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Dynamical Systems in Neuroscience:

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Synchronization (see www.izhikevich.com) 487From ϕ ′ n = ω we f<strong>in</strong>d that ω 0 = ω, i.e., the common frequency is the frequency ofthe free oscillation of the last, uncoupled oscillator. The phase lag along the cha<strong>in</strong>,χ = ϕ i+1 − ϕ i , satisfies n − 1 identical conditions 0 = H + (χ). Thus, the travel<strong>in</strong>g wavewith a constant phase shift exists when H + has a zero cross<strong>in</strong>g with positive slope, <strong>in</strong>contrast to Fig. 10.26. The sign of χ, and not the direction of coupl<strong>in</strong>g, determ<strong>in</strong>es thedirection of wave propagation.10.3.3 NetworksNow let us consider weakly connected networks (10.11) with arbitrary, possibly allto all coupl<strong>in</strong>g. To study synchronized states of the network, we need to determ<strong>in</strong>ewhether the correspond<strong>in</strong>g phase model (10.15) has equilibria and exam<strong>in</strong>e their stabilityproperties. A vector φ = (φ 1 , . . . , φ n ) is an equilibrium of (10.15) whenn∑0 = ω i + H ij (φ j − φ i ) for all i . (10.19)j≠1It is stable when all eigenvalues of the l<strong>in</strong>earization matrix (Jacobian) at φ have negativereal parts, except one zero eigenvalue correspond<strong>in</strong>g to the eigenvector along thecircular family of equilibria (φ plus a phase shift is a solution of (10.19) too s<strong>in</strong>ce thephase differences φ j − φ i are not affected).In general, determ<strong>in</strong><strong>in</strong>g the stability of equilibria is a difficult problem. Ermentrout(1992) found a simple sufficient condition. Namely, if• a ij = H ′ ij(φ j − φ i ) ≥ 0, and• the directed graph def<strong>in</strong>ed by the matrix a = (a ij ) is connected, (i.e., each oscillatoris <strong>in</strong>fluenced, possibly <strong>in</strong>directly, by every other oscillator),then the equilibrium φ is neutrally stable, and the correspond<strong>in</strong>g limit cycle x(t + φ)of (10.11) is asymptotically stable.Another sufficient condition was found by Hoppensteadt and Izhikevich (1997). Itstates that if system (10.15) satisfies• ω 1 = · · · = ω n = ω(identical frequencies), and• H ij (−χ) = −H ji (χ) (pair-wise odd coupl<strong>in</strong>g)for all i and j, then the network dynamics converge to a limit cycle. On the cycle, alloscillators have equal frequencies 1 + εω and constant phase deviations.The proof follows from the observation that (10.15) is a gradient system <strong>in</strong> therotat<strong>in</strong>g coord<strong>in</strong>ates ϕ = ωτ + φ, with the energy functionE(φ) = 1 n∑ n∑∫ χR ij (φ j − φ i ) , where R ij (χ) = H ij (s) ds .2i=1j=1One can check that dE(φ)/dτ = − ∑ (φ ′ i) 2 ≤ 0 along the trajectories of (10.15), withequality only at equilibria.0

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