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Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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280 Simple Modelsspikemembrane potential, VE threshE leakE KtimeFigure 8.1: Leaky <strong>in</strong>tegrate-and-fire neuron with noisy <strong>in</strong>put. The spike is addedmanually for aesthetic purposes and to fool the reader <strong>in</strong>to believ<strong>in</strong>g that this is aspik<strong>in</strong>g neuron.“spik<strong>in</strong>g models”; the spikes <strong>in</strong> the next two figures, as well as <strong>in</strong> hundreds of scientificpapers devoted to these models, are drawn by hand. The quadratic <strong>in</strong>tegrate-and-firemodel is the simplest truly spik<strong>in</strong>g model.8.1.1 Integrate-and-fireThe leaky <strong>in</strong>tegrate-and-fire model (Lapicque 1907, Ste<strong>in</strong> 1967, Tuckwell 1988) is anidealization of a neuron hav<strong>in</strong>g Ohmic leakage current and a number of voltage-gatedcurrents that are completely de-activated at rest. Subthreshold behavior of such aneuron can be described by the l<strong>in</strong>ear differential equationC ˙V = I −Ohmic leakage{ }} {g leak (V − E leak ) ,where all parameters have the same biophysical mean<strong>in</strong>gs as <strong>in</strong> the previous chapters.When the membrane potential V reaches the threshold value E thresh , the voltagesensitivecurrents <strong>in</strong>stantaneously activate, the neuron is said to fire an action potential,and V is reset to E K , as <strong>in</strong> Fig. 8.1. After appropriate re-scal<strong>in</strong>g, the leaky <strong>in</strong>tegrateand-firemodel can be written <strong>in</strong> the form˙v = b − v , if v = 1, then v ← 0, (8.1)where the rest<strong>in</strong>g state is v = b, the threshold value is v = 1 and the reset value isv = 0. Apparently, the neuron is excitable when b < 1 and fires a periodic spike tra<strong>in</strong>when b > 1 with period T = − ln(1 − 1/b) (verify that).The <strong>in</strong>tegrate-and-fire neuron illustrates a number of important neuro-computationalproperties:• All-or-none spikes. S<strong>in</strong>ce the shape of the spike is not simulated, all spikes areimplicitly assumed to be identical <strong>in</strong> size and duration.

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