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Dynamical Systems in Neuroscience:

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Synchronization (see www.izhikevich.com) 491- +spike- + - +spike spikex(t)spikex(t)=-cot tspike-1 1000x'=-1+x2 x'=0+x2 x'=1+x2Figure 10.34: Phase portraits and typical oscillations of the quadratic <strong>in</strong>tegrate-and-fireneuron ẋ = I + x 2 with x ∈ R ∪ {±∞}. Parameter: I = −1, 0, +1.x'=1+x2A0xnormalized PRC (PRC( ,A)/A)100A=1A=2A=3A=0.1phase of oscillation,TFigure 10.35: The dependence of PRC of the quadratic <strong>in</strong>tegrate-and-fire model on thestrength of the pulse A.on <strong>in</strong>variant circle (SNIC) bifurcation studied <strong>in</strong> Sect. 6.1.2. Appropriate rescal<strong>in</strong>g ofthe membrane potential and time converts the model <strong>in</strong>to the normal formx ′ = 1 + x 2 , x ∈ R .Because of the quadratic term, x escapes to the <strong>in</strong>f<strong>in</strong>ity <strong>in</strong> a f<strong>in</strong>ite time, produc<strong>in</strong>ga spike depicted <strong>in</strong> Fig. 10.34. If we identify −∞ and +∞, then x exhibits periodicspik<strong>in</strong>g of <strong>in</strong>f<strong>in</strong>ite amplitude. Such a spik<strong>in</strong>g model is called quadratic <strong>in</strong>tegrate-and-fire(QIF) neuron; see also Sect. 8.1.3 for some generalizations of the model.Strong pulseThe solution of this system start<strong>in</strong>g at the spike, i.e., at x(0) = ±∞, isx(t) = − cot t ,as the reader can check by differentiat<strong>in</strong>g. It is a periodic function with T = π,hence, we can <strong>in</strong>troduce the phase of oscillation via the relation x = − cot ϑ. Thecorrespond<strong>in</strong>g PRC can be found explicitly (see Ex. 9) and it has the formPRC (ϑ, A) = π/2 + atan (A − cot ϑ) − ϑ ,

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