The Cerebellum

Acknowledgments ix Introduction xi Section I: The Neuronal Machine 1 1 Structure and Physiology 3 Anatomy of the Cerebellar Cortex 3 Physiology of the Cerebellar Cortex 6 Subdivisions of the Cerebellum 10 The Gatekeepers: Vestibular and Deep Cerebellar Nuclei 13 Afferent Connections of the Cerebellum 15 Efferent Connections of the Cerebellum 19 References 21 Further Reading 25 2 Operating the Machine 27 Learning in the Cerebellar Cortex 28 Pattern Recognition by the Cerebellum 30 Neural Networks 32 The Cerebellum as Part of a "Control System" 32 Multiple Sites for Cerebellar Learning? 34 The Cerebellar Clock 36 Conclusions 38 References 38 Further Reading 40 3 Plasticity in the Cerebellar Cortex 41 Cerebellar Long-Term Depression 41 The Calcium Trigger 42 The Synaptic Conversation 44 The Memory Trace 46 What About Potentiation? 48 Other Sites of Plasticity 51 Interneurons 52 Conclusions 55 References 55 Further Reading 59 4 Adjusting the Memory Trace 61 Consolidation Mechanisms 61 Memory Transfer and Synaptic Plasticity 63 Mossy-Fiber Collaterals in the DCN 64 Intrinsic Plasticity in the DCN 66 Other Changes to the Memory Trace 67 References 68 Section II: Motor Learning 71 5 Learning a New Motor Response 73 Learning in the Cerebellar Cortex 75 Cerebellar LTD and Learning 76 The Engram for the NMR 79 Conclusions 82 References 82 Further Reading 85 6 Recalibration for Fine Motor Control 87 A Stable Platform for Vision 87 Adjusting an Orienting Movement 95 Adjusting a Tracking Movement 97 Conclusions 99 References 99 Further Reading 102 7 Perfecting Limb Movements by Motor Learning 103 Updating Dynamic Models 104 Throwing and Pointing 107 Sequence Learning 109 Stepping and Changes to gait 111 The Agile Mouse 113 Conclusions 114 References 114 Further Reading 117 Section III: Precision Control 119 8 Coordination 121 Precise Ocular Coordination 122 Coordinating the Eyes and the Head 126 Error Correction for Limb Movements 127 Planning for Multiple Joints 129 Internal Models Revisited 132 Conclusions 134 References 134 Further Reading 137 9 Balance and Locomotion 139 Cerebellar Ataxia 140 Signals from the Inner Ear 142 Coordinating Locomotion 146 Navigation 148 Conclusions 150 References 150 Further Reading 152 10 Timing 153 Timing Using Discharge Rates 154 Timing Using Synchronous Firing 161 Conclusions 165 References 166 Further Reading 168 Section IV: Interpreting the World 169 11 Intelligence and Language 173 General Intelligence 175 Executive Function 177 Problem-solving 181 Speech and Language 183 Possible Mechanisms 187 Conclusions 188 References 188 Further Reading 191 12 Sensing, Feeling, and Interacting 193 Sensory Perception 194 Attending to theWorld 198 Prediction 200 Mental Imagery 202 Social Skills 204 Conclusions 208 References 208 Further Reading 211 Summary: What does the cerebellum do? 213 Index 215