The Integrative Functions of The Basal Ganglia

Cover
Half Title
Title Page
Copyright Page
Dedication
Table of Contents
Preface
About the Author
Chapter 1 Introduction
1.1 Discovering the Basal Ganglia
1.2 Fortunes of the BG
1.3 Extrapyramidal System
1.4 Basic Motif of Cerebral Organization
1.5 Action Selection
1.6 Reinforcement Learning
1.7 Behavior: The Neglected Problem
1.8 Explaining BG Function
1.9 Summary
References
Chapter 2 Anatomical Organization of the Basal Ganglia
2.1 What's in a Name?
2.2 Topographical Inputs to the BG
2.2.1 Limbic Striatum
2.2.2 Associative Striatum
2.2.3 Sensorimotor Striatum
2.3 Chemical Compartments
2.4 Sources of Corticostriatal Projections
2.5 Convergence and Divergence
2.6 Thalamostriatal Projections
2.7 Dopaminergic Projections to the Striatum
2.8 Direct and Indirect Pathways
2.9 BG Output Nuclei
2.10 Globus Pallidus External Segment (GPe)
2.11 Globus Pallidus Internal Segment (GPi)
2.12 Ventral Pallidum
2.13 Ventral Tegmental Area (VTA)
2.14 Substantia Nigra
2.14.1 Nigral Outputs
2.14.2 Substantia Nigra Pars Lateralis
2.15 Subthalamic Nucleus (STN)
2.16 Summary
References
Chapter 3 Synaptic Transmission and Plasticity in the Basal Ganglia
3.1 Striatal Neurons
3.1.1 Spiny Projection Neurons (SPNs)
3.1.2 Fast-Spiking Interneurons (FSIs)
3.1.3 Low-Threshold Spiking Interneurons (LTSIs)
3.1.4 Cholinergic Interneurons (CINs)
3.2 Pallidal Neurons
3.2.1 GPe Neurons
3.2.2 GPi Neurons
3.2.3 Nigral Neurons
3.3 Glutamate
3.3.1 Corticostriatal Transmission
3.3.2 Thalamostriatal Transmission
3.4 Gamma-Aminobutyric Acid (GABA)
3.4.1 Feedforward Inhibition in the Striatum
3.4.2 GABAergic Transmission in the Pallidum
3.4.3 Lateral Inhibition
3.5 Dopamine
3.5.1 How DA Influences Striatal Outputs
3.5.2 Dopaminergic Modulation of GABAergic Transmission
3.6 Acetylcholine (ACh)
3.7 Endocannabinoid (eCB)
3.8 Adenosine
3.9 Synaptic Plasticity in the BG
3.9.1 Induction of Plasticity
3.10 Striatal LTP
3.11 Striatal LTD
3.12 LTP or LTD?
3.13 Functional Implications of Striatal Synaptic Plasticity
3.14 Summary
References
Chapter 4 Current Ideas on BG Function
4.1 Parallel Loops
4.2 Convergence and Divergence
4.3 Interaction Between Loops
4.4 Disinhibition
4.5 Rate Model
4.6 Action Selection Models
4.6.1 Focused Selection
4.6.2 Central Selection
4.6.3 Problems with Action Selection
4.7 Reinforcement Learning
4.8 The Challenge of Behavioral Analysis
4.9 Summary
References
Chapter 5 Behavior and Control
5.1 Insufficiency Principle
5.2 Solutions to the Calculation Problem
5.3 The Definition of Control
5.4 Computing in a Control System
5.5 Misunderstanding Control
5.6 Hierarchical Control
5.7 Beyond Sensorimotor Transformations
5.8 Reinforcement and Teleology
5.9 Neural Signaling and Control Systems
5.10 Information and Coding
5.11 Summary
References
Chapter 6 The Place of the BG in the Hierarchy
6.1 Posture and Movement
6.1.1 Parkinsonian Rigidity
6.2 Control of Muscle Tension and Length
6.2.1 Muscle Length Control and the Fusimotor System
6.3 Bidirectional Control
6.3.1 Stiffness Control
6.3.2 Bandwidth Limitations
6.4 Reticulospinal Pathway
6.5 Position Controllers for Orientation
6.6 Midbrain Contributions to Orienting
6.7 Nigrocollicular Pathway and Eye Movements
6.8 BG Regulation of Steering and Orienting
6.9 SNr and Postural Control
6.10 BG and Locomotion
6.11 SNr and Position Coordinates
6.11.1 SNr and Representation of Position Vectors
6.12 VTA Output and Head Position
6.13 Functional Significance of BG Outputs
6.13.1 Two-Way Comparison Functions
6.14 Summary
References
Chapter 7 Transition Control
7.1 Velocity Control
7.2 DA and Kinematics
7.3 Position Control Versus Velocity Control
7.4 Vector Integration to Endpoint (VITE) Model
7.5 A Neural Integrator in the BG
7.5.1 Integrator Dynamics
7.6 Direct and Indirect Pathways
7.7 Neurobiological Implementation of Integration
7.8 Dopamine and Gain Control
7.8.1 Bradykinesia and Akinesia
7.8.2 Adaptive Gain
7.9 Adaptive Gain and Reinforcement of Action Parameters
7.10 Corticostriatal Circuit and Transition Control
7.11 Compared with Other Models of BG Function
7.12 Limitations in Previous Experimental Designs
7.13 Summary
References
Chapter 8 Higher-Order Transitions and Cognition
8.1 Event Repetition and Control of Tempo
8.2 Regulation of Rhythmic Behavior
8.2.1 Licking
8.3 Related Rates and Gear Coupling
8.4 Interval Timing
8.4.1 BG and Timing
8.4.2 DA Modulation of Timing
8.4.3 Direct and Indirect Pathways
8.5 Serial Order
8.6 Dorsolateral Striatum and Grooming
8.7 Sequence Learning
8.8 Imagination
8.8.1 BG and the Imagination Mode
8.8.2 Mental Rotation
8.9 Working Memory
8.10 Summary
References
Chapter 9 Motivation
9.1 Aspects of Motivation
9.2 Limbic BG and Reward
9.3 Valence and Bidirectional Control
9.4 Distinct Accumbens Outputs Regulate Reparatory and Consummatory Behaviors
9.5 Reinforcement
9.6 Self-Stimulation
9.7 Effort Exertion
9.7.1 Limbic BG and Effort Regulation
9.7.2 DA Contribution to Effort
9.7.3 Conflating Reward Rate and Performance
9.8 DA and Force Generation
9.9 A Labile Motivational Hierarchy
9.10 Parallel BG Networks and Motivated Behaviors
9.11 Summary
References
Chapter 10 Actions and Goals
10.1 Approaching a Goal
10.2 Compulsory Approach
10.3 A Striatal Circuit for Relationship Control and Continuous Pursuit
10.4 Approach Behavior and Feedback
10.5 Learning to Approach
10.6 Contingency, Associative Structures, and Analysis of Conditioning Experiments
10.7 Goal-Directed Actions
10.8 Neural Basis of Action–Outcome Learning
10.8.1 Striatal Activity Modulated by Reward Expectancy
10.8.2 Associative Cortico-BG Network and A–O Learning
10.8.3 Posterior Dorsomedial Striatum
10.8.4 Direct and Indirect Pathways
10.8.5 Synaptic Plasticity in pDMS
10.8.6 The Role of DA in Goal-Directed Actions
10.9 BG Contributions to Neuroprosthetic Control
10.10 Summary
References
Chapter 11 Corticostriatal Contributions to Habits and Behavioral Automaticity
11.1 Multiple Memory Systems
11.2 Place and Response
11.3 BG and Procedural Learning
11.4 Limitations of Multiple Memory Systems Framework
11.5 From Actions to Habits
11.6 Sensorimotor Striatum and Habit Formation
11.6.1 Plasticity Mechanisms Underlying Habits
11.7 Development of Automaticity
11.8 Habits and Skills
11.8.1 Reduced Attentional Demand
11.8.2 Effector Specificity
11.9 Nature of Feedback
11.10 Compulsive Behavior
11.11 Summary
References
Chapter 12 Dopamine and Reinforcement Learning
12.1 Prediction Errors
12.1.1 Rescorla–Wagner Model
12.1.2 Temporal Difference Algorithm
12.2 Principles of RL
12.3 Phasic DA and RPE
12.4 Results that Challenge the RPE Hypothesis
12.5 Value, Performance, and RPE
12.6 Phasic DA and Performance
12.7 Is DA Necessary and Sufficient for Learning?
12.8 Learning and Performance
12.9 Adaptive Gain and Reinforcement
12.10 Reinforcement Learning Versus Control
12.11 Summary
References
Chapter 13 Reorganization, Exploration, and Plasticity
13.1 Learning From a Control Perspective
13.2 Exploration and Reorganization
13.3 BG and Exploration
13.4 Bird Song Learning
13.5 BG Contributions to Bird Song
13.6 DA and Bird Song
13.7 Lessons From Bird Song
13.8 Neural Plasticity and Reorganization
13.8.1 Adaptive Gain and Induction of Long-Term Plasticity
13.8.2 Changes in DA Dependence with Learning
13.9 Summary
References
Chapter 14 Interpretation of Clinical Symptoms
14.1 Analysis of Symptoms
14.1.1 Impaired Input Function
14.1.2 Impaired Output Function
14.1.3 Change in Gain
14.1.4 Oscillations
14.2 Postural Control Deficits
14.3 Bradykinesia, Akinesia, and Paradoxical Kinesia
14.4 Deficits in Locomotion
14.5 Deep Brain Stimulation (DBS)
14.6 Hyperkinetic Symptoms
14.6.1 Chorea
14.6.2 L-DOPA–Induced Dyskinesia (LID)
14.6.3 Reduced Damping
14.6.4 Loss of Selectivity
14.6.5 Loss of Feedback
14.7 Perseveration, Stereotypy, and Compulsion
14.8 Attentional Deficits
14.9 Psychosis and Schizophrenia
14.9.1 Positive and Negative Symptoms
14.9.2 Hallucinations
14.9.3 Delusions
14.10 Summary
References
Chapter 15 Synthesis
15.1 Behavior and Control
15.2 Kinematics as a Gateway to Understanding BG Function
15.3 Cortex Versus BG
15.4 BG Outputs and the Coordination Problem
15.5 Direct and Indirect Pathways
15.6 DA and Adaptive Gain
15.7 Motivational Hierarchy
15.8 Goal Seeking and Control of Relationship
15.9 Higher Functions
15.10 Learning and Reorganization
15.11 Gaps in Understanding
15.12 A New Vista and the Way Forward
References
Index