
Dynamical Systems in Neuroscience
The Geometry of Excitability and Bursting
Paperback | 22 January 2010
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orIn order to model neuronal behavior or to interpret the results of modeling studies, neuroscientists must call upon methods of nonlinear dynamics. This book offers an introduction to nonlinear dynamical systems theory for researchers and graduate students in neuroscience. It also provides an overview of neuroscience for mathematicians who want to learn the basic facts of electrophysiology.
Dynamical Systems in Neuroscience presents a systematic study of the relationship of electrophysiology, nonlinear dynamics, and computational properties of neurons. It emphasizes that information processing in the brain depends not only on the electrophysiological properties of neurons but also on their dynamical properties. The book introduces dynamical systems, starting with one- and two-dimensional Hodgkin-Huxley-type models and continuing to a description of bursting systems. Each chapter proceeds from the simple to the complex, and provides sample problems at the end. The book explains all necessary mathematical concepts using geometrical intuition; it includes many figures and few equations, making it especially suitable for non-mathematicians. Each concept is presented in terms of both neuroscience and mathematics, providing a link between the two disciplines.
Nonlinear dynamical systems theory is at the core of computational neuroscience research, but it is not a standard part of the graduate neuroscience curriculum-or taught by math or physics department in a way that is suitable for students of biology. This book offers neuroscience students and researchers a comprehensive account of concepts and methods increasingly used in computational neuroscience.
An additional chapter on synchronization, with more advanced material, can be found at the author's website, www.izhikevich.com.
Explains the relationship of electrophysiology, nonlinear dynamics, and the computational properties of neurons, with each concept presented in terms of both neuroscience and mathematics and illustrated using geometrical intuition.
In order to model neuronal behavior or to interpret the results of modeling studies, neuroscientists must call upon methods of nonlinear dynamics. This book offers an introduction to nonlinear dynamical systems theory for researchers and graduate students in neuroscience. It also provides an overview of neuroscience for mathematicians who want to learn the basic facts of electrophysiology.
Dynamical Systems in Neuroscience presents a systematic study of the relationship of electrophysiology, nonlinear dynamics, and computational properties of neurons. It emphasizes that information processing in the brain depends not only on the electrophysiological properties of neurons but also on their dynamical properties. The book introduces dynamical systems, starting with one- and two-dimensional Hodgkin-Huxley-type models and continuing to a description of bursting systems. Each chapter proceeds from the simple to the complex, and provides sample problems at the end. The book explains all necessary mathematical concepts using geometrical intuition; it includes many figures and few equations, making it especially suitable for non-mathematicians. Each concept is presented in terms of both neuroscience and mathematics, providing a link between the two disciplines.
Nonlinear dynamical systems theory is at the core of computational neuroscience research, but it is not a standard part of the graduate neuroscience curriculum-or taught by math or physics department in a way that is suitable for students of biology. This book offers neuroscience students and researchers a comprehensive account of concepts and methods increasingly used in computational neuroscience.
An additional chapter on synchronization, with more advanced material, can be found at the author's website, www.izhikevich.com.
Industry Reviews
A unique contribution to the theoretical neuroscience literature that can serve as a useful reference for audiences ranging from quantitatively skilled undergraduates interested in mathematical modeling, to neuroscientists at all levels, to graduate students and even researchers in the field of theoretical neuroscience.
-- Jonathan E. Rubin * Mathematical Review *| Preface | p. xv |
| Introduction | p. 1 |
| Neurons | p. 1 |
| What Is a Spike? | p. 2 |
| Where Is the Threshold? | p. 3 |
| Why Are Neurons Different, and Why Do We Care? | p. 6 |
| Building Models | p. 6 |
| Dynamical Systems | p. 8 |
| Phase Portraits | p. 8 |
| Bifurcations | p. 11 |
| Hodgkin Classification | p. 14 |
| Neurocomputational properties | p. 16 |
| Building Models (Revisited) | p. 20 |
| Review of Important Concepts | p. 21 |
| Bibliographical Notes | p. 21 |
| Electrophysiology of Neurons | |
| Ions | p. 25 |
| Nernst Potential | p. 26 |
| Ionic Currents and Conductances | p. 27 |
| Equivalent Circuit | p. 28 |
| Resting Potential and Input Resistance | p. 29 |
| Voltage-Clamp and I-V Relation | p. 30 |
| Conductances | p. 32 |
| Voltage-Gated Channels | p. 33 |
| Activation of Persistent Currents | p. 34 |
| Inactivation of Transient Currents | p. 35 |
| Hyperpolarization-Activated Channels | p. 36 |
| The Hodgkin-Huxley Model | p. 37 |
| Hodgkin-Huxley Equations | p. 37 |
| Action Potential | p. 41 |
| Propagation of the Action Potentials | p. 42 |
| Dendritic Compartments | p. 43 |
| Summary of Voltage-Gated Currents | p. 44 |
| Review of Important Concepts | p. 49 |
| Bibliographical Notes | p. 50 |
| Exercises | p. 50 |
| One-Dimensional Systems | |
| Electrophysiological Examples | p. 53 |
| I-V Relations and Dynamics | p. 54 |
| Leak + Instantaneous INa, p | p. 55 |
| Dynamical Systems | p. 57 |
| Geometrical Analysis | p. 59 |
| Equilibria | p. 60 |
| Stability | p. 60 |
| Eigenvalues | p. 61 |
| Unstable Equilibria | p. 61 |
| Attraction Domain | p. 62 |
| Threshold and Action Potential | p. 63 |
| Threshold and Action Potential | p. 63 |
| Bistability and Hysteresis | p. 66 |
| Phase Portraits | p. 67 |
| Topological Equivalence | p. 68 |
| Local Equivalence and the Hartman-Grobman Theorem | p. 69 |
| Bifurcations | p. 70 |
| Saddle-Node (Fold) Bifurcation | p. 74 |
| Slow Transition | p. 75 |
| Bifurcation Diagram | p. 77 |
| Bifurcations and I-V Relations | p. 77 |
| Quadratic Integrate-and-Fire Neuron | p. 80 |
| Review of Important Concepts | p. 82 |
| Bibliographical Notes | p. 83 |
| Exercises | p. 83 |
| Two-Dimensional Systems | |
| Planar Vector Fields | p. 89 |
| Nullclines | p. 92 |
| Trajectories | p. 94 |
| Limit Cycles | p. 96 |
| Relaxation Oscillators | p. 98 |
| Equilibria | p. 99 |
| Stability | p. 100 |
| Local Linear Analysis | p. 101 |
| Eigenvalues and Eigenvectors | p. 102 |
| Local Equivalence | p. 103 |
| Classification of Equilibria | p. 103 |
| Example: FitzHugh-Nagumo Model | p. 106 |
| Phase Portraits | p. 108 |
| Bistability and Attraction Domains | p. 108 |
| Stable/Unstable Manifolds | p. 109 |
| Homoclinic/Heteroclinic Trajectories | p. 111 |
| Saddle-Node Bifurcation | p. 113 |
| Andronov-Hopf Bifurcation | p. 116 |
| Review of Important Concepts | p. 121 |
| Bibliographical Notes | p. 122 |
| Exercises | p. 122 |
| Conductance-Based Models and Their Reductions | |
| Minimal Models | p. 127 |
| Amplifying and Resonant Gating Variables | p. 129 |
| INa,p+IK -Model | p. 132 |
| INa,t -Model | p. 133 |
| INa, p+Ih -Model | p. 136 |
| Ih+IKir -Model | p. 138 |
| IK+IKir -Model | p. 140 |
| IA -Model | p. 142 |
| Ca2+ -Gated Minimal Models | p. 147 |
| Reduction of Multidimensional Models | p. 147 |
| Hodgkin-Huxley model | p. 147 |
| Equivalent Potentials | p. 151 |
| Nullclines and I-V Relations | p. 151 |
| Reduction to Simple Model | p. 153 |
| Review of Important Concepts | p. 156 |
| Bibliographical Notes | p. 156 |
| Exercises | p. 157 |
| Bifurcations | |
| Equilibrium (Rest State) | p. 159 |
| Saddle-Node (Fold) | p. 162 |
| Saddle-Node on Invariant Circle | p. 164 |
| Supercritical Andronov-Hopf | p. 168 |
| Subcritical Andronov-Hopf | p. 174 |
| Limit Cycle (Spiking State) | p. 178 |
| Saddle-Node on Invariant Circle | p. 180 |
| Supercritical Andronov-Hopf | p. 181 |
| Fold Limit Cycle | p. 181 |
| Homoclinic | p. 185 |
| Other Interesting Cases | p. 190 |
| Three-Dimensional Phase Space | p. 190 |
| Cusp and Pitchfork | p. 192 |
| Bogdanov-Takens | p. 194 |
| Relaxation Oscillators and Canards | p. 198 |
| Bautin | p. 200 |
| Saddle-Node Homoclinic Orbit | p. 201 |
| Hard and Soft Loss of Stability | p. 204 |
| Bibliographical Notes | p. 205 |
| Exercises | p. 210 |
| Neuronal Excitability | |
| Excitability | p. 215 |
| Bifurcations | p. 216 |
| Hodgkin's Classification | p. 218 |
| Classes 1 and 2 | p. 221 |
| Class 3 | p. 222 |
| Ramps, Steps, and Shocks | p. 224 |
| Bistability | p. 226 |
| Class 1 and 2 Spiking | p. 228 |
| Integrators vs. Resonators | p. 229 |
| Fast Subthreshold Oscillations | p. 230 |
| Frequency Preference and Resonance | p. 232 |
| Frequency Preference in Vivo | p. 237 |
| Thresholds and Action Potentials | p. 238 |
| Threshold Manifolds | p. 240 |
| Rheobase | p. 242 |
| Postinhibitory Spike | p. 242 |
| Inhibition-Induced Spiking | p. 244 |
| Spike Latency | p. 246 |
| Flipping from an Integrator to a Resonator | p. 248 |
| Transition Between Integrators and Resonators | p. 251 |
| Slow Modulation | p. 252 |
| Spike Frequency Modulation | p. 255 |
| I-V Relation | p. 256 |
| Slow Subthreshold Oscillation | p. 258 |
| Rebound Response and Voltage Sag | p. 259 |
| AHP and ADP | p. 260 |
| Review of Important Concepts | p. 264 |
| Bibliographical Notes | p. 264 |
| Exercises | p. 265 |
| Simple Models | |
| Simplest Models | p. 267 |
| Integrate-and-Fire | p. 268 |
| Resonate-and-Fire | p. 269 |
| Quadratic Integrate-and-Fire | p. 270 |
| Simple Model of Choice | p. 272 |
| Canonical Models | p. 278 |
| Cortex | p. 281 |
| Regular Spiking (RS) Neurons | p. 282 |
| Intrinsically Bursting (IB) Neurons | p. 288 |
| Multi-Compartment Dendritic Tree | p. 292 |
| Chattering (CH) Neurons | p. 294 |
| Low-Threshold Spiking (LTS) Interneurons | p. 296 |
| Fast Spiking (FS) Interneurons | p. 298 |
| Late Spiking (LS) Interneurons | p. 300 |
| Diversity of Inhibitory Interneurons | p. 301 |
| Thalamus | p. 304 |
| Thalamocortical (TC) Relay Neurons | p. 305 |
| Reticular Thalamic Nucleus (RTN) Neurons | p. 306 |
| Thalamic Interneurons | p. 308 |
| Other Interesting Cases | p. 308 |
| Hippocampal CA1 Pyramidal Neurons | p. 308 |
| Spiny Projection Neurons of Neostriatum and Basal Ganglia | p. 311 |
| Mesencephalic V Neurons of Brainstream | p. 313 |
| Stellate Cells of Entorhinal Cortex | p. 314 |
| Mital Neurons of the Olfactory Bulb | p. 316 |
| Review of Important Concepts | p. 319 |
| Bibliographical Notes | p. 319 |
| Exercises | p. 321 |
| Bursting | |
| Electrophysiology | p. 325 |
| Example: The INa,p+IK+IK(M)-Model | p. 327 |
| Fast-Slow Dynamics | p. 329 |
| Minimal Models | p. 332 |
| Central Pattern Generators and Half-Center Oscillators | p. 334 |
| Geometry | p. 335 |
| Fast-Slow Bursters | p. 336 |
| Phase Portraits | p. 336 |
| Averaging | p. 339 |
| Equivalent Voltage | p. 341 |
| Hysteresis Loops and Slow Waves | p. 342 |
| Bifurcations ôResting Bursting Tonic Spikingö | p. 344 |
| Classification | p. 347 |
| Fold/Homoclinic | p. 350 |
| Circle/Circle | p. 354 |
| Fold/Fold Cycle | p. 364 |
| Fold/Hopf | p. 365 |
| Fold/Circle | p. 366 |
| Neurocomputational Properties | p. 367 |
| How to Distinguish? | p. 367 |
| Integrators vs. Resonators | p. 368 |
| Bistability | p. 368 |
| Bursts as a Unit of Neuronal Information | p. 371 |
| Chirps | p. 372 |
| Synchronization | p. 373 |
| Review of Important Concepts | p. 375 |
| Bibliographical Notes | p. 376 |
| Exercises | p. 378 |
| Synchronization | |
| Solutions to Exercises | p. 387 |
| References | p. 419 |
| Index | p. 435 |
| Synchronization (www.izhikevich.com) | |
| Pulsed Coupling | p. 444 |
| Phase of Oscillation | p. 444 |
| Isochrons | p. 445 |
| PRC | p. 446 |
| Type 0 and Type 1 Phase Response | p. 450 |
| Poincare Phase Map | p. 452 |
| Fixed Points | p. 453 |
| Synchronization | p. 454 |
| Phase-Locking | p. 456 |
| Arnold Tongues | p. 456 |
| Weak Coupling | p. 458 |
| Winfree's Approach | p. 459 |
| Kuramoto's Approach | p. 460 |
| Malkin's Approach | p. 461 |
| Measuring PRCs Experimentally | p. 462 |
| Phase Model for Coupled Oscillators | p. 465 |
| Synchronization | p. 467 |
| Two Oscillators | p. 469 |
| Chains | p. 471 |
| Networks | p. 473 |
| Mean-Field Approximations | p. 474 |
| Examples | p. 475 |
| Phase Oscillators | p. 475 |
| SNIC Oscillators | p. 477 |
| Homoclinic Oscillators | p. 482 |
| Relaxation Oscillators and FTM | p. 484 |
| Bursting Oscillators | p. 486 |
| Review of Important Concepts | p. 488 |
| Bibliographical Notes | p. 489 |
| Solutions | p. 497 |
| Table of Contents provided by Publisher. All Rights Reserved. |
ISBN: 9780262514200
ISBN-10: 0262514206
Series: Computational Neuroscience
Published: 22nd January 2010
Format: Paperback
Language: English
Number of Pages: 458
Audience: General Adult
For Ages: 18+ years old
Publisher: RANDOM HOUSE US
Country of Publication: US
Dimensions (cm): 25.5 x 17.8 x 2.7
Weight (kg): 0.82
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