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Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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Chapter 1IntroductionThis chapter highlights some of the most important concepts developed <strong>in</strong> the book.First, we discuss several common misconceptions regard<strong>in</strong>g the spike-generation mechanismof neurons. Our goal is to motivate the reader <strong>in</strong>to th<strong>in</strong>k<strong>in</strong>g of a neuron notonly <strong>in</strong> terms of ions and channels, as many biologists do, and not only <strong>in</strong> terms of<strong>in</strong>put/output relationship, as many theoreticians do, but also as a nonl<strong>in</strong>ear dynamicalsystem that looks at the <strong>in</strong>put through the prism of its own <strong>in</strong>tr<strong>in</strong>sic dynamics. Weask such questions as “what makes a neuron fire?” or “where is the threshold”, andthen outl<strong>in</strong>e the answers us<strong>in</strong>g geometrical theory of dynamical systems.From a dynamical systems po<strong>in</strong>t of view, neurons are excitable because they arenear a transition, called bifurcation, from rest<strong>in</strong>g to susta<strong>in</strong>ed spik<strong>in</strong>g activity. Whilethere is a huge number of possible ionic mechanisms of excitability and spike-generation,there are only four different bifurcation mechanisms that can result <strong>in</strong> such a transition.Consider<strong>in</strong>g the geometry of phase portraits at these bifurcations, we can understandmany computational properties of neurons, such as the nature of threshold and all-ornonespik<strong>in</strong>g, the co-existence of rest<strong>in</strong>g and spik<strong>in</strong>g states, the orig<strong>in</strong> of spike latencies,post-<strong>in</strong>hibitory spikes, the mechanism of <strong>in</strong>tegration and resonance, etc. Moreover, wecan understand how these properties are <strong>in</strong>terrelated, why some are equivalent andsome are mutually exclusive.1.1 NeuronsIf somebody were to put a gun to the head of the author of this book and ask himto name the s<strong>in</strong>gle most important concept <strong>in</strong> bra<strong>in</strong> science, he would say it is theconcept of a neuron. There are only 10 11 or so neurons <strong>in</strong> the human bra<strong>in</strong>, muchfewer than the number of non-neural cells such as glia. Yet neurons are unique <strong>in</strong> thesense that only they can transmit electrical signals over long distances. From neuronallevel we can go down to cell biophysics, to molecular biology of gene regulation, etc.From neuronal level we can go up to neuronal circuits, to cortical structures, to thewhole bra<strong>in</strong>, and f<strong>in</strong>ally to the behavior of the organism. So, let us see how much weunderstand of what is go<strong>in</strong>g on at the level of <strong>in</strong>dividual neurons.1

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