Series Foreword |
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xiii | |
Acknowledgments |
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xv | |
Preface |
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xvii | |
1 Preliminary Material |
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1 | (58) |
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1 | (3) |
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1.1.1 The Cell, the Circuit, and the Brain |
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1 | (1) |
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1.1.2 Physics of Electrical Circuits |
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1 | (1) |
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1.1.3 Mathematical Preliminaries |
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2 | (2) |
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1.1.4 Writing Computer Code |
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4 | (1) |
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1.2 The Neuron, the Circuit, and the Brain |
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4 | (7) |
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4 | (3) |
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7 | (1) |
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8 | (3) |
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1.3 Physics of Electrical Circuits |
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11 | (3) |
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1.3.1 Terms and Properties |
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11 | (1) |
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1.3.2 Pumps, Reservoirs, and Pipes |
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12 | (1) |
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1.3.3 Some Peculiarities of the Electrical Properties of Neurons |
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13 | (1) |
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1.4 Mathematical Background |
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14 | (22) |
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1.4.1 Ordinary Differential Equations |
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15 | (9) |
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1.4.2 Vectors, Matrices, and Their Basic Operations |
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24 | (4) |
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1.4.3 Probability and Bayes' Theorem |
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28 | (8) |
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1.5 Introduction to Computing and MATLAB |
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36 | (15) |
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37 | (1) |
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38 | (2) |
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1.5.3 Allocation of Memory |
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40 | (1) |
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1.5.4 Using the Colon (:) Symbol |
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41 | (1) |
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42 | (1) |
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42 | (1) |
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1.5.7 Vector and Matrix Operations in MATLAB |
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43 | (1) |
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44 | (2) |
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46 | (1) |
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47 | (1) |
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1.5.11 Some Operations Useful for Modeling Neurons |
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48 | (1) |
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1.5.12 Good Coding Practice |
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49 | (2) |
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1.6 Solving Ordinary Differential Equations (ODEs) |
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51 | (8) |
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1.6.1 Forward Euler Method |
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51 | (1) |
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1.6.2 Simulating ODEs with MATLAB |
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52 | (2) |
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1.6.3 Solving Coupled ODEs with Multiple Variables |
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54 | (1) |
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1.6.4 Solving ODEs with Nested for Loops |
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55 | (1) |
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1.6.5 Comparing Simulation Methods |
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55 | (1) |
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1.6.6 Euler-Mayamara Method: Forward Euler with White Noise |
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56 | (3) |
2 The Neuron and Minimal Spiking Models |
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59 | (30) |
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2.1 The Nernst Equilibrium Potential |
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59 | (3) |
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2.2 An Equivalent Circuit Model of the Neural Membrane |
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62 | (4) |
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2.2.1 Depolarization versus Hyperpolarization |
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65 | (1) |
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2.3 The Leaky Integrate-and-Fire Model |
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66 | (4) |
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2.3.1 Specific versus Absolute Properties of the Cell |
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68 | (1) |
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2.3.2 Firing Rate as a Function of Current (f-I Curve) of the Leaky Integrate-and-Fire Model |
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69 | (1) |
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2.4 Tutorial 2.1: The f-I Curve of the Leaky Integrate-and-Fire Neuron |
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70 | (2) |
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2.5 Extensions of the Leaky Integrate-and-Fire Model |
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72 | (4) |
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72 | (2) |
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2.5.2 Spike-Rate Adaptation (SRA) |
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74 | (2) |
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2.6 Tutorial 2.2: Modeling the Refractory Period |
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76 | (2) |
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2.7 Further Extensions of the Leaky Integrate-and-Fire Model |
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78 | (3) |
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2.7.1 Exponential Leaky Integrate-and-Fire (ELIF) Model |
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78 | (1) |
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2.7.2 Two-Variable Models: The Adaptive Exponential Leaky Integrate-and-Fire (AELIF) Neuron |
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79 | (2) |
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2.7.3 Limitations of the LIF Formalism |
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81 | (1) |
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2.8 Tutorial 2.3: Models Based on Extensions of the LIF Neuron |
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81 | (5) |
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2.9 Appendix: Calculation of the Nernst Potential |
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86 | (3) |
3 Analysis of Individual Spike Trains |
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89 | (44) |
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3.1 Responses of Single Neurons |
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89 | (11) |
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89 | (3) |
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3.1.2 Time-Varying Responses and the Peristimulus lime Histogram (PSTH) |
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92 | (1) |
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3.1.3 Neurons as Linear Filters and the Linear-Nonlinear Model |
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93 | (3) |
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3.1.4 Spike-Triggered Average |
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96 | (1) |
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3.1.5 White-Noise Stimuli for Receptive Field Generation |
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96 | (2) |
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3.1.6 Spatiotemporal Receptive Fields |
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98 | (2) |
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3.2 Tutorial 3.1: Generating Receptive Fields with Spike-Triggered Averages |
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100 | (4) |
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3.3 Spike-Train Statistics |
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104 | (6) |
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3.3.1 Coefficient of Variation (CV) of Interspike Intervals |
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105 | (2) |
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107 | (1) |
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3.3.3 The Homogeneous Poisson Process: A Random Point Process for Artificial Spike Trains |
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108 | (1) |
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3.3.4 Comments on Analyses and Use of Dummy Data |
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109 | (1) |
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3.4 Tutorial 3.2: Statistical Properties of Simulated Spike Trains |
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110 | (3) |
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3.5 Receiver-Operating Characteristic (ROC) |
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113 | (8) |
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3.5.1 Producing the ROC Curve |
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113 | (2) |
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3.5.2 Optimal Position of the Threshold |
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115 | (3) |
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3.5.3 Uncovering the Underlying Distributions from Binary Responses: Recollection versus Familiarity |
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118 | (3) |
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3.6 Tutorial 3.3: Receiver-Operating Characteristic of a Noisy Neuron |
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121 | (2) |
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3.7 Appendix A: The Poisson Process |
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123 | (5) |
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3.7.1 The Poisson Distribution |
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123 | (2) |
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3.7.2 Expected Value of the Mean of a Poisson Process |
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125 | (1) |
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3.7.3 Fano Factor of the Poisson Process |
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125 | (1) |
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3.7.4 The Coefficient of Variation (CV) of the ISI Distribution of a Poisson Process |
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126 | (1) |
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3.7.5 Selecting from a Probability Distribution: Generating ISIS for the Poisson Process |
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127 | (1) |
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3.8 Appendix B: Stimulus Discriminability |
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128 | (5) |
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3.8.1 Optimal Value of Threshold |
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129 | (1) |
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3.8.2 Calculating the Probability of an Error |
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130 | (1) |
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3.8.3 Generating a Z-Score from a Probability |
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130 | (3) |
4 Conductance-Based Models |
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133 | (40) |
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4.1 Introduction to the Hodgkin-Huxley Model |
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133 | (4) |
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4.1.1 Positive versus Negative Feedback |
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134 | (2) |
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4.1.2 Voltage Clamp versus Current Clamp |
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136 | (1) |
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4.2 Simulation of the Hodgkin-Huxley Model |
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137 | (10) |
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138 | (1) |
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4.2.2 Full Set of Dynamical Equations for the Hodgkin-Huxley Model |
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139 | (1) |
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4.2.3 Dynamical Behavior of the Hodgkin-Huxley Model: A Type-II Neuron |
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140 | (7) |
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4.3 Tutorial 4.1: The Hodgkin-Huxley Model as an Oscillator |
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147 | (3) |
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4.4 The Connor-Stevens Model: A Type-I Model |
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150 | (4) |
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4.5 Calcium Currents and Bursting |
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154 | (2) |
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4.5.1 Thalamic Rebound and the T-Type Calcium Channel |
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155 | (1) |
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4.6 Tutorial 4.2: Postinhibitory Rebound |
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156 | (3) |
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4.7 Modeling Multiple Compartments |
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159 | (7) |
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4.7.1 The Pinsky-Rinzel Model of an Intrinsic Burster |
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160 | (1) |
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4.7.2 Simulating the Pinsky-Rinzel Model |
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160 | (3) |
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4.7.3 A Note on Multicompartmental Modeling with Specific Conductances versus Absolute Conductances |
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163 | (3) |
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166 | (1) |
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4.8 Hyperpolarization-Activated Currents (Ih) and Pacemaker Control |
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166 | (2) |
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4.9 Dendritic Computation |
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168 | (2) |
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4.10 Tutorial 4.3: A Two-Compartment Model of an Intrinsically Bursting Neuron |
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170 | (3) |
5 Connections between Neurons |
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173 | (38) |
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173 | (6) |
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5.1.1 Electrical Synapses |
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173 | (1) |
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174 | (5) |
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5.2 Modeling Synaptic Transmission through Chemical Synapses |
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179 | (3) |
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5.2.1 Spike-Induced Transmission |
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179 | (2) |
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181 | (1) |
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182 | (3) |
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5.3.1 Short-Term Synaptic Depression |
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183 | (1) |
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5.3.2 Short-Term Synaptic Facilitation |
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183 | (1) |
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5.3.3 Modeling Dynamical Synapses |
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184 | (1) |
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5.4 Tutorial 5.1: Synaptic Responses to Changes in Inputs |
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185 | (2) |
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5.5 The Connectivity Matrix |
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187 | (6) |
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5.5.1 General Types of Connectivity Matrices |
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189 | (1) |
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5.5.2 Cortical Connections: Sparseness and Structure |
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190 | (1) |
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191 | (2) |
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5.6 Tutorial 5.2: Detecting Circuit Structure and Nonrandom Features within a Connectivity Matrix |
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193 | (3) |
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5.7 Oscillations and Multistability in Small Circuits |
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196 | (1) |
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5.8 Central Pattern Generators |
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197 | (6) |
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5.8.1 The Half-Center Oscillator |
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199 | (1) |
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5.8.2 The Triphasic Rhythm |
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199 | (1) |
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5.8.3 Phase Response Curves |
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200 | (3) |
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5.9 Tutorial 5.3: Bistability and Oscillations from Two LIF Neurons |
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203 | (2) |
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5.10 Appendix: Synaptic Input Produced by a Poisson Process |
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205 | (6) |
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5.10.1 Synaptic Saturation |
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205 | (3) |
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5.10.2 Synaptic Depression |
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208 | (1) |
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5.10.3 Synaptic Facilitation |
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209 | (1) |
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5.10.4 Notes on Combining Mechanisms |
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209 | (2) |
6 Firing-Rate Models and Network Dynamics |
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211 | (46) |
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211 | (2) |
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6.2 Simulating a Firing-Rate Model |
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213 | (4) |
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6.2.1 Meaning of a Unit and Dale's Principle |
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216 | (1) |
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6.3 Recurrent Feedback and Bistability |
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217 | (10) |
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6.3.1 Bistability from Positive Feedback |
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217 | (4) |
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6.3.2 Limiting the Maximum Firing Rate Reached |
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221 | (1) |
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6.3.3 Dynamics of Synaptic Response |
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222 | (1) |
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6.3.4 Dynamics of Synaptic Depression and Facilitation |
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223 | (2) |
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6.3.5 Integration and Parametric Memory |
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225 | (2) |
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6.4 Tutorial 6.1: Bistability and Oscillations in a Firing-Rate Model with Feedback |
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227 | (2) |
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6.5 Decision-Making Circuits |
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229 | (7) |
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6.5.1 Decisions by Integration of Evidence |
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232 | (1) |
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6.5.2 Decision-Making Performance |
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233 | (2) |
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6.5.3 Decisions as State Transitions |
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235 | (1) |
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235 | (1) |
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6.6 Tutorial 6.2: Dynamics of a Decision-Making Circuit in Two Modes of Operation |
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236 | (2) |
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6.7 Oscillations from Excitatory and Inhibitory Feedback |
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238 | (4) |
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6.8 Tutorial 6.3: Frequency of an Excitatory-Inhibitory Coupled Unit Oscillator and PING |
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242 | (3) |
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6.9 Orientation Selectivity and Contrast Invariance |
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245 | (5) |
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246 | (4) |
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6.10 Ring Attractors for Spatial Memory and Head Direction |
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250 | (4) |
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6.10.1 Dynamics of the Ring Attractor |
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252 | (2) |
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6.11 Tutorial 6.4: Orientation Selectivity in a Ring Model |
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254 | (3) |
7 An Introduction to Dynamical Systems |
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257 | (36) |
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7.1 What Is a Dynamical System? |
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257 | (1) |
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7.2 Single Variable Behavior and Fixed Points |
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258 | (3) |
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258 | (2) |
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7.2.2 Requirement for Oscillations |
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260 | (1) |
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7.3 Models with Two Variables |
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261 | (6) |
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7.3.1 Nullclines and Phase-Plane Analysis |
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262 | (2) |
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7.3.2 The Inhibition-Stabilized Network |
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264 | (3) |
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7.3.3 How Inhibitory Feedback to Inhibitory Neurons Impacts Stability of States |
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267 | (1) |
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7.4 Tutorial 7.1: The Inhibition-Stabilized Circuit |
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267 | (2) |
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7.5 Attractor State Itinerancy |
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269 | (2) |
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269 | (1) |
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7.5.2 Noise-Driven Transitions in a Bistable System |
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270 | (1) |
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7.6 Quasistability and Relaxation Oscillators: The FitzHugh-Nagumo Model |
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271 | (4) |
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7.7 Heteroclinic Sequences |
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275 | (1) |
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275 | (7) |
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7.8.1 Chaotic Systems and Lack of Predictability |
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277 | (2) |
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7.8.2 Examples of Chaotic Neural Circuits |
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279 | (3) |
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282 | (6) |
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7.9.1 Power-Law Distributions |
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283 | (1) |
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7.9.2 Requirements for Criticality |
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284 | (3) |
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7.9.3 A Simplified Avalanche Model with a Subset of the Features of Criticality |
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287 | (1) |
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7.10 Tutorial 7.2: Diverse Dynamical Systems from Similar Circuit Architectures |
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288 | (2) |
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7.11 Appendix: Proof of the Scaling Relationship for Avalanche Sizes |
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290 | (3) |
8 Learning and Synaptic Plasticity |
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293 | (46) |
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293 | (4) |
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8.1.1 Modeling Hebbian Plasticity |
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296 | (1) |
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8.2 Tutorial 8.1: Pattern Completion and Pattern Separation via Hebbian Learning |
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297 | (3) |
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8.3 Spike-Timing Dependent Plasticity (STDP) |
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300 | (9) |
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302 | (2) |
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8.3.2 Synaptic Competition via STDP |
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304 | (1) |
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8.3.3 Sequence Learning via STDP |
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305 | (1) |
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305 | (3) |
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8.3.5 A Note on Spike-Timing Dependent Plasticity |
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308 | (1) |
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8.3.6 Mechanisms of Spike-Timing Dependent Synaptic Plasticity |
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309 | (1) |
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8.4 More Detailed Empirical Models of Synaptic Plasticity |
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309 | (2) |
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8.5 Tutorial 8.2: Competition via STDP |
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311 | (2) |
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313 | (6) |
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8.6.1 Firing-Rate Homeostasis |
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314 | (2) |
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8.6.2 Homeostasis of Synaptic Inputs |
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316 | (1) |
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8.6.3 Homeostasis of Intrinsic Properties |
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317 | (2) |
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319 | (7) |
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321 | (1) |
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8.7.2 Reward Prediction Errors and Reinforcement Learning |
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322 | (2) |
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8.7.3 The Weather-Prediction Task |
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324 | (1) |
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8.7.4 Calculations Required in the Weather-Prediction Task |
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325 | (1) |
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8.8 Tutorial 8.3: Learning the Weather-Prediction Task in a Neural Circuit |
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326 | (3) |
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8.9 Eyeblink Conditioning |
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329 | (2) |
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8.10 Tutorial 8.4: A Model of Eyeblink Conditioning |
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331 | (4) |
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8.11 Appendix A: Rate-Dependent Plasticity via STDP between Uncorrelated Poisson Spike Trains |
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335 | (1) |
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8.12 Appendix B: Rate-Dependence of Triplet STDP between Uncorrelated Poisson Spike Trains |
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336 | (3) |
9 Analysis of Population Data |
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339 | (30) |
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9.1 Principal Component Analysis (PCA) |
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340 | (6) |
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9.1.1 PCA for Sorting of Spikes |
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341 | (1) |
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9.1.2 PCA for Analysis of Firing Rates |
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342 | (1) |
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342 | (3) |
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9.1.4 The Procedure of PCA |
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345 | (1) |
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9.2 Tutorial 9.1: Principal Component Analysis of Firing-Rate Trajectories |
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346 | (2) |
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9.3 Single-Trial versus Trial-Averaged Analyses |
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348 | (1) |
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9.4 Change-Point Detection |
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349 | (2) |
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351 | (1) |
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9.5 Hidden Markov Modeling (HMM) |
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351 | (4) |
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9.6 Tutorial 9.2: Change-Point Detection for a Poisson Process |
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355 | (2) |
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9.7 Decoding Position from Multiple Place Fields |
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357 | (5) |
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9.8 Appendix A: How PCA Works: Choosing a Direction to Maximize the Variance of the Projected Data |
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362 | (4) |
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9.8.1 Carrying out PCA without a Built-in Function |
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364 | (2) |
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9.9 Appendix B: Determining the Probability of Change Points for a Poisson Process |
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366 | (3) |
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366 | (1) |
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9.9.2 Evaluating the Change Point, Method 1 |
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367 | (1) |
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9.9.3 Evaluating the Change Point, Method 2 |
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367 | (2) |
References |
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369 | (12) |
Index |
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381 | |