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