Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience: Dynamical Systems in Neuroscience:

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474 Synchronization (see www.izhikevich.com)Figure 10.20: Yoshiki Kuramoto in1988, while he was visiting Jim Murray’sinstitute at University of Oxford.rewrite the corresponding Poincare phase map (10.3)PRC{ }} {ϑ(t n+1 ) = {ϑ(t n ) + Z(ϑ(t n )) εp(t n )h +h} mod T} {{ }Ain the formϑ(t n + h) − ϑ(t n )hwhich is a discrete version of= Z(ϑ(t n ))εp(t n ) + 1 ,˙ϑ = 1 + εZ(ϑ) · p(t) , (10.6)in the limit h → 0.To be consistent with all the examples in the previous section, we implicitly assumehere that p(t) perturbs only the first, voltage-like variable x 1 of the state vector x =(x 1 , . . . , x m ) ∈ R m and Z(ϑ) is the corresponding sensitivity function. However, thephase model (10.6) is also valid for an arbitrary input p(t) = (p 1 (t), . . . , p m (t)). Indeed,let Z i describe the linear response to perturbations of the ith state variable x i , andZ(ϑ) = (Z 1 (ϑ), . . . , Z m (ϑ)) denote the corresponding linear response vector-function.Then, the combined phase shift Z 1 p 1 + · · · + Z m p m is just the dot product Z · p in(10.6).10.2.2 Kuramoto’s approachConsider the unperturbed (ε = 0) oscillator (10.5), and let the function ϑ(x) denotethe phases of points near its limit cycle attractor. Obviously, isochrons are the levelcontours of ϑ(x) since the function is constant on each isochron. Differentiating thefunction using the chain rule yieldsdϑ(x)dt= grad ϑ · dxdt= grad ϑ · f(x) ,

Synchronization (see www.izhikevich.com) 475xgrad (x)isochronf(x)limit cycleFigure 10.21: Geometrical interpretationof the vector grad ϑ.where grad ϑ = (ϑ x1 (x), . . . , ϑ xm (x)) is the gradient of ϑ(x) with respect to the statevector x = (x 1 , . . . , x m ) ∈ R m . However,dϑ(x)dtnear the limit cycle, because isochrons are mapped to isochrons by the flow of thevector-field f(x). Therefore, we get a useful equality= 1grad ϑ · f(x) = 1 . (10.7)Figure 10.21 shows a geometrical interpretation of grad ϑ(x): it is the vector based atpoint x, normal to the isochron of x and with the length equal to the number densityof isochrons at x. Its length can also be found from (10.7).Kuramoto (1984) applied the chain rule to the perturbed system (10.5)dϑ(x)dt= grad ϑ · dxdtand, using (10.7), obtained the phase model= grad ϑ · {f(x) + εp(t)} = grad ϑ · f(x) + ε grad ϑ · p(t) ,˙ϑ = 1 + ε grad ϑ · p(t) , (10.8)which has the same form as (10.6). Subtracting (10.8) from (10.6) yields (Z(ϑ) −grad ϑ) · p(t) = 0. Since this is valid for any p(t), we conclude that Z(ϑ) = grad ϑ;see also Ex. 6. Thus, Kuramoto’s phase model (10.8) is indeed equivalent to Winfree’smodel (10.8).10.2.3 Malkin’s approachYet another equivalent method of reduction of weakly perturbed oscillators to theirphase models follows from Malkin theorem (1949,1956), which we state in the simplestform below. The most abstract form and its proof is provided by Hoppensteadt andIzhikevich (1997).Malkin’s theorem. Suppose the unperturbed (ε = 0) oscillator in (10.5) has anexponentially stable limit cycle of period T . Then its phase is described by the equation˙ϑ = 1 + εQ(ϑ) · p(t) , (10.9)

474 Synchronization (see www.izhikevich.com)Figure 10.20: Yoshiki Kuramoto <strong>in</strong>1988, while he was visit<strong>in</strong>g Jim Murray’s<strong>in</strong>stitute at University of Oxford.rewrite the correspond<strong>in</strong>g Po<strong>in</strong>care phase map (10.3)PRC{ }} {ϑ(t n+1 ) = {ϑ(t n ) + Z(ϑ(t n )) εp(t n )h +h} mod T} {{ }A<strong>in</strong> the formϑ(t n + h) − ϑ(t n )hwhich is a discrete version of= Z(ϑ(t n ))εp(t n ) + 1 ,˙ϑ = 1 + εZ(ϑ) · p(t) , (10.6)<strong>in</strong> the limit h → 0.To be consistent with all the examples <strong>in</strong> the previous section, we implicitly assumehere that p(t) perturbs only the first, voltage-like variable x 1 of the state vector x =(x 1 , . . . , x m ) ∈ R m and Z(ϑ) is the correspond<strong>in</strong>g sensitivity function. However, thephase model (10.6) is also valid for an arbitrary <strong>in</strong>put p(t) = (p 1 (t), . . . , p m (t)). Indeed,let Z i describe the l<strong>in</strong>ear response to perturbations of the ith state variable x i , andZ(ϑ) = (Z 1 (ϑ), . . . , Z m (ϑ)) denote the correspond<strong>in</strong>g l<strong>in</strong>ear response vector-function.Then, the comb<strong>in</strong>ed phase shift Z 1 p 1 + · · · + Z m p m is just the dot product Z · p <strong>in</strong>(10.6).10.2.2 Kuramoto’s approachConsider the unperturbed (ε = 0) oscillator (10.5), and let the function ϑ(x) denotethe phases of po<strong>in</strong>ts near its limit cycle attractor. Obviously, isochrons are the levelcontours of ϑ(x) s<strong>in</strong>ce the function is constant on each isochron. Differentiat<strong>in</strong>g thefunction us<strong>in</strong>g the cha<strong>in</strong> rule yieldsdϑ(x)dt= grad ϑ · dxdt= grad ϑ · f(x) ,

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