) |
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4.3.3 Pleasant and painful touch |
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72 | (2) |
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74 | (2) |
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4.4 Representing potential secondary reinforcers |
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76 | (19) |
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4.4.1 The requirements of the representation |
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76 | (4) |
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4.4.2 Objects, and not their reward and punishment associations or value, are represented in the inferior temporal visual cortex |
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80 | (2) |
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4.4.3 Object representations |
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82 | (1) |
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4.4.4 Invariant representations of faces and objects in the inferior temporal visual cortex |
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83 | (11) |
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4.4.5 Face expression, gesture and view represented in a population of neurons in the cortex in the superior temporal sulcus |
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94 | (1) |
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4.4.6 The brain mechanisms that build the appropriate view-invariant representations of objects required for learning emotional responses to objects, including faces |
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94 | (1) |
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4.5 The orbitofrontal cortex |
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95 | (64) |
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4.5.1 Historical background |
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95 | (2) |
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97 | (2) |
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99 | (1) |
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4.5.4 Effects of damage to the orbitofrontal cortex |
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100 | (2) |
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4.5.5 Neurophysiology and functional neuroimaging of the orbitofrontal cortex |
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102 | (40) |
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4.5.6 The human orbitofrontal cortex |
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142 | (9) |
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4.5.7 A neurophysiological and computational basis for stimulusreinforcer association learning and reversal in the orbitofrontal cortex |
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151 | (7) |
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4.5.8 Executive functions of the orbitofrontal cortex |
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158 | (1) |
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159 | (30) |
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4.6.1 Associative processes involved in emotion-related learning |
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159 | (6) |
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4.6.2 Connections of the amygdala |
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165 | (2) |
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4.6.3 Effects of amygdala lesions |
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167 | (7) |
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4.6.4 Neuronal activity in the primate amygdala to reinforcing stimuli |
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174 | (7) |
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4.6.5 Responses of these amygdala neurons to novel stimuli that are reinforcing |
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181 | (1) |
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4.6.6 Neuronal responses in the amygdala to faces |
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182 | (2) |
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4.6.7 Evidence from humans |
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184 | (4) |
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188 | (1) |
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189 | (9) |
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4.7.1 Introduction and overview of the anterior cingulate cortex |
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189 | (2) |
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4.7.2 Anterior cingulate cortex anatomy and connections |
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191 | (1) |
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4.7.3 Anterior cingulate cortex functional neuroimaging and neuronal activity |
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191 | (4) |
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4.7.4 Anterior cingulate cortex lesion effects |
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195 | (1) |
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4.7.5 Mid-cingulate cortex, the cingulate motor area, and action-outcome learning |
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196 | (2) |
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4.8 Value-related decision-making and medial prefrontal cortex area 10 |
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198 | (2) |
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4.8.1 Decision-making between the value of odours |
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198 | (1) |
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4.8.2 Decision-making between the value of thermal somatosensory stimuli |
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199 | (1) |
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4.8.3 Value-related decision-making and the medial prefrontal cortex area 10: further evidence |
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199 | (1) |
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200 | (3) |
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4.10 Human brain imaging investigations of mood and depression |
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203 | (5) |
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4.11 Output pathways for emotional responses |
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208 | (5) |
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4.11.1 The autonomic and endocrine systems |
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208 | (1) |
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4.11.2 Motor systems for implicit responses, including the basal ganglia |
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209 | (1) |
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4.11.3 Output systems for explicit responses to emotional stimuli |
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209 | (1) |
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4.11.4 Basal forebrain and hypothalamus |
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209 | (1) |
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4.11.5 Basal forebrain cholinergic neurons |
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210 | (2) |
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4.11.6 Noradrenergic neurons |
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212 | (1) |
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4.12 Effects of emotion on cognitive processing and memory |
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213 | (5) |
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4.13 Laterality effects in human emotional processing |
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218 | (3) |
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221 | (3) |
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5 Food reward value, pleasure, hunger, and appetite |
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224 | (53) |
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224 | (1) |
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5.2 Peripheral signals for hunger and satiety |
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224 | (3) |
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5.3 The control signals for hunger and satiety |
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227 | (9) |
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5.3.1 Sensory-specific satiety |
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227 | (5) |
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232 | (1) |
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5.3.3 Duodenal chemosensors |
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233 | (1) |
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5.3.4 Glucostatic hypothesis |
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233 | (1) |
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5.3.5 Hormonal signals related to hunger and satiety, and their effects on the hypothalamus |
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233 | (2) |
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5.3.6 Conditioned appetite and satiety |
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235 | (1) |
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5.4 The brain control of eating and reward |
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236 | (33) |
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236 | (8) |
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5.4.2 Brain mechanisms for taste reward value |
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244 | (12) |
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5.4.3 Convergence between taste and olfactory processing to represent flavour |
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256 | (1) |
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5.4.4 Brain mechanisms for the reward produced by the odour of food |
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257 | (5) |
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5.4.5 The responses of orbitofrontal cortex taste and olfactory neurons to the sight of food: expected value neurons |
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262 | (1) |
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5.4.6 Functions of the amygdala and temporal cortex in feeding |
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262 | (4) |
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5.4.7 Functions of the orbitofrontal cortex in feeding |
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266 | (3) |
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5.4.8 Output pathways for feeding |
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269 | (1) |
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5.5 Obesity, bulimia, and anorexia |
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269 | (6) |
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269 | (1) |
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5.5.2 Brain processing of the sensory properties and pleasantness of food |
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270 | (1) |
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271 | (1) |
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5.5.4 Sensory-specific satiety |
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272 | (1) |
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5.5.5 Fixed meal times, and the availability of food |
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272 | (1) |
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5.5.6 Food saliency, and portion size |
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273 | (1) |
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5.5.7 Energy density of food |
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273 | (1) |
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273 | (1) |
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273 | (1) |
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273 | (1) |
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274 | (1) |
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5.5.12 Cognitive factors, and attention |
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274 | (1) |
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5.5.13 Compliance with information about risk factors for obesity |
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274 | (1) |
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5.6 Conclusions on reward, affective responses to food, and the control of appetite |
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275 | (2) |
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6 Pharmacology of emotion, reward, and addiction; the basal ganglia |
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277 | (46) |
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277 | (2) |
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279 | (8) |
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6.2.1 Dopamine and brain-stimulation reward |
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279 | (1) |
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6.2.2 Self-administration of dopaminergic substances, and addiction |
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279 | (2) |
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6.2.3 Behaviours associated with the release of dopamine |
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281 | (1) |
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6.2.4 The activity of dopaminergic neurons and reward |
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282 | (5) |
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287 | (30) |
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6.3.1 Systems-level architecture of the basal ganglia |
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288 | (1) |
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6.3.2 Effects of basal ganglia damage |
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289 | (2) |
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6.3.3 Neuronal activity in the striatum |
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291 | (13) |
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6.3.4 What computations are performed by the basal ganglia? |
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304 | (2) |
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6.3.5 How do the basal ganglia perform their computations? |
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306 | (8) |
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6.3.6 Synthesis on the role of dopamine in reward and addiction |
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314 | (1) |
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6.3.7 Synthesis: emotion, dopamine, reward, punishment, and action selection in the basal ganglia |
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315 | (2) |
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6.4 Opiate reward systems, analgesia, and food reward |
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317 | (1) |
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6.5 Pharmacology of depression in relation to brain systems involved in emotion |
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318 | (1) |
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6.6 Pharmacology of anxiety in relation to brain systems involved in emotion |
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319 | (1) |
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320 | (1) |
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6.8 Overview of behavioural selection and output systems involved in emotion |
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320 | (3) |
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7 Sexual behaviour, reward, and brain function; sexual selection of behaviour |
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323 | (45) |
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323 | (2) |
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7.2 The ultimate explanation for the reward value of sex |
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325 | (4) |
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7.3 Mate selection, attractiveness, and love |
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329 | (6) |
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329 | (2) |
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331 | (3) |
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7.3.3 Pair-bonding, and love |
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334 | (1) |
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7.4 Parental attachment, care, and parent-offspring conflict |
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335 | (1) |
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7.5 Sperm competition and its consequences for sexual behaviour |
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336 | (7) |
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7.6 Concealed ovulation and its consequences for sexual behaviour |
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343 | (1) |
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7.7 Sexual selection of sexual and non-sexual behaviour |
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344 | (4) |
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7.7.1 Sexual selection and natural selection |
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344 | (3) |
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7.7.2 Non-sexual characteristics may be sexually selected for courtship |
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347 | (1) |
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7.8 Individual differences in sexual rewards |
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348 | (7) |
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349 | (2) |
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7.8.2 How might different types of behaviour be produced by natural selection altering the relative reward value of different stimuli in different individuals? |
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351 | (2) |
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7.8.3 How being tuned to different types of reward could help to produce individual differences in sexual behaviour |
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353 | (2) |
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7.9 The neural reward mechanisms that might mediate some aspects of sexual behaviour |
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355 | (7) |
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7.10 Neural basis of sexual behaviour |
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362 | (5) |
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367 | (1) |
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8 Decision-making mechanisms |
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368 | (86) |
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368 | (1) |
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8.2 Decision-making in an attractor network |
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369 | (4) |
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8.2.1 An attractor decision-making network |
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369 | (2) |
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8.2.2 An integrate-and-fire implementation of the attractor network for probabilistic decision-making |
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371 | (2) |
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8.3 Mean-field analysis of the attractor decision-making network |
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373 | (2) |
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8.4 Stability, energy landscapes, and noise |
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375 | (2) |
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8.5 Neurophysiology of vibrotactile decision-making |
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377 | (3) |
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8.6 Probabilistic decision-making by the integrate-and-fire attractor model |
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380 | (10) |
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8.6.1 Integrate-and-fire simulations of decision-making |
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380 | (1) |
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8.6.2 Decision-making on single trials |
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380 | (2) |
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8.6.3 The probabilistic nature of the decision-making |
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382 | (1) |
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8.6.4 Probabilistic decision-making and Weber's law |
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383 | (3) |
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386 | (1) |
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8.6.6 Finite-size noise effects |
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387 | (3) |
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8.7 Confidence in decisions |
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390 | (27) |
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8.7.1 The model of decision-making |
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391 | (2) |
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8.7.2 Neuronal responses on difficult vs easy trials, and decision confidence |
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393 | (4) |
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8.7.3 Decision times of the neuronal responses |
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397 | (1) |
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397 | (1) |
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8.7.5 Simulation of fMRI signals: haemodynamic convolution of synaptic activity |
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397 | (2) |
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8.7.6 Prediction of the BOLD signals on difficult vs easy decision-making trials |
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399 | (3) |
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8.7.7 Neuroimaging investigations of task difficulty, and confidence |
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402 | (3) |
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8.7.8 Correct decisions vs errors, and confidence |
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405 | (12) |
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8.8 Decisions based on confidence in one's decisions: self-monitoring |
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417 | (15) |
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8.8.1 Decisions about confidence estimates |
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417 | (1) |
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8.8.2 A theory for decisions about confidence estimates |
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417 | (7) |
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8.8.3 Decisions about confidence estimates: neurophysiological evidence |
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424 | (2) |
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8.8.4 Decisions about decisions: self-monitoring |
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426 | (1) |
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8.8.5 Synthesis: decision confidence, noise, neuronal activity, the BOLD signal, and self-monitoring |
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427 | (5) |
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432 | (1) |
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8.10 Comparison with other models of decision-making |
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433 | (2) |
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8.11 Applications and implications of this approach to decision-making |
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435 | (19) |
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8.11.1 Multiple decision-making systems in the brain |
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435 | (2) |
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8.11.2 Distributed decision-making |
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437 | (1) |
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8.11.3 Predicting a decision before the evidence is provided |
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437 | (2) |
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439 | (1) |
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439 | (1) |
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8.11.6 The evolutionary utility of probabilistic choice |
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440 | (1) |
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8.11.7 Unpredictable behaviour |
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441 | (1) |
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441 | (1) |
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442 | (1) |
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8.11.10 Decision-making with sequential inputs and with postponed responses |
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442 | (1) |
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8.11.11 Decision-making between the emotional and rational systems |
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442 | (1) |
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8.11.12 Dynamical neuropsychiatry: schizophrenia |
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442 | (7) |
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8.11.13 Dynamical neuropsychiatry: obsessive-compulsive disorder |
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449 | (3) |
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8.11.14 Decision-making, oscillations, and communication through coherence |
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452 | (2) |
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9 Neuroeconomics and decision-making |
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454 | (29) |
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454 | (1) |
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455 | (1) |
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9.3 Neoclassical economics |
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456 | (2) |
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9.3.1 Utility functions, WARP, and GARP |
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456 | (1) |
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9.3.2 Expected Utility Theory |
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457 | (1) |
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9.3.3 Random Utility Models |
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457 | (1) |
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9.4 Behavioural economics |
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458 | (5) |
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458 | (1) |
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9.4.2 Risk seeking over losses |
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458 | (1) |
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459 | (4) |
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463 | (20) |
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9.5.1 Overview of neuroeconomics |
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463 | (6) |
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9.5.2 A common scale of value for different goods in the orbitofrontal cortex, but no conversion to a common currency |
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469 | (4) |
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9.5.3 Absolute value and relative value are both represented in the orbitofrontal cortex |
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473 | (4) |
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9.5.4 The representation of expected reward value |
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477 | (1) |
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9.5.5 Delay of reward, emotional choice, and rational choice |
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477 | (2) |
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9.5.6 The representation of negative reward prediction error |
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479 | (1) |
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9.5.7 The representation of positive reward prediction error |
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479 | (1) |
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9.5.8 Reward prediction error, temporal difference error, and choice |
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480 | (1) |
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481 | (2) |
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10 Emotional feelings and consciousness: a theory of consciousness |
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483 | (35) |
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483 | (1) |
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10.2 A Higher-Order Syntactic Thought (HOST) theory of consciousness |
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484 | (12) |
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10.2.1 Multiple routes to action |
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484 | (3) |
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10.2.2 A computational hypothesis of consciousness |
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487 | (2) |
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10.2.3 Adaptive value of processing in the system that is related to consciousness |
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489 | (1) |
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490 | (1) |
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491 | (1) |
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492 | (1) |
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10.2.7 Consciousness and causality |
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493 | (2) |
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10.2.8 Consciousness, a computational system for higher-order syntactic manipulation of symbols, and a commentary or reporting functionality |
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495 | (1) |
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10.3 Selection between conscious vs unconscious decision-making, and free will |
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496 | (8) |
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10.3.1 Dual major routes to action: implicit and explicit |
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496 | (6) |
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10.3.2 The Selfish Gene vs The Selfish Phenotype |
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502 | (2) |
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10.3.3 Decision-making between the implicit and explicit systems |
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504 | (1) |
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504 | (2) |
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506 | (1) |
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10.6 Content and meaning in representations |
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507 | (2) |
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10.7 The causal role of consciousness: a theory of the relation between the mind and the brain |
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509 | (2) |
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10.8 Comparison with other theories of consciousness |
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511 | (7) |
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10.8.1 Higher-order thought theories |
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511 | (2) |
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10.8.2 Oscillations and temporal binding |
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513 | (1) |
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10.8.3 A high neural threshold for information to reach consciousness |
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514 | (1) |
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10.8.4 James-Lange theory and Damasio's somatic marker hypothesis about feelings |
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515 | (1) |
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10.8.5 LeDoux's approach to emotion and consciousness |
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515 | (1) |
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10.8.6 Panksepp's approach to emotion and consciousness |
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515 | (1) |
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10.8.7 Global workspace theories of consciousness |
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516 | (1) |
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10.8.8 Monitoring and consciousness |
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516 | (2) |
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11 Conclusions, and broader issues |
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518 | (26) |
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518 | (7) |
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525 | (10) |
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11.2.1 Selection of mainly autonomic responses, and their classical conditioning |
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525 | (1) |
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11.2.2 Selection of approach or withdrawal, and their classical conditioning; fixed action patterns |
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526 | (1) |
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11.2.3 Selection of fixed stimulus-response habits |
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526 | (1) |
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11.2.4 Selection of arbitrary behaviours to obtain goals, action-outcome learning, and emotional learning |
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526 | (1) |
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11.2.5 The roles of the prefrontal cortex in the selection of action, in decision-making, and in attention |
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527 | (6) |
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11.2.6 Selection of actions by explicit rational thought |
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533 | (2) |
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535 | (4) |
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11.4 Emotion and aesthetics |
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539 | (3) |
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542 | (2) |
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A Neural networks and emotion-related learning |
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544 | (45) |
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A.1 Neurons in the brain, the representation of information, and neuronal learning mechanisms |
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544 | (11) |
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544 | (1) |
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A.1.2 Neurons in the brain, and their representation in neuronal networks |
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|
544 | (1) |
|
A.1.3 A formalism for approaching the operation of single neurons in a network |
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|
545 | (2) |
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A.1.4 Synaptic modification |
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|
547 | (2) |
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A.1.5 Long-Term Potentiation and Long-Term Depression |
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|
549 | (4) |
|
A.1.6 Distributed representations |
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|
553 | (2) |
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A.2 Pattern association memory |
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|
555 | (17) |
|
A.2.1 Architecture and operation |
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|
556 | (2) |
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558 | (3) |
|
A.2.3 The vector interpretation |
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|
561 | (1) |
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|
562 | (3) |
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A.2.5 Prototype extraction, extraction of central tendency, and noise reduction |
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565 | (1) |
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|
565 | (1) |
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A.2.7 Local learning rule |
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|
566 | (5) |
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A.2.8 Implications of different types of coding for storage in pattern associators |
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|
571 | (1) |
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A.3 Autoassociation memory: attractor networks |
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572 | (8) |
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A.3.1 Architecture and operation |
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|
573 | (2) |
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A.3.2 Introduction to the analysis of the operation of autoassociation networks |
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|
575 | (1) |
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575 | (5) |
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A.4 Coupled attractor networks |
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|
580 | (2) |
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A.5 Reinforcement learning |
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|
582 | (7) |
|
A.5.1 Associative reward-penalty algorithm of Barto and Sutton |
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|
583 | (1) |
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A.5.2 Error correction or delta rule learning, and classical conditioning |
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|
584 | (1) |
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A.5.3 Temporal Difference (TD) learning |
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585 | (4) |
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589 | (30) |
|
B.1 Overview of different models of decision-making |
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|
589 | (21) |
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B.1.1 Sequential-sampling models: sequential probability ratio test, drift-diffusion, and race models |
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|
589 | (6) |
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B.1.2 Biologically motivated rate models |
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|
595 | (2) |
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597 | (11) |
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B.1.4 Distinguishing model approaches |
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|
608 | (2) |
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B.2 Synaptic facilitation and sequential decision-making |
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|
610 | (2) |
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B.3 Synaptic facilitation, graded firing rates, and postponed decisions |
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|
612 | (2) |
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B.4 The integrate-and-fire formulation used in the model of decisionmaking |
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|
614 | (2) |
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B.5 The mean-field approach used in the model of decision-making |
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|
616 | (2) |
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B.6 The model parameters used in the simulations of decision-making |
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|
618 | (1) |
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|
619 | (68) |
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|
621 | (58) |
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|
679 | (8) |
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|
687 | |