12.07.2015 Views

Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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282 Simple Models1thresholdyz resetx0-1x-1 0 1100 20 40 60 80 100time, msFigure 8.2: Resonate-and-fire model with b = −0.05, ω = 0.25 and z reset = i. The spikewas added manually.spike. Young did not specify what happens after the spike. The resonate-and-firemodel is the Young model with the resett<strong>in</strong>g: if V ≥ V thresh , then V ← V reset andW ← W reset , where V rest and W reset are some parameters.When the rest<strong>in</strong>g state is a stable focus, the model can be recast <strong>in</strong> complex coord<strong>in</strong>atesasż = (b + iω)z + I ,where b + iω ∈ C is the complex eigenvalue of the rest<strong>in</strong>g state, z = x + iy ∈ C is thecomplex-valued variable describ<strong>in</strong>g damped oscillations with frequency ω around therest<strong>in</strong>g state. The real part, x, is a current-like variable. It describes the dynamicsof the resonant current and synaptic currents. The imag<strong>in</strong>ary part, y is a voltage-likevariable. The neuron is said to fire a spike when y reaches the threshold y = 1. Thus,the threshold is a horizontal l<strong>in</strong>e on the complex plane that passes through i ∈ C, as<strong>in</strong> Fig. 8.2, though other choices are also possible. After fir<strong>in</strong>g the spike, the variablez is reset to z reset .The resonate-and-fire model illustrates the most important features of resonators:damped oscillations, frequency preference, post-<strong>in</strong>hibitory (rebound) spikes, and Class2 excitability. It cannot have susta<strong>in</strong>ed subthreshold oscillations of membrane potential.Integrate-and-fire and resonate-and-fire neurons do not contradict, but complementeach other. Both are l<strong>in</strong>ear, and hence are useful when we prove theorems and deriveanalytical expressions. They have the same flaws limit<strong>in</strong>g their applicability, which wediscussed earlier. In contrast, two simple models described below are difficult to treatanalytically, but because of their universality they should be the models of choice whenlarge-scale simulations are concerned.8.1.3 Quadratic <strong>in</strong>tegrate-and-fireSubstitut<strong>in</strong>g −v by +v 2 <strong>in</strong> (8.1) results <strong>in</strong> the quadratic <strong>in</strong>tegrate-and-fire model˙v = b + v 2 , if v = v peak , then v ← v reset , (8.2)

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