Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience: Dynamical Systems in Neuroscience:

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482 Synchronization (see www.izhikevich.com)frequency lockingin-phaseentrainment(1:1 frequency locking)synchronizationphase lockinganti-phaseFigure 10.28: Various degrees of locking of oscillators.The coupled oscillators above are said to be frequency locked when there is a periodictrajectory on the 2-torus, which is called a torus knot. It is said to be of type (p, q)if ϑ 1 makes p rotations while ϑ 2 makes q rotations, and p and q are relatively primeintegers, i.e., do not have a common divisor greater than 1. Torus knots of type (p, q)produce p:q frequency locking, e.g., 2:3 frequency locking in Fig. 10.27. A 1:1 frequencylocking is called entrainment. There could be many periodic orbits on the torus, withstable orbits between unstable ones. Since the orbits on the 2-torus cannot intersect,they all are knots of the same type, resulting in the same p:q frequency locking.Let us follow a trajectory on the torus and count the number of rotations of thephase variables. The limit of the ratio of rotations as t → ∞ is independent on thetrajectory we follow, and it is called the rotation number of the torus flow. It is rationalif and only if there is a (p, q) periodic orbit, in which case the rotation number is p/q.An irrational rotation number implies there are no periodic orbits, and it corresponds toa quasi-periodic or multifrequency torus flow. Oscillators exhibit phase drifting in thiscase. Denjoy (1932) proved that such coupled oscillators are topologically equivalentto the uncoupled system ˙ϑ 1 = r, ˙ϑ2 = 1 with irrational r.Suppose the oscillators are frequency locked; that is, there is a p : q limit cycleattractor on the torus. We say that the oscillators are p : q phase locked ifqϑ 1 (t) − pϑ 2 (t) = conston the cycle. The value of the constant determines whether the locking is in-phase(const= 0), anti-phase (const= T/2, half-period), or out-of-phase. Frequency lockingdoes not necessarily imply phase locking: The (2, 3) torus knot in Fig. 10.27b correspondsto phase locking, whereas that in Fig. 10.27c does not. Frequency lockingwithout phase locking is called phase trapping. Finally, synchronization is a 1:1 phaselocking. The phase difference ϑ 2 − ϑ 1 is also called phase lag or phase lead. Therelationships between all these definitions are shown in Fig. 10.28.Frequency locking, phase locking, entrainment, and synchronization of a networkof n > 2 oscillators is the same as pair-wise locking, entrainment, and synchronization

Synchronization (see www.izhikevich.com) 483Figure 10.29: A major part of computational neuroscience concerns coupled oscillators.of the oscillators comprising the network. In addition, a network can exhibit partialsynchronization, when only a subset of oscillators is synchronized.Synchronization of oscillators with nearly identical frequencies is described by thephase model (10.15). Existence of one equilibrium of (10.15) implies the existence ofthe entire circular family of equilibria, since translation of all ϕ i by a constant phaseshift does not change the phase differences ϕ j − ϕ i and hence the form of (10.15).This family corresponds to a limit cycle of (10.11), on which all oscillators, x i (t + ϕ i ),have equal frequencies and constant phase shifts, i.e., they are synchronized, possiblyout-of-phase.10.3.1 Two oscillatorsConsider (10.11) with n = 2, describing two coupled oscillators, as in Fig. 10.29. Letus introduce the “slow” time τ = εt and rewrite the corresponding phase model (10.15)in the formϕ ′ 1 = ω 1 + H 1 (ϕ 2 − ϕ 1 ) ,ϕ ′ 2 = ω 2 + H 2 (ϕ 1 − ϕ 2 ) ,where ′ = d/dτ is the derivative with respect to slow time. Let χ = ϕ 2 − ϕ 1 denotethe phase difference between the oscillators. Then the two-dimensional system abovebecomes one-dimensionalχ ′ = ω + G(χ) , (10.17)

Synchronization (see www.izhikevich.com) 483Figure 10.29: A major part of computational neuroscience concerns coupled oscillators.of the oscillators compris<strong>in</strong>g the network. In addition, a network can exhibit partialsynchronization, when only a subset of oscillators is synchronized.Synchronization of oscillators with nearly identical frequencies is described by thephase model (10.15). Existence of one equilibrium of (10.15) implies the existence ofthe entire circular family of equilibria, s<strong>in</strong>ce translation of all ϕ i by a constant phaseshift does not change the phase differences ϕ j − ϕ i and hence the form of (10.15).This family corresponds to a limit cycle of (10.11), on which all oscillators, x i (t + ϕ i ),have equal frequencies and constant phase shifts, i.e., they are synchronized, possiblyout-of-phase.10.3.1 Two oscillatorsConsider (10.11) with n = 2, describ<strong>in</strong>g two coupled oscillators, as <strong>in</strong> Fig. 10.29. Letus <strong>in</strong>troduce the “slow” time τ = εt and rewrite the correspond<strong>in</strong>g phase model (10.15)<strong>in</strong> the formϕ ′ 1 = ω 1 + H 1 (ϕ 2 − ϕ 1 ) ,ϕ ′ 2 = ω 2 + H 2 (ϕ 1 − ϕ 2 ) ,where ′ = d/dτ is the derivative with respect to slow time. Let χ = ϕ 2 − ϕ 1 denotethe phase difference between the oscillators. Then the two-dimensional system abovebecomes one-dimensionalχ ′ = ω + G(χ) , (10.17)

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