Dynamical Systems in Neuroscience:
Dynamical Systems in Neuroscience: Dynamical Systems in Neuroscience:
470 Synchronization (see www.izhikevich.com)synchronization (1:1)poincare phase mapphase-lockedphase-locking (2:2)n+1=f nsync0nFigure 10.14: Co-existence of synchronized and phase-locked solutions corresponds toco-existence of stable fixed point and a stable periodic orbit of the Poincare phase map.correspond to stable equilibria in the iterates ϑ k+1 = f q (ϑ k ), where f q = f ◦ f ◦ · · · ◦ fis the composition of f with itself q times. Geometrically, studying such maps is likeconsidering every q-th input pulse in Fig. 10.12b and ignoring all the intermediatepulses.Since maps can have co-existence of stable fixed points and periodic orbits, varioussynchronized and phase-locking states can co-exist in response to the same input pulsetrain, as in Fig. 10.14. The oscillator converges to one of the states depending on theinitial phase of oscillation, but could be switched between the states by a transientinput.10.1.9 Arnold tonguesTo synchronize an oscillator, the input pulse train must have period T s sufficiently nearthe oscillator’s free-running period T so that the graph of PRC and the horizontal linein Fig. 10.13 intersect. The amplitude of the function |PRC (ϑ, A)| decreases as thestrength of the pulse A decreases, because weaker pulses produce weaker phase shifts.Hence the region of existence of a synchronized state shrinks as A → 0, and it looks likea horn or a tongue on the (T s , A)-plane depicted in Fig. 10.15, called Arnold tongue.Each p:q-phase-locked state has its own region of existence (p:q-tongue in the figure),also shrinking to a point pT/q on the T s -axis. The larger the order of locking, p + q,the narrower the tongue, and the more difficult it is to observe such a phase-lockedstate numerically, let alone experimentally.The tongues can overlap, leading to the co-existence of phase-locked states, as inFig. 10.14. If A is sufficiently large, the Poincare phase map (10.3) becomes noninvertible,i.e., it has a region of negative slope, and there is a possibility of chaoticdynamics (Glass and MacKey 1988).In Fig. 10.16 we illustrate the cycle slipping phenomenon that occurs when the inputperiod T s drifts away from the 1:1 Arnold tongue. The fixed point of the Poincarephase map corresponding to the synchronized state undergoes a fold bifurcation anddisappears. Similarly to the case of saddle-node on invariant circle bifurcation, the
Synchronization (see www.izhikevich.com) 471amplitude of stimulation, A1:61:4synchronization1:2 3:21:11:3 2:32:15:20T/4 T/3 T/2 3T/4 T 5T/4 3T/2 7T/4 2T 9T/4 5T/2period of stimulation, TsFigure 10.15: Arnold tongues are regions of existence of various phase-locked states onthe “period-strength” plane.cycle slippingT/2Poincare phase mapn+1=f( n)0?ghost ofattractor-T/2-T/2 0 n T/2Figure 10.16: Cycle slipping phenomenon at the edge of the Arnold tongue correspondingto a synchronized state.fold fixed point becomes a ghost attractor that traps orbits and keeps them near thesynchronized state for a long period of time. Eventually the orbit escapes, the synchronizedstate is briefly lost, and then the orbit returns to the ghost attractor to betrapped again. Such an intermittently synchronized orbit typically corresponds to ap:q-phase-locked state with high order of locking p + q.10.2 Weak CouplingIn this section we consider dynamical systems of the formẋ = f(x) + εp(t) , (10.5)describing periodic oscillators, ẋ = f(x), forced by a time-depended input εp(t), e.g.,from other oscillators in a network. The positive parameter ε measures the overallstrength of the input, and it is assumed to be sufficiently small, denoted as ε ≪ 1. Wedo not assume ε → 0 here. In fact, most of the results in this section can be cast in
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- Page 452 and 453: 442 Referencesterneurons mediated b
- Page 454 and 455: 444 ReferencesDickson C.T., Magistr
- Page 456 and 457: 446 ReferencesGuckenheimer J. (1975
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- Page 464 and 465: 454 ReferencesTuckwell H.C. (1988)
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470 Synchronization (see www.izhikevich.com)synchronization (1:1)po<strong>in</strong>care phase mapphase-lockedphase-lock<strong>in</strong>g (2:2)n+1=f nsync0nFigure 10.14: Co-existence of synchronized and phase-locked solutions corresponds toco-existence of stable fixed po<strong>in</strong>t and a stable periodic orbit of the Po<strong>in</strong>care phase map.correspond to stable equilibria <strong>in</strong> the iterates ϑ k+1 = f q (ϑ k ), where f q = f ◦ f ◦ · · · ◦ fis the composition of f with itself q times. Geometrically, study<strong>in</strong>g such maps is likeconsider<strong>in</strong>g every q-th <strong>in</strong>put pulse <strong>in</strong> Fig. 10.12b and ignor<strong>in</strong>g all the <strong>in</strong>termediatepulses.S<strong>in</strong>ce maps can have co-existence of stable fixed po<strong>in</strong>ts and periodic orbits, varioussynchronized and phase-lock<strong>in</strong>g states can co-exist <strong>in</strong> response to the same <strong>in</strong>put pulsetra<strong>in</strong>, as <strong>in</strong> Fig. 10.14. The oscillator converges to one of the states depend<strong>in</strong>g on the<strong>in</strong>itial phase of oscillation, but could be switched between the states by a transient<strong>in</strong>put.10.1.9 Arnold tonguesTo synchronize an oscillator, the <strong>in</strong>put pulse tra<strong>in</strong> must have period T s sufficiently nearthe oscillator’s free-runn<strong>in</strong>g period T so that the graph of PRC and the horizontal l<strong>in</strong>e<strong>in</strong> Fig. 10.13 <strong>in</strong>tersect. The amplitude of the function |PRC (ϑ, A)| decreases as thestrength of the pulse A decreases, because weaker pulses produce weaker phase shifts.Hence the region of existence of a synchronized state shr<strong>in</strong>ks as A → 0, and it looks likea horn or a tongue on the (T s , A)-plane depicted <strong>in</strong> Fig. 10.15, called Arnold tongue.Each p:q-phase-locked state has its own region of existence (p:q-tongue <strong>in</strong> the figure),also shr<strong>in</strong>k<strong>in</strong>g to a po<strong>in</strong>t pT/q on the T s -axis. The larger the order of lock<strong>in</strong>g, p + q,the narrower the tongue, and the more difficult it is to observe such a phase-lockedstate numerically, let alone experimentally.The tongues can overlap, lead<strong>in</strong>g to the co-existence of phase-locked states, as <strong>in</strong>Fig. 10.14. If A is sufficiently large, the Po<strong>in</strong>care phase map (10.3) becomes non<strong>in</strong>vertible,i.e., it has a region of negative slope, and there is a possibility of chaoticdynamics (Glass and MacKey 1988).In Fig. 10.16 we illustrate the cycle slipp<strong>in</strong>g phenomenon that occurs when the <strong>in</strong>putperiod T s drifts away from the 1:1 Arnold tongue. The fixed po<strong>in</strong>t of the Po<strong>in</strong>carephase map correspond<strong>in</strong>g to the synchronized state undergoes a fold bifurcation anddisappears. Similarly to the case of saddle-node on <strong>in</strong>variant circle bifurcation, the