Dynamical Systems in Neuroscience:
Dynamical Systems in Neuroscience: Dynamical Systems in Neuroscience:
466 Synchronization (see www.izhikevich.com)(a)membrane potential, V(c)phase, n101 2 3 4 5 6 7?T stime, tT345 6 71K + gating variable, b(b)0.60.40.2(d)Tmod Tn+10314-7-80 -60 -40 -20 0 20membrane potential, V102Poincare phase map315,6,74stablefixedpoint021 2 3 4 5 6 7pulse number, n020 10nTFigure 10.10: Description of synchronization of I Na + I K -oscillator in Fig. 10.4 usingPoincare phase map.10.1.5 Poincare phase mapThe phase resetting curve (PRC) describes the response of an oscillator to a singlepulse, but it can be used to study its response to a periodic pulse train using thefollowing “stroboscopic” approach. Let ϑ n denote the phase of oscillation at the timethe nth input pulse arrives. Such a pulse resets the phase by PRC (ϑ n ), so that the newphase right after the pulse is ϑ n +PRC (ϑ n ); see Fig. 10.9. Let T s denote the period ofpulsed stimulation. Then the phase of oscillation before the next, (n + 1)th pulse, isϑ n +PRC (ϑ n ) + T s . Thus, we have a stroboscopic mapping of a circle to itselfϑ n+1 = (ϑ n + PRC (ϑ n ) + T s ) mod T (10.3)called Poincare phase map (two pulse-coupled oscillators are considered in Ex. 11).Knowing the initial phase of oscillation ϑ 1 at the first pulse, we can determine ϑ 2 , thenϑ 3 , etc. The sequence {ϑ n } with n = 1, 2, . . . , is called the orbit of the map, and it isquite easy to find numerically.Let us illustrate this concept using the I Na + I K -oscillator with PRC shown inFig. 10.4. Its free-running period is T ≈ 21.37 ms, and the period of stimulationin Fig. 10.10a is T s = 18.37, which results in the Poincare phase map depicted inFig. 10.10d. The cobweb in the figure is the orbit going from ϑ 1 to ϑ 2 to ϑ 3 , etc. Notice
Synchronization (see www.izhikevich.com) 467that the phase ϑ 3 cannot be measured directly from the voltage trace in Fig. 10.10abecause pulse 2 changes the phase, so it is not the time since the last spike when pulse3 arrives. The Poincare phase map (10.3) takes into account such multiple pulses.The orbit approaches a point (called fixed point, see below) that corresponds to asynchronized or phase-locked state.A word of caution is in order. Recall that PRCs are measured on the limit cycleattractor. However, each pulse displaces the trajectory away from the attractor, asin Fig. 10.5. To use the PRC formalism to describe the effect of the next pulse, theoscillator must be given enough time to relax back to the limit cycle attractor. Thus,if the period of stimulation T s is too small, or the attraction to the limit cycle is tooslow, or the stimulus amplitude is too large, then the Poincare phase map may be notan appropriate tool to describe the phase dynamics.10.1.6 Fixed pointsTo understand the structure of orbits of the Poincare phase map (10.3), or any othermapϑ n+1 = f(ϑ n ) , (10.4)we need to find its fixed pointsϑ = f(ϑ)(ϑ is a fixed point),which are analogues of equilibria of continuous dynamical systems. Geometrically, afixed point is the intersection of the graph of f(ϑ) with the diagonal line ϑ n+1 = ϑ n ;see Fig. 10.10d or Fig. 10.11. At such a point, the orbit ϑ n+1 = f(ϑ n ) = ϑ n is fixed.A fixed point ϑ is asymptotically stable if it attracts all nearby orbits, i.e., if ϑ 1 is ina sufficiently small neighborhood of ϑ, then ϑ n → ϑ as n → ∞, as in Fig. 10.11, left.The fixed point is unstable if any small neighborhood of the point contains an orbitdiverging from it, as in Fig. 10.11, right.The stability of the fixed point is determined by the slopem = f ′ (ϑ)of the graph of f at the point, which is called Floquet multiplier of the mapping. It playsthe same role as the eigenvalue λ of an equilibrium of a continuous dynamical system.Mnemonically, the relationship between them is µ = e λ , to that the fixed point is stablewhen |m| < 1 (λ < 0) and unstable when |m| > 1 (λ > 0). Fixed points bifurcatewhen |m| = 1 (λ is zero or purely imaginary). They lose stability via flip bifurcation (adiscrete analogue of Andronov-Hopf bifurcation) when m = −1 and disappear via foldbifurcation (a discrete analogue of saddle-node bifurcation) when m = 1. The formerplays an important role in the period-doubling phenomenon illustrated in Fig. 10.14,bottom trace. The latter plays an important role in the cycle slipping phenomenonillustrated in Fig. 10.16.
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- Page 452 and 453: 442 Referencesterneurons mediated b
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Synchronization (see www.izhikevich.com) 467that the phase ϑ 3 cannot be measured directly from the voltage trace <strong>in</strong> Fig. 10.10abecause pulse 2 changes the phase, so it is not the time s<strong>in</strong>ce the last spike when pulse3 arrives. The Po<strong>in</strong>care phase map (10.3) takes <strong>in</strong>to account such multiple pulses.The orbit approaches a po<strong>in</strong>t (called fixed po<strong>in</strong>t, see below) that corresponds to asynchronized or phase-locked state.A word of caution is <strong>in</strong> order. Recall that PRCs are measured on the limit cycleattractor. However, each pulse displaces the trajectory away from the attractor, as<strong>in</strong> Fig. 10.5. To use the PRC formalism to describe the effect of the next pulse, theoscillator must be given enough time to relax back to the limit cycle attractor. Thus,if the period of stimulation T s is too small, or the attraction to the limit cycle is tooslow, or the stimulus amplitude is too large, then the Po<strong>in</strong>care phase map may be notan appropriate tool to describe the phase dynamics.10.1.6 Fixed po<strong>in</strong>tsTo understand the structure of orbits of the Po<strong>in</strong>care phase map (10.3), or any othermapϑ n+1 = f(ϑ n ) , (10.4)we need to f<strong>in</strong>d its fixed po<strong>in</strong>tsϑ = f(ϑ)(ϑ is a fixed po<strong>in</strong>t),which are analogues of equilibria of cont<strong>in</strong>uous dynamical systems. Geometrically, afixed po<strong>in</strong>t is the <strong>in</strong>tersection of the graph of f(ϑ) with the diagonal l<strong>in</strong>e ϑ n+1 = ϑ n ;see Fig. 10.10d or Fig. 10.11. At such a po<strong>in</strong>t, the orbit ϑ n+1 = f(ϑ n ) = ϑ n is fixed.A fixed po<strong>in</strong>t ϑ is asymptotically stable if it attracts all nearby orbits, i.e., if ϑ 1 is <strong>in</strong>a sufficiently small neighborhood of ϑ, then ϑ n → ϑ as n → ∞, as <strong>in</strong> Fig. 10.11, left.The fixed po<strong>in</strong>t is unstable if any small neighborhood of the po<strong>in</strong>t conta<strong>in</strong>s an orbitdiverg<strong>in</strong>g from it, as <strong>in</strong> Fig. 10.11, right.The stability of the fixed po<strong>in</strong>t is determ<strong>in</strong>ed by the slopem = f ′ (ϑ)of the graph of f at the po<strong>in</strong>t, which is called Floquet multiplier of the mapp<strong>in</strong>g. It playsthe same role as the eigenvalue λ of an equilibrium of a cont<strong>in</strong>uous dynamical system.Mnemonically, the relationship between them is µ = e λ , to that the fixed po<strong>in</strong>t is stablewhen |m| < 1 (λ < 0) and unstable when |m| > 1 (λ > 0). Fixed po<strong>in</strong>ts bifurcatewhen |m| = 1 (λ is zero or purely imag<strong>in</strong>ary). They lose stability via flip bifurcation (adiscrete analogue of Andronov-Hopf bifurcation) when m = −1 and disappear via foldbifurcation (a discrete analogue of saddle-node bifurcation) when m = 1. The formerplays an important role <strong>in</strong> the period-doubl<strong>in</strong>g phenomenon illustrated <strong>in</strong> Fig. 10.14,bottom trace. The latter plays an important role <strong>in</strong> the cycle slipp<strong>in</strong>g phenomenonillustrated <strong>in</strong> Fig. 10.16.