Introduction to Neural Dynamics and Signal Transmission Delay

(5.3.12)

φ = θ +ST +2nj,

(5.3.13)

where j is an integer chosen so that 0 < φ, θ < 2π. Assume R = \b\ < 1. If r > 1 then by (5.3.12) we have ρ < 1, which is a contradiction since ρ = -Jr2 + s2 > r. Therefore, R = \b\ < 1 implies r < 1, and thusReA = r - 1 < 0. If/? = \b\ = l a n d b φ 1,then(5.3.12)impliespe {r ~ X)T = 1. This, together with ρ > r, implies either r < l o r p = r = l. In the latter case, we have s = 0 and hence Θ = 0, φ = 0 and b = Re1^ = 1, a contradiction. This proves the sufficiency. To show the necessity, we fix b so that \b\ = R > 1. Choose s = Λ/R2 — 1, θ - arctan VR2 - 1 e (0, f ) , r - Φ-θ+2π and λ = is. Then at the above values of τ and λ, both sides of (5.3.10) are iy/R2 - 1 = - 1 + Re1'* β-^φ-θ+1π)

= - 1 + Re1'* .

This shows that for the given r, (5.3.10) has a zero of zero real part. This completes the proof. Remark 5.3.8. The work of Belair [1993] suggested that for a fixed τ > 0, all zeros of (5.3.10) have negative real parts if and only if b lies inside the domain bounded by the arcs of Archimedean spirals R = (φ— f ) / τ and R = — φ)/τ for j < φ < π. Such a domain, called the zone of stability, is a teardrop shaped region, and approaches the half plane Re(fr) < 1 in the limit as τ —> 0 + , and the circle of radius 1 as r —• oo. See Figure 5.3.1. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

103

5.3 Delay-induced instability

0

Figure 5.3.1. A teardrop region as the zone of stability of λ + 1 = be~^ T for a fixed τ > 0 and its limiting regions: a half plane when r —• 0 and the unit circle when τ —»• oo.

The above discussion shows that it is possible for the delay τ to destabilize an equilibrium which is asymptotically stable in the absence of time-delay, and a necessary and sufficient condition for the occurrence of delay-induced instability of the zero solution of (5.3.2) in terms of o(W) can be stated as s(W) < l < p(W), Ρ

(5.3.14)

where s(W) = max{ReX; λ e σ(Ψ)}, p(W) = max{|X|;Ae^(W)}. By contrast, if W) > 1 /β, then the zero solution is unstable for all τ; if \/β > p(W) and j φ a(W) then the zero solution is asymptotically stable for all τ. Recall that we say matrix W has the Perron property if p{W) e σ(Ψ), that is, p(W) = s(W). Therefore, equation (5.3.14) can be interpreted as (see Belair, Campbell and van den Driessche [ 1996]) Delay-induced instabilityfor the zero solution of equation (5.3.2) is impossible for any choice of the neuron gain if and only if the connection matrix W has the Perron property. Recall that directed graph (digraph) of a real η xn matrix W = (ti>,7) consists of a set of vertices { 1 , 2 , . . . , « } and a set of directed edges, with edge set {e ji; wij φ 0}. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5 Signal transmission delays

A cycle of length k in this graph is a set of edges {e ili2}, { e ,2,-3}, . . . , {e ikil} with vertices i\, ... , ik distinct. The signed digraph of W, D(W), is obtained from the digraph by attaching to each edge ( j , i) the sign of . The cycle is positive (respectively negative) according as the product u>,-2,·, ιυ; 3 ,· 2 ... is positive (respectively negative). A network is called frustrated if the signed digraph of its connection matrix W has a negative cycle. In Belair, Campbell and van den Driessche [1996], it is proved that Delay-induced instability of the zero solution of equation (5.3.2) is possible only if the network is frustrated. This result shows that frustration is essential for delay-induced instability, confirming the numerical observation in Marcus and Westervelt [1988,1989]. Example 5.3.9. Let W

(° =\-a \o

1 0 1

0 \ 1— α I , 0

)

(5.3.15)

where 0 < a < 1, be the connection matrix of a frustrated 3-neuron system. For general a, a(W) = {0, ± V 1 -2a}. When a = 0.5, a(W) = {0} and thus W has the Perron property for this value of a, and the zero solution of equation (5.3.2) is linearly asymptotically stable for all delays. When a = 0.25, σ (W) = {0, ±V2/2} and thus for all r, when β < Λ/2, the zero solution of equation (5.3.2) is linearly asymptotically stable, and when β > Λ/2, it is unstable. By contrast, when a = 0.75, a(W) = {0, ±ΪΛ/2/2), thus W does not have the Perron property, and delay-induced instability of the zero solution of equation (5.3.2) occurs for Β > Λ/2 at some value of r > 0. A network in which every cycle in the connection matrix is negative is called fully frustrated. The example below gives a fully frustrated network. Example 5.3.10. Let the signed digraph of the η χ η network connection matrix W consist of a single neuron (labeled 1) connected by negative 2-cycle to each of η — 1 other neurons (labeled 2, ..., n). The product of each 2-cycle, namely, wiktvki, lc = 2, . . . , η is negative and a(W) = {0, ±i Σλ=2\ w \k^k\ I}· This is a fully frustrated network and exhibits delay-induced instability for β > 1/ \ w\kw>k\ I· When the connection matrix W is normalized as Σ]=\ I wij I = 1 for all i = 1 , . . . , n, Belair, Campbell and van den Driessche [1996] obtained the following quantitative result for a network that is not frustrated: Theorem 5.3.11. Assume that W is normalized and is not frustrated. Then the zero solution of (5.3.2) is linearly asymptotically stable for all delays if β < 1, and is unstable for all delays if β > 1. Example 5.3.12. Theorem 5.3.11 can be applied to the all-excitatory connection matrix with Wij = \/{n — 1) for i φ j and wu = 0. In fact, Wu [1998] proved, for Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.4 Hopf bifurcation of periodic solutions

105

the nonlinear equation (5.3.2), that the zero solution is a global attractor when β < 1, whereas two nonzero asymptotically stable equilibria appear if β > 1.

5.4 Hopf bifurcation of periodic solutions The discussions in Section 5.3 show how delay-induced instability occurs in a network of neurons through the change of the signs of the real parts of some characteristic values of the linearization at a given equilibrium when the delay passes through a critical value. Typically, such a change of the signs of the real part of a characteristic value is related to the occurrence of a pair of purely imaginary characteristic values and, as we illustrate in this section, such a change leads to the occurrence of a branch of periodic solutions — the so-called Hopf bifurcation. For the sake of simplicity, we will first concentrate on the simple scalar equation x{t) = -x(t)

+ F(x(t - τ))

(5.4.1)

with the time delay τ > 0 as bifurcation parameter, where F e C 3 (M; R) and F(0) = 0. Such an equation can be regarded as the model for a single neuron with selffeedback. Alternatively, such an equation describes the synchronized activity of a group of neurons described by the additive model (5.3.1). While the signal function is usually assumed to be a smooth sigmoid which changes its concavity at zero, we emphasize that in many applications it is important not to assume that F"{0) = 0. This is specially useful when we consider the dynamics of the model equation HO = -y(0

+ f(y(t -τ)) + /

near an equilibrium y* such that _y* = f(y*) + I, where / is an input of constant value. A change of variable χ = y — y* leads to a scalar equation of (5.4.1) with F(£) = / ( £ + y*) - y* + I for ξ e R. It is then obvious that F"(0) = f"(y*) may be nonzero even if / is a sigmoid and even if / " ( 0 ) = 0. We first rescale the time t by t - » ί / r . This provides the normalization of the delay so that x(t) = —Tx(t) + rF(x(t

- 1)).

(5.4.2)

Let β = F'(0). Then the linearization at the equilibrium χ = 0 is x{t) = -rx(t)

+ xßxit

- 1)

(5.4.3)

and the associated characteristic equation is 0 = Α(ζ;τ,β)

= ζ + τ -rße'z.

(5.4.4)

As illustrated in last section, the analysis of the distribution of zeros of the transcendental function in (5.4.4) yields information about the stability of the equilibrium χ = 0. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5 Signal transmission delays

In this section, we consider the critical values of τ such that (5.4.4) has a pair of purely imaginary zeros. Let ζ = ico with ω > 0 and substitute this into (5.4.4). We have ΐω + τ — rßicoso) — i sin ω) = 0, from which it follows that I = β cos ω

(5.4.5)

ω = —τβ sin ω.

(5.4.6)

and Consequently, ω = a>k(ß) =

2kn + arccos 4 ß , 2(k + 1)7Γ — arccos ^

for β < — 1, for β > 1

(5.4.7)

and COk(ß) T = xk{ß) = ^ = =

for 1^1 > 1

(5.4.8)

for a nonnegative integer k (denoted by k e No). This proves the following Lemma 5.4.1. The characteristic equation (5.4.4) has a pair of purely imaginary conjugate complex solutions ζ = ±i ω with ω > Oifandonly if\ß\ > 1 and ω = a>k(ß) and τ = Tkiß) for some k € No. The functions ω* and r* are C1 -smooth and a)k satisfies wk € ((2k + |)7r, (2k + 2)π)

for k e No and β > 1,

(ok(ß) e ((2k + \)π, (2k + 1)tt)

fork e N0 and β < - 1 .

xk is decreasing in β > 1 and increasing in β < — 1 for each fixed k e No- Furthermore, 2π rk+l(ß) - Tk(ß) = for β € R \ [ - 1 , 1] andk € N 0 y/ß2 ~ 1 and lim r^(^) = 0,

lim rk(ß) = oo.

To apply the standard Hopf bifurcation theory, one needs to check the so-called "transversality condition" formulated in the next result. Lemma 5.4.2. Let \ß\ > 1 and fix k e No- Then there is an ε > 0 and a simple characteristic root z(t) = a(r) + ib(r) of (5.4.4) for |τ — xk(ß)\ < ε such that ζ is C 1 in τ, a(Tk(ß)) = 0, b(rk(ß)) = cok(ß) and £a(rk(ß)) > 0. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

107

5.4 Hopf bifurcation of periodic solutions Proof.

The existence of ε and z(r) follow from the implicit function theorem since ^Α(ζ\τ,β)\ z = i a ) k ( f i ) < T = 1 . k ( f i ) = 1 + Tße~z\z=ia,k(ß)yZ=Xk(ß) φ 0. Substituting ζ(τ) into (5.4.4) and differentiating with respect to r, we get z'(r) +

from which and follows that /

tkße~lü>k =

a'tik) = Re

ße~

1 -

ia>k + Xk

z i T )

k

z M

z ' ( T ) =

(here we write r* =

+ Tk)T

0,

τ^(β)

and ω^ =

ωι^β))

it

2

I 0)l· ( l + i c o

+ T ß e ~

= —ρ k

T

k

[(1 +

(Or.

^

Tk)2+

^ > 0.

ω\\

From the results in Lemmas 5.4.1 and 5.4.2 and using some detailed analysis, from XI.2 in Diekmann, van Gils, Verduyn Lunel and Walther [1995], we can have a complete picture about the distribution of zeros of (5.4.4) shown in Figure 5.4.1. Namely, one observes that the characteristic equation (5.4.4) has τ

Figure 5.4.1. Schematic picture of the distribution partition of the (β, r)-plane. Integer numbers in each region bounded by r = tkiß) shows the number of characteristic values with positive real parts.

(i) only solutions with negative real parts if r > 0 and β e [—1,1] or if ^ < —1 and 0 < r < το(β) (so the equilibrium χ = 0 is asymptotically stable); (ii) exactly 2k + 1 solutions with positive real parts if β > 1 and T k - i ( ß ) < τ < Tk(ß) for k > 1, and exactly one solution with positive real part if β > 1 and 0 < r < r0(/3); Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5 Signal transmission delays

(iii) exactly 2k solutions with positive real parts provided that Tk-i(ß) < τ < τ^(β) with β < — 1 and k e Ν (the set of positive integers). In other words, r^ (β) is the sequence of critical values of the delay where the "unstable dimension" of the equilibrium χ = 0 increases by 2. Lemmas 5.4.1 and 5.4.2 also provide the verification of the conditions required in the standard Hopf bifurcation of delay differential equations (see, Hale and Verduyn Lunel [1993], p. 332) and thus we have

Theorem 5.4.3. Let \ß\ > 1. Then for each k e No, α Hopf bifurcation for equation (5.4.1) occurs at Xk(ß)· In particular, there exists a sequence {τ"}^=1 such that τ" —> χkiß) as η οο and that for each τ = τη, equation (5.4.1) has a periodic solution Xk^nof the period pktTn suchthat pk^n -> = JUL— andxk,T"(t) —> v 0 uniformly for t —Ε as η —>• oo. It is desirable to determine the direction of the above Hopf bifurcation, i.e., to answer the question whether the bifurcating branch of periodic solutions exists locally for τ > τ k i ß ) (supercritical

bifurcation) or r < Xk(ß) (subcriticalbifurcation).

Also,

it is important to describe the stability of the bifurcating periodic solutions. A general approach to the solutions of the above questions is to compute and analyze the reduced system on the center manifold associated with the conjugate pair of characteristic values ±ia)k(ß), an alternative approach is that based on the method of LyapunovSchmidt developed in Stech [1985]. Two general algorithms are available in the literature to compute the center manifold. One is provided by Belair and Campbell with a Maple implementation and another is provided in Faria and Magalhäes [ 1995] to calculate the reduced system on the center manifold in normal forms. Giannakopoulos and Zapp [1999] applied the algorithm of Faria and Maghalhäes and derived the following normal form equation, in polar coordinates (ρ, ξ), for the reduced system on the center manifold Ρ = (τ - τ*)£α(τ*)ρ + Kp1 + 0((τ - η)2ρ + \(ρ, τ - τ*)|4), ξ = -ωΐί + 0(\(τ -Tk,p)\)

Κ

- -'

with _ F'"{0)τ 1 + β2τ 2 β(1+2τ + β2τ2) (F"(0))2t β2τ(ί\β2 + 6β-2) + (2β3 + 13^2 + 4 / 3 - 4 ) 2 β2(5β + 4)(/ϊ — 1)(1 +2τ + β2τ2) Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

ΐ Ρ <

109

5.4 Hopf bifurcation of periodic solutions and

Κ =

F"'( 0)τ

1 + β2τ

β(1 + 2r +

2

β2τ2)

(F"( 0 ) ) 2 r β2τ(1 ϊβ3 + 35β2 + 24β - 6) + (-2β4

+ 1\β3+ Α3β2 + 24β - 12)

β2{β - \){5β2 + \5β + 12)(1 +2τ+

β2τ2)

ή β > 1. Since ^ α ( τ ^ ) > 0, the sign of the factor Κ determines the direction and (orbitally asymptotical) stability of the bifurcating periodic solutions. Note also that ±K

< 0

if and only if

± F"'(0)F'(0) <

±(F"(Q))2Ck{ß)

with Cjfc(^) =

\\β2+6β-2 ι ? (ß+\)2 (5ß+A)(ß — \) (5ß+4,)(\+ß2Zk) 11/?3+35£2+24/?-6 (ß+lKß2-3) ? 03-l)(5/32+15/?+12) (5/52+15/5+12)(l+^2r,0

fnrfl^-1 " , „ , ^

We then get

Theorem 5.4.4. Fix

keK

(i) If F"'(0)F'(0) > (F"(0))2Ck(ß), then the Hopf bifurcation is subcritical the bifurcating periodic solutions are unstable.

and

(ii) If F"'(0)F'(0) < (F"(0))2Ck(ß), then the Hopf bifurcation is supercritical. The bifurcating periodic solutions are stable if β < — 1 andk — 0, and unstable if β < -\andk>\ or if β > 1. From the definition of Ck(ß), we can easily verify that ( C I ) if β < — 1 or β > 7 3 , then the sequence {Ck(ß))keN0 strictly increasing;

is positive, bounded and

(C2) if 1 < β < λ/3, the sequence {Ck(ß))k€N 0 is positive, bounded and strictly decreasing; (C3) if β = 7 3 , then the sequence {Ck(ß)}keN0

is a positive constant sequence.

Consequently, we have the following

Corollary 5.4.5. Let [ F " ( 0 ) ] 2 + [F'"(0)] 2 φ 0. (i) If F'" (0) F' (0) < 0 then all Hopf bifurcations are

supercritical;

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5 Signal transmission delays

(ii) If F"{0) = 0 and F w (0)F'(0) > 0 then all Hopf bifurcations are subcritical; (iii) IfF"(0)

φ 0 and F"'(0)F'(0)

> 0 then

(a) for β e (—oo, —1) U (%/3, oo), either all Hopf bifurcations are supercritical, or all Hopf bifurcations are subcritical, or there exists a unique kc G Ν such that all Hopf bifurcations occurring at τ = τ* (β) are subcritical for k < kc and supercritical for k > kc; (b) for β e (1, >/3), either all Hopf bifurcations are supercritical, or all Hopf bifurcations are subcritical, or there exists a unique kc e Ν such that all Hopf bifurcations occurring at τ = Xk(ß) are subcritical for k > kc and supercritical for k < kc. Figure 5.4.2 lists all possible bifurcation scenarios.

Figure 5.4.2. All possible bifurcation scenarios if [F"(0)] 2 + [F"'{Ο)]2 φ 0.

So far the analysis in Sections 5.3 and 5.4 is local in nature: the asymptotic stability of the equilibrium and the existence of a local Hopf bifurcation come from the study of the characteristic values of the linearization of the network at considered equilibrium and the conclusion about the behaviors holds true only for trajectories in a small neighborhood of the equilibrium. In particular, the local Hopf bifurcation theorem (Theorem 5.4.3) gives the existence of periodic solutions near (0, τ^β)) in the space C([—1, 0]; R) χ R. This is a local result in the sense that (i) the amplitude of the periodic solution is small and (ii) the existence of a periodic solution is guaranteed only for τ near Tk(ß). The issue of global existence (for all r > r k ( ß ) ) of largeamplitude periodic solutions is related to the study of the global continuation which Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.4 Hopf bifurcation of periodic solutions

111

we briefly describe below. Let S = {(φ, τ, ρ) g C([—1,0]; Μ) χ R+ χ R+; χ is a non-constant /^-periodic solution of (5.4.2) with XQ = φ) 5 = the closure of 5 in C ( [ - l , 0]; l ) x R + x R+. For each k € No, let Sk denote the maximal connected component of S in C ( [ - l , 0]; 1 ) χ l + χ R + which contains (0, rk(ß), Ιπ/y/ß2 - 1). Assuming F e C 2 (R, R) with |F'(0)| > 1 and assuming that F(x) = χ for some χ e R implies F'ix) Φ 1, Giannakopoulos and Zapp [1998] applied the global Hopf bifurcation theorem in Erbe, Geba, Krawcewicz and Wu [1992] based on the S'-equivariant degree theory to obtain the following: Theorem 5.4.6. For each k e No, the maximal connected component Sk is unbounded. Moreover, for each (φ, τ, β) € Sjt with φ Φ 0 there exists a non-constant p-periodic solution Χ of (5.4.1) with XQ = φ and ρ > 2r if β < —1 and k = 0, Ρ € (ifc+J' tf ß < — 1 andk > 1 and ρ € ^jtTT· it+j) tf ß > 1 and k € N. To ensure the unboundedness from above of the r-component of S* (this is equivalent to ensuring the existence of a non-constant periodic solution for each τ > Tk(ß)), we need further conditions to guarantee certain a-priori bounds for the super-norm norm of all possible periodic solutions. This can be achieved if we assume that there exists a compact interval / C R with 0 e int(/) and F(I) c 7 6

(5.4.10)

and assume the negative feedback condition xF(x)<0

for * e / \ {0}.

(5.4.11)

More precisely, one has Corollary 5.4.7. Assume (5.4.10) holds. Then (i) if β < —1, then for each k € Ν and τ > Tk(ß) there exists φ € C([— 1,0]; I)

(ii) if β > 1, then for each k e No and τ > Xk(fi) there exists φ 6 C([— 1, 0]; I)

(iii) if β < —1 and, in addition, (5.4.11) holds, then for each τ > το(β) there exists ψ e C([— 1, 0]; /) and ρ > 2r such that (ψ, τ, ρ) e Sk. 6

Note that a commonly used signal function which is bounded satisfies this condition.

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5 Signal transmission delays

We now provide several examples to illustrate the above local and global results. All figures below are schematic. For the bifurcation diagrams based on a numerical continuation method for delay differential equations where the periodic solutions are approximated by Fourier polynomials, see Giannakopoulos and Zapp [1998], Example 5.4.8. We consider the sigmoid nonlinearity 1 F(x) = Fi(x) := 2 1 + exp(-4x)

3 4'

Clearly, — | < F(x) < | for all JC e R, F is strictly increasing and has exactly one turning point at 0 (F"(0) = 0, F'"(0) < 0). Moreover, F(0) = 0, β = F'{0) = 2 > 1 and F has three fixed points χ = 0, χ « 0.644, —Jc. By Theorem 5.4.4, we conclude that a supercritical Hopf bifurcation occurs at (0, τ*(λ)) for each k G NOCorollary 5.4.7 shows that for eachfixed& e Noandforallr > τ* (λ) there is a periodic solution χ belonging to S* (in the sense explained in Corollary 5.4.7). Furthermore, we know that the period ρ of χ satisfies ρ e ^FTT' it+j)· Figures 5.4.3 and 5.4.4 show that along the branches |x| = sup feR |x(/)| is monotonically increasing in τ, but ρ/τ does not change much.

Figure 5.4.3. Schematic picture of branches of periodic solutions in the (r, |x|)plane. All bifurcations are supercritical and the corresponding solutions are bounded by x.

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5.4 Hopf bifurcation of periodic solutions

113

ρ/τ

Figure 5.4.4. Schematic picture of branches of periodic solutions in the τ)-plane.

Example 5.4.9. We now use the nonlinearity F(x) = —F\(x) with F\ as in the first example, χ = 0 is the only fixed point of F, and/J = F'(0) = — | < — 1, F"{0) = 0, F"'{0) > 0. Therefore, a supercritical Hopf bifurcation occurs at τ = τ^(λ), k € No- Since F satisfies (5.4.10) and (5.4.11) with I = [—x, x], where χ is defined in Example 5.4.8, we obtain that the τ-component of Sk is unbounded from above. The period ρ of the solution χ belonging to Sk is larger than 2 τ (the corresponding solution is said to be slowly oscillating) for k = 0 and ρ € f ° r k > \. Numerical simulations indicate that the branches are monotonically increasing in r and for every τ > το (λ) there is a unique slowly oscillating periodic solution. Example 5.4.10. We now provide an example which demonstrates the change of the direction of Hopf bifurcations. Let F(x) = - 5 7 5 arctan(x + loVS) + 575arctan(10\/5). F Then F(0) = 0, β := F'(0) = < - 1 , F"{0) = = 14 We have -125751501 · 9716 « C0(ß) < Ci(ß) ^ 1.49952. We can show that a Hopf bifurcation occurs at τ = for each k e No- The first Hopf bifurcation at r = το(β) is subcritical, all other Hopf bifurcations are supercritical (see Figure 5.4.5).

Example 5.4.11. We now consider an example where the function F does not satisfy condition (5.4.10). Consider F(x) = —15 sinh(;c). Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5 Signal transmission delays

Figure 5.4.5. A subcritical Hopf bifurcation occurs at τ = r^iß) = 4.66712 and a supercritical Hopf bifurcation occurs at τ = τ\ ( β ) = 15.82280.

It holds that F(0) = 0, β := F'(0) = - 1 5 , F"(0) = 0 and F'"(0) = - 1 5 . F is strictly decreasing but does not satisfy condition (5.4.10). χ = 0 is the only fixed point of F. Using Theorem 5.4.4 we get a subcritical Hopf bifurcation at each r = Tkiß). From Theorem 5.4.6 we know that the branches of periodic solutions Sk bifurcating from (χ = 0, r = τ^(β)) are unbounded but we do not know whether the component of Sk is unbounded. Numerically simulations show that the branches are bounded in τ. The numerically computed ρ-periodic solution jt*(f; r) belonging to Sk exists only forO < τ < Tic(ß) and o o a s r —> 0. Furthermore, for the period ρ of xo(t\ τ) it was observed that §r —>· 4 as r —> 0. Example 5.4.12. The last example deals with the case in which F does not satisfy the negative feedback condition (5.4.11). We consider F(x) = 2

*+ 1 — 2. 1 + (χ + l) 1 0

This equation was introduced by Mackey and Glass [1977] as a mathematical model for the generation of white blood cells. F is a bounded function with F(0) = 0, β = F'(0) = - 4 < - 1 , F " ( 0 ) = —5 and F"'(0) = 255. Using Theorem 5.4.4 we get a supercritical Hopf bifurcation at (x = 0, f = Xk(ß)) for every k e No. From Corollary 5.4.7 we know that for k > 1 the τ-component of Sk is unbounded from above. Since (5.4.11) is not satisfied, we can not investigate So by applying Theorem 5.3.7 directly. Numerical simulations show that the branch So of slowly oscillating periodic solutions undergoes a cascade of period doubling bifurcations. Moreover, numerical calculations indicate the existence of chaotic attractors for sufficiently large delay τ. We note that the general procedure described here for the existence of a local bifurcation and its global continuation of a scalar equation can, in principle, be extended to networks of arbitrarily large number of neurons, but the practical implementation Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

115

of this procedure is far from trivial and it is quite useful to use the algebraic manipulation language Maple to perform the calculations. In Belair, Campbell and van der Driessche [1996], this was used to perform a center manifold reduction for a frustrated network of 3 neurons given by 3 « i ( 0 = -Ui(t)

+ ^ W i j f ( u j ( t - τ)),

1 < i < 3

7=1

with /

0 10\ w = (Wij) = - 1 . 2 5 0 1 \ 1.25 1 0 / and f(u) = β tanh(«). Here, a(W) = {1, —0.5 ± /} and delay-induced instability occurs for 1 > β > giving rise to a Hopf bifurcation. Specially, for β = 0.96, this occurs at τ = 4.265 where the linearization has a pair of purely imaginary characteristic values ±ϊω with ω = 0.3899. Thus at this point, the solutions to the system of 3 delay differential equations may be approximated by the flow on a two-dimensional center manifold governed by a system of two ordinary differential equations whose normal form in polar coordinates is ρ = -0.541 ρ3, θ = 0.3899. This shows that the bifurcation is supercritical and the delay-induced stable periodic solutions occur for r > 4.265. Finally, we mention that a general symmetric Hopf bifurcation theory was developed and applied to obtain the existence of periodic solutions with some spatialtemporal patterns for networks of neurons with delayed feedback and symmetric connection topology in Wu [1998]. See also Huang and Wu [2001] and Wu, Faria and Huang [1999] for various waves induced by signal delays.

5.5 A network of two neurons: McCulloch-Pitts nonlinearity In the subsequent sections, we consider a system of two neurons connected to each other. Denoting the activation of the neurons by xi and X2, their decay rates by γ\ and Y2, their connection weights by w\ and W2, the signal delays related to each connection by ri and T2 and the constant inputs received by each neuron by I\ and I2, respectively (shown in Figure 5.5.1). Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

116

5 Signal transmission delays

h

h (u>2, n)

Figure 5.5.1. A network of two neurons with inputs ( / ι , I 2 ), connection weights (w\,w2) and signal delays (τι, τ2).

Let / denote the signal function, then we have the following system of delay differential equations *i(0

= ~Y\x\it)

+ I\ + w\f(x2(t

-

τι)),

* 2 ( 0 = ~Y2Xi(t)

+ h + w2f(xi(t

-

r2)).

Two-neuron networks have played an important role in the analysis of delayinduced changes in the dynamics of neural networks. Despite the low number of neurons in the system, in many instances, two-neuron networks with delay display the same behavior as larger networks and many techniques developed to deal with twoneuron networks can carry over to large size networks such as those used in practical applications. Therefore, a two-neuron net can be used as prototypes to improve our understanding of the influence of delay. In this section, we introduce the work of Huang and Wu [1999b, 2000] where a two-neuron artificial network with delayed feedback from one neuron to another is considered. Assuming the signal transmission is of a digital (McCulloch-Pitts) nature, the model then describes a combination of analog and digital signal processing in the network and takes the form of a system of delay differential equations with discontinuous nonlinearity. In what follows, we present a detailed analysis of the dynamics of the inhibitory network starting from non-oscillatory states, and show that the trajectories are either synchronized and ultimately periodic or converge to a desynchronized equilibrium. We will also show that large threshold is one of the main sources for the synchronization of the network considered. More precisely, we consider the following model χ = -μχ

+ f(y(t

-

Ti)),

y = -py

+ /(*('

-

τ2)),

(5.5.1)

with μ > 0, τι > 0 and τ2 > 0. We point out that in the case where / is piecewise Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

117

linear, i.e., /(£) = + 1| - \ξ - 1|) for ξ e R, (5.5.1) is the simplest form of the cellular neural networks introduced by Chua and Yang [1988] which have found interesting applications in, for example, image processing of moving objects and which can be implemented by a delayed cloning-template (see Roska and Chua [1992]). We consider the case where the signal transmission is of digital nature: a neuron is either fully active or completely inactive. Thus the signal transmission is of McCulloch-Pitts type and we have if ξ > σ, if £ < σ

a b

(5.5.2)

for a, b, σ e R and α Φ b. Therefore, the model describes a combination of analog and digital signal processing. We only consider the case where b > a (The case where b < a can be considered similarly). To simplify our presentation, let us first rescale the variables by r

-

ßt,

τ =

μ

τ

- ψ .

('*) = & > ( ' + * _ 2μ _ b+a ° ~ b-a" b-a *

f */£\ _ J

2 f (b-a y , b+a\ _ b+a Κζ) — b_a J y 2μ $ 2μ ) b-a '

f )

b+a b-a '

and then drop the * to get χ = - x + f(y(t -

τ)),

y = - y + f(x(t -

τ))

(5.5.3)

with -1

m

j l

if ξ > σ , if I < σ. if

(5.5.4)

The discontinuity of / makes it difficult to apply directly the theory of dynamical systems to model equation (5.5.3). But, the simple form of (5.5.3) and (5.5.4) enables us to carry out a direct elementary analysis of the dynamics of the network due to its obvious connection with the following systems of linear nonhomogeneous ordinary differential equations χ = —χ — 1, (5.5.5)

{

y =

- y - l

(5.5.6) (5.5.7) (5.5.8) As usual, we use X = C([—r, 0]; R ) as the phase space. Note that for each given initial value Φ = (ψ, ψ ) Τ € Χ, one can solve system (5.5.3) on intervals [0, r], Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5 Signal transmission delays

[τ, 2τ], . . . successively to obtain a unique mapping ( χ φ ,

: [—τ, oo) —• R 2

such that χφ |[_τ ο]= φ, |[_ r ,o]= ψ, {χΦ, y*1*)7 is continuous for all t > 0, differentiable and satisfies (5.5.3) for almost every t > 0. This gives a unique solution of (5.5.3) defined for all t > —τ. Our goal is to determine the limiting behaviors of ( χ φ , >»Φ)Γ as t —>- oo for any Φ e X, but here we shall concentrate on the case where φ — σ and ψ — σ have no sign change on [—r, 0] (initial values are either above or below the normalized threshold σ). Namely, we consider Φ € U Xp~ U χ - ' + υ χ - · - = Χ σ given by χ*'* = {Φ € χ- Φ = (φ, f)T,
C± and f e C±)

with ,,+ _ Ι ' σ ~~ y '

φ : [—τ, 0] —[σ, oo) is continuous and
and C

a

=

φ : [—r, 0] —• (—oo, σ] is continuous and
Clearly, all constant initial values (except σ) are contained in Χσ. Our analysis shows that once we restrict Φ to Χσ, the behaviors of (χΦ(ί), }>φ(ΐ))Τ as t —>· oo are completely determined by the value Op(0), ψ(0)) τ and the size of the threshold σ. More precisely, we have Theorem 5.5.1. Assume that |σ| < 1, Φ = (φ, ψ)τ f(0). Then χΦ(ί) = y*(t)forallt > 0 αηάχφ(ί) τ + ln[l + τ Φe (respectively, Φ e X~~), where q : Μ minimal period ω = In ( j ^ e T — l) + In —

e Χ£'+ U and φ(0) = = γφ(ί) = q(t) for all t > + ln[l - ip(0)] - ln(l -a))if R is a periodic function of the l).

Clearly, if Φ = (φ, ψ)τ e X is synchronous {i.e., φ = ψ) then the solution (χΦ, y 0 ) 7 : [—τ, oo) —> R 2 is synchronized, that is, χφ(ί) = y 0, due to the uniqueness of the initial value problem of (5.5.3). The above result shows that the solution (* φ , of (5.5.3) is synchronized even if the initial value Φ is asynchronous but φ(ϋ) = ψ φ ) and (φ, ψ ) τ G X p + U • Obviously, a synchronized solution of (5.5.3) is characterized by the scalar equation z = -z + f(z(t - r))

(5.5.9)

and Theorem 5.5.1 shows that a solution of (5.5.9) with initial value in C^ is eventually periodic with the minimal period ω = In — l) + In — l) if |σ| < 1. It is interesting to note that the periodic function q and its minimal period ω are both independent of the choice of the initial value of Φ 6 X^^ with φ(0) = ψ(0). Moreover, note that ^ —> 1 as τ —> oo. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

119

Theorem5.5.2. Let \σ\ < 1 and Φ = (φ,ψ)τ G Χσ. Then (χΦ(ί), γφ(ί))Τ ( - 1 , L)R if either ψ(0) > σ and we show by direct calculations that the solution (* φ , is convergent to (—1, l ) r as t —> oo if (^>(0), ψΦ))τ G Rj and Φ = (φ, ψ)τ G Χσ'+', region Rn consists of (<^(0), ψ(0))τ with either φφ) > σ, ψφ) > σ and ψφ) > + ex - 1 or φφ) < σ, ψφ) < σ and 0, (χΦ(ΐ*), }>φ(ί*))Γ G Rj and ( * Φ ( Ί * + · ) , / » ( ' * + 0 ) Γ l [ - r , 0 ] G if (φφ),ψφ))τ G Rn and Φ = τ (φ,ψ) e U region R m consists of those above the diagonal but not in Ri U Rn, and we also show that there exists t* > 0 so that (.χ φ (ί*), }>Φ0*))Γ G Rj and (χ φ (ί* + ·), / V + ·)) Γ Ι[-τ,0]6 if G Rm and Φ = {ψ, ψ ) τ G X p + U In summary, the solution ( χ φ , >' Φ ) Γ of (5.5.3) starting from any point above the diagonal must enter the region Rj eventually and (* φ (ί* + ·), yΦ{t* + ·)) Ι[-τ,θ]£ forsomei* > 0, and therefore must be convergent to (—1, 1)^ as t —> oo. The region below the diagonal in the ( 1 is completely different from that with \σ\ < 1. In particular, we notice the role of synchronization of large threshold. Theorem 5 . 5 . 3 . Let \σ\ > 1, and let Φ = (φ, ψ)τ G Χσ. Then, as t —oo, (χΦ(ί), γφ(ί))Τ (1, l)r ifa > 1, and (χφ{ί), γφ(ί)) ( - 1 , - 1 ) Γ if α < - 1 . In the case where \σ\ = 1, we notice that the dynamics of the network undergoes change when the ratio (if σ = 1) or fyojli (if ο = —1) passes certain critical values. Theorem 5 . 5 . 4 . Let σ = (i) (**(r), y*(0)

Τ

1

and Φ = (φ, ψ)τ e Χσ. Then (1,

l ) r if any one of the following cases occur: (a) Φ G X-'~; (b) Φ G X+<+ and e~τ < < e*; (c) Φ G X~'+ and ψφ) = 1; (d) Φ G X+- αηάφ{0) = 1; Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

120

5 Signal transmission delays

(ii) (χ φ (ί), }> φ (ί)) Τ aand na

( - 1 , 1 ) Γ if either Φ G X~'+ and ψφ) φ 1 or Φ G X+'+

ψ(0)+1

T ?(0)+l > ee '·

(iii) (* φ (ί), (1, - 1 ) Γ if either Φ e X+~ αηάφ{ 0) φ 1 or Φ G X+' + »(Q)+i < e e-r aand na p(0)+l ^ · Theorem 5.5.5. Let σ = — 1 and Φ = (ψ, ·ψ)τ G Χσ. Then vT

(i) (x*(t),y*(t)) 0 or Φ

G

χ-~

^

( - 1 , - 1 ) τ ^ ΐ ^ Φ e X+' + and[0(O) + l][r/r(O) + l] φ

and e~z <

< ex;

(ii) (χφ(ί), >,Φ(θ)Γ -* (—1, l)r if any one of the following cases occurs: (a) Φ € X--+; (b) Φ e

and

(c) Φ G Xp+and (iii) (λ:φ(0, y*(0)T (a) Φ

G

<

;

φ(0) = - 1 ;

~~* (1- — 1)Γ if any one of the following occurs:

X+'~;

(b) Φ G X-'~ and

>

^

(c) Φ G X+'+ and f(0) = 1. Regarding the threshold σ as a bifurcation parameter, we show in the above results that as σ moves from the interior of interval (—1,1) to its exterior, the dynamics of model equation undergoes transition from either ultimate periodicity and synchronization or convergence to asynchronous equilibria (1, — 1) T and (— 1,1) T to synchronization and convergence to synchronous equilibria (1, l ) r and (—1, —l) r . These results seem to indicate that large threshold is the main source for synchronization in the network discussed. We now provide the proofs of the above results. We first note that, by solving (5.5.3) on intervals [0, τ], [r, 2 r ] , . . . successively for each Φ = (φ, ψ)τ G Χσ, the unique solution ( χ φ , >>φ)7 : [—τ, σο) —> Μ2 satisfies (i) ( χ φ , )>Φ)Γ : [—τ, οο) -»• Μ2 is continuous; (ii) χΦ(ί) and γΦ(ί) are differentiate for all t > 0 such that [x(i - r ) - a][y(t - r) - σ] φ 0;

(iii) (χφ(ί), γφ(ί))Τ

satisfies (5.5.3) for all t > 0 with [x(t - τ) - a][y(t - τ ) - σ ] φ 0 .

Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

121

Note also that, by the variation-of-constants formula, on the interval [/*, t], the solution of the equation ζ = -ζ

—1

is given by z{t)

= e'^Hzit*)

+ 1] - 1

or *(*> + 1 z(t*) + 1

e-(t-n

=

and the solution of the equation z = - z + 1

is given by Z(t)

= e-^Hzit*)

or

z(t)-]_

1] + 1

e_(t_r)

=

- 1

Z(t*)

Consequently, each ( * φ , >>φ) following straight lines:

-

: [0, oo) —» R 2 moves alternately along one of the

x(t)

+ 1 _

x(t*)

+ 1

y(t)

+ 1 "

y(t*)

+ 1'

xit)

+ 1 _

x{t*)

+ 1

y(t)

-

1 ~

y(t*)

-

1'

x{t)

-

1 _

x(t*)

-

1

y(0

-

ι ~ y(t*)

-

x(t)

-

1 _

x(t*)

-

y(t)

+ 1 ~

y(t*)

+ 1'

l'

1

In other words, a trajectory is a broken line. As an illustrative example, we consider Φ = (φ, f ) T e X + ' + with |σ| < 1 and
=

ψ%+\

until

= ( χ Φ 0 ) , y
'ι +

< φ(θ)+ι

moves along the straight line

<

=

where t\ is the first zero of (jc — σ ) 0 - σ) in [0, oo),

then it moves along the line

until t2 + τ, where t2 is the second

=

zero of (x — a)(y — σ) in [0, oo), and then it moves along the line until t3 + r, where t^ is the third zero of (λ — σ)(γ towards ( - 1 , l ) r along the line

=

— σ) in [0, oo), and then it moves

=

Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

122

5 Signal transmission delays

5.5.1. We only consider the case where Φ — {φ,ψ)τ e The case where Φ = (φ, ψ)τ € can be dealt with analogously. Using equation (5.5.3), we can easily obtain that χΦ(ί) = γ φ ( ΐ ) for all t > 0. Therefore, it suffices to show that the solution χ of the equation Proof

of Theorem

χ = - x + f{xit -

τ))

(5.5.10)

with the initial condition χ |[_ r q]= φ € C+ is eventually periodic with the minimal period In { f e e * - l) + In ( j ^ e * - l). Let t\ be the first nonnegative zero of χ — σ. Then for t e (0, t\ + τ), except at most finitely many points, we have χ = —χ —

1,

from which and the continuity of the solution it follows that x(t)

= e-'[
t e [ 0 , t\ +

+ l ] - l ,

r]

and, in particular, Λ(ί0

=e-,1[^(0) + 1 ] - 1 = σ .

This implies t\ =1η[^(0) + 1 ] - 1 η ( 1 + σ ) and x(

h

+ τ) = e~(tl+T)(
+ 1) -

+ (1 + a)e~T

1 = -1

< σ.

Also χίι+τ(θ)

:= x(ti

+ τ + θ) = e-^+z+eH
= (1 +σ)
(Γ+
+ 1) -

1

- 1< σ

for θ e (—r, 0], which means Jt i l + r e C~. To move to the next step where we explicitly construct a solution of (5.5.10) beyond [0, t\ + τ], we start with a new initial value φ* = x f l + r . Let ?2 be the first zero after t\ of χ — a. Then t2 > h + τ and on (t\ + τ, ti + t ) , we have χ

=

—χ

+ 1

and hence x(t)

= e~^-T)[x(ti

+ τ) -

1] + 1

= [ίί>(0) + l ] ( l — γ-j·—er\e~' V 1+ σ /

+

1

for t e [ti + τ, t2 + τ]. In particular, X(t2)

= (φ(0)

+

l)(l - —

+

Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

1= σ

123

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

implies t2 = ln(p(0) + 1) + In

- l ) - ln(l - or).

Also, Λ

χ,2+τ(θ)

= x(t2

+ τ + θ) = =

(φφ)

+ l)(l -

Y^e

1 - (1 -

T

)e~

( t 2 + T + e )

+ 1

> σ

for θ € (—r, 0], which means χ(2+τ € C+. Repeating the above arguments inductively, we can show that there exists a sequence {i„} (n = 1, 2 , . . . ) such that h = ln(p(0) + 1) - ln(l + σ), t2 = ln(?(0) + 1) + I n t2n-l

~

= t2n-3 + In

t2n = tln-2 x{tn)

=

x(hn-\

~

~

+ In ( j ^ e *

ln

+

ln(1

( f h ^ ~

l ) + In ( j ^ e *

-

~

' n

' '

n

^ -

2

> 2

'

σ, + r ) = - 1 + (1 +

σ)β~τ,

-a)e~T,

x(t2n

+ T) = 1 - (1

x(t)

= (
X(t)

= g-U-^-i-*\x{t2n-\ + τ) -

-

1,

ί€[0,ίι+τ],

= - ( , ( 0 ) + 1)

1] + 1

- ι ) " " 1 e - ' + 1,

- l)" t

x(t)

σ)

~

= e-^"'T\x(t2n

+ Τ) + 1] -

€

1 + τ, ί 2η + τ],

1

t € [t2n + τ, t2n+1

+ τ].

See Figure 5.5.2. Consequently, we can see that χ(t) is periodic for t > t\ + τ, and that its minimal period is ω =

tln+\

-

tln-\

+h

-1'2"-1+ln ( ΐ Τ Τ ' -

0

Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

,2

- 0] " "-'

124

5 Signal transmission delays

Figure 5.5.2. The solution (5.5.3) with |σ| < 1 and the initial value (φ, ψ)τ Xp+ and
€

This completes the proof. Proof of Theorem 5.5.2. We only give the proof for the case of ψ(0) >
V·0·11'

which imply that χτ (θ) = χ(τ + Θ) < σ and yT(9) = y{t +Θ) > σ for θ G (—τ, 0), and so χτ G C~ and yT G C+. Repeating this argument on [τ, 2τ], [2τ, 3τ], . . . , consecutively, we obtain that xt G C~ and yt G C+ for all t > 0. Therefore, (x(t), y(t))T satisfies (5.5.6) for almost all t > 0, from which it follows that (5.5.11) holds for all t G [0, oo), and hence (x(t), y(t))T -> (— 1, l ) r as t oo. Case 2. We now consider the case where φ, ψ G C+ and ψ(0) > φ(0)βτ +eT — 1. The idea is simple, namely, we show that if t\ is the first zero of (χ — σ)(γ — σ) in [0, oo) then jc il+T G C~ and y / l + T G C+, then the conclusion follows from Case 1. By the initial condition and (5.5.4), (x(t), y(t))T must satisfy system (5.5.5) for t G [0, τ] except at most finitely many t. It follows that for t G [0, τ] i x ( 0 = C^(0) + I)«"' - 1, \y(t) = 0) + 1 ) β - ' - 1 . Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

(5.5.12)

125

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

Let t\ be the first zero of (χ — a)(y — σ) in [0, oo). Then (x(t), y(t))T satisfies system (5.5.5) for t e [0, t\ + τ] except at most finitely many t, and so expression (5.5.12) holds for t e [0, t\ + τ]. On the other hand, = [(φ(0) + 1)β~ί-1-σ][(ψ(0)

(x(t)-a)(y(t)-a)

+ \)β-(-l-σ]

= 0 (5.5.13)

implies t = ln(^(0) + 1) - ln(l + σ)

or

t = ln(^(0) + 1) - ln(l + σ).

In view of ^(0) > 1η(φ(0) + 1 ) + r. Therefore, h = ln(^>(0) + 1) - ln(l +σ)

and

h+τ

< ln(^(0) + 1) - ln(l + σ).

This, together with (5.5.12), implies that *(ίΐ + τ) = - 1 + (1 + σ)β~τ Φ(0) + 1 Ψ y(ti + r) = -1 + χ,ι+τ(θ)

< σ,

; ' (1 + σ)e φ(0) + 1

= χ(ΐχ +τ

> σ,

+ θ) = (y>(0) + 1)β~(ίι+τ+θ)

= -1 + (1 + σ)β~(τ+θ)

< σ

- 1

for θ € (—r, 0]

and yn+ΑΘ)

= y(n +τ+θ)

= ( f ( 0 ) + l)e~(ti+T+9) θ

> - 1 + (1 +σ)β~

> σ

- 1

for 0 e [—τ, 0).

This shows that xt] +T e C~ andy^+r € C+. So, by Case 1, we have (x(t), (—1,1) T as t oo.

y(0)T

Case 3. We next consider the case where φ, ψ e C~ and^(0) < ψ(0)ετ —eT +1. For this case, the proof is similar to that of Case 2, and thus is omitted. Case 4. We now consider the case where φ, ψ e Cj~ and φ(0) < ψ(0) < τ

β φ(0) + eT - 1. Let

_

+ 1 ^(0) + 1 " T Then
1 < β < eT. Using a similar argument as that in Case 2, we can show that (jc(r), y(t))T satisfies (5.5.5) for t € [0, t\ + τ] except at most finitely many t, where t\ = ln[^(0) + 1] — ln(l + o), Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

126

5 Signal transmission delays

and that (5.5.12) holds for all t e [0, t\ + r]. This, together with the fact that 1 < β < eT, implies that + τ) = - 1 + (1 + σ )e~T < σ, and

y(ti + t ) = - 1 + β (I + a)e~T < σ,

y(fl) = (f(0) + l)e~tl - 1 = - 1 + 0 ( 1 + o r ) > σ.

Therefore, there exists t2 e (ii, t\ + τ) such that y(t2) = σ. Clearly, t2 is the second zero of — a)(y — σ) in [0, oo). From (5.5.13) we can get t2 = ln(^(0) + 1) - ln(l + σ) = ij + ln/3. By(5.5.12), wehavex(i —τ) < σ andy(t — τ) > σίοτί e (ti+T,t2 + r). Itfollows from (5.5.3) and (5.5.4) that (x (t), y(t))T satisfies system (5.5.6) fori € (t\+x, t2+r). Therefore, for t e [t\ + τ, f2 + t], we have x(t) = e-^-tl~T)[x(ti y(t) = e-C-'i-^lyOi

+ r) + 1] - 1 = (1 + a)e~^-tl)

- 1, T

+ r) - 1] + 1 = [(1 + σ)β - 2e ]e~+

1. (5.5.14)

Hence, 1+ σ x(t2 + r) = -l + —— e~T ß

and

y(t2 + τ) = 1

2eT-(l ß

+σ)β -e

T

.

Note that t2 + τ < t\ + 2τ, and that for t e (t2 + τ, t\ + 2r) we have x(t - τ) = (φ(0) + l)e~ ( t ~ T ) - 1 < (
y(t - r) = (f(0) +

- 1 < (^(0) + l)e~'

< σ,

- 1 = σ.

So (x(t), y(t))T must satisfy system (5.5.7) for t e (t2 + r, t\ + 2r). It follows that for t e [t2 + τ, t\ + 2r] we have x(t) = e- ( t - t 2 ~ T ) [x(t 2 + τ ) - 1 ] + 1 = [ 1 + σ - 20e r ]
+ τ) - 1] + 1 = [(1 + σ)β - 2eT]e-^-^

+ 1. (5.5.15) Let t3 be the third zero of (* — σ)(γ — σ) in [0, oo), i.e., the first zero after t2 of (χ — σ)(γ — σ). Then (x{t), y(t))T satisfies system (5.5.7) for t e (t2 + r, /3 + r), and (5.5.15) holds for alii 6 [ί2 + τ,ί3 + τ], On the other hand, from (5.5.14), (5.5.15) and the fact β > 1, by solving the equation (x(t) — a)(y(t) — σ) = 0, we can get t3 = ti+ ln(2eT - (1 + σ)β) - ln(l - σ). This, together with (5.5.15), implies that x(t3 + τ) = 1 - l ßT f 2e - (1 + σ)β

- σ)e~T

and

y(t3 + r ) = 1 - (1 -

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σ)β~τ.

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

127

If < β < er, then χ + τ) < σ, moreover, from (5.5.14) and (5.5.15), it is easy to see that χί3+τ(θ) = x(ts + r + θ) < σ and y f3+T (0) = y(t$ + τ + θ) > σ for θ € (—r, 0). This shows that ·χ ί3+τ e C~ and y, 3 + r g C+. So, by Case 1, we have (jc(f), y(t))T ( - 1 , l ) r as t oo. See Figure 5.5.3. y

Figure 5.5.3. The trajectory of (5.5.3) with |σ| < 1 and the initial value (φ, ψ)Τ € Xp+

with 1 < β < eT and

< β < eT, where β denotes the ratio

t i S l l ! , Pi represents the point (*(*,· + r),

+ r)) for i = 1, 2, 3.

We next consider the case where 1 < β <

- For this case, we have

x(ti + r) > σ. Note that x(t2 + τ) = — 1 + t4 € (t2 + τ, t·} + τ) such that x f a ) = σ, that is, [1 + σ - 2ßeT]e-^~tl)

< σ, therefore there exists

+ 1 =

σ.

This, together with 1 < β < ' f f f f i f , implies that t3 < t4 = ti + \n[2ßer

-

( 1 + σ ) ] - l n ( l - σ) < t3 + τ .

Clearly, is the fourth zero of (χ — σ)(γ — σ) in [0, oo). Again according to (5.5.14) and (5.5.15), we have x(t — τ) < σ and y(t — z) > afort e (/3 + τ, f4 + t). Itfollows from (5.5.3) and (5.5.4) that (x(t), y(t))T satisfies system (5.5.6) fori e te+τ, tt+τ). Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

128

5 Signal transmission delays

Therefore, for t e

+ τ, fy + τ] we have x(t) = + r) + 1] - 1 _ 4^-4^+l-V (,_„) _ 1 — e A> 1—σ

(5.5.16)

= [(1 + σ)£ - 2 e r ] e - ( i " f l ) + 1. In particular, x(t4 + r) = -1 + and y(i 4 + τ) = 1 -

4e

- 4ßeT + 1 - σ< 2ße* - (1 + σ)

2eT - (1 + σ)β 2βετ - (1 + σ)

(1

-σ)β~τ.

Note that t4 + τ < ts + 2τ, and that from (5.5.15) we have x(t — r) > σ and y(t — τ) > σ for t € (t4 + τ, tj, + 2τ). Consequently, (x(t), y(t))T satisfies system (5.5.5) for t G {t4 + r, tj + 2r). Therefore, for t e[t4 + τ, tj + 2τ] we have x(t) = e-^-k~T)[x(t4 + τ) + 1] - 1 _ 4e2r_4ßer + 1_a2 ((_u) e A' — l—σ y(t) = e-^~THy(t4 + r) + 1] - 1 2r T _ Aße —4e +(1—σ2)β —fr—fi) _ ι — e A1—σ

(5.5.17)

Let ts be the fifth zero of (jc — a)(y — σ) in [0, oo), i.e., the first zero after t4 of (x — a)(y — σ). Then (x(t), y(t))T satisfies system(5.5.5)fori e (ί4 + τ, ts + r), and so (5.5.17) holds for all t e [ί4 + τ, ί 5 + τ], On the other hand, from (5.5.15), (5.5.16), (5.5.17) and the fact that β > 1, by solving the equation (x(t) — a)(y(t) — σ) = 0, we can obtain t5 = h+

ln[4elT - 4ßez + 1 - σ2] - ln(l - σ 2 ).

This, together with (5.5.17), implies that x(ts + r) = -1 + (1 and y(f5 + T) = - 1 + ff

^ ß

<

+σ)β~τ

4ße2x - 4eT + (1 -

σ2)β

4e2z - 4ßeT + 1 - σ2

(1+aJe-

1 3 W 1 ' t h e n '5 > t3 + r and y(t5 + r) > σ. Moreover,

from (5.5.16) and (5.5.17), it is easy to see that * , 5 + r ( 0 ) = x(ts + τ + θ) < σ and Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

129

(-1,1)

Figure 5.5.4. (
The trajectory of (5.5.3) with |σ| < 1 and the initial value

X + ' + with \ < β < ex and ^ g f f i

e

where β denotes the ratio y(ti + τ)) for i = 1, 2, 3, 4, 5.

>V5+T(0)

=

y(ts

+ τ + θ)

< β <

, Pi represents the point (*(*,• +

τ),

> σ

for θ € (—τ, 0). This shows that * ί 5 + τ € C~ and >V5+T e C+. So, by Case 1, we have (x(t), y ( 0 ) r ( - 1 , l ) r as t oo. See Figure 5.5.4. It now remains to consider the case where 1 < β < we have y(ts + τ) < σ. Again note that β > 1 implies y(t4

+ τ) = 1 -

2eT — (1 + σ)β η τ /ι ι T 2ße - (1 + σ )

. For this case

> 1 - (1 - σ)β~τ

-

Thus, there exists te € (ί4 + τ, ts + r) such that y(te) = o, that is y(te)

=

AßelT

-

4 e r + β{\

σ2)

-

-te-ti)

_ ι

=

σ

.

1 - σ

This implies that t6 = ti + ln[AßelT

Let

- AeT + (1 40e2r 4€2τ

- 4eT _

Αβετ

σ2)β]

-

σ2)β

+ (1 +

1 _

1η(1

σ

2

Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

-

σ2).

>

σ.

130

5 Signal transmission delays

Then y(t5 + r) =-l

+ (l+a)F(ß)e~T

and

t6 = ts + In F(ß).

Note that x(h + r) = x(t5 + r) = - 1 + (1 + σ)β~τ. Replacing β and t\ by Fiß) and ts, respectively, and applying the same procedure as above, we can obtain the following: (I) For t e (ts + r, t(, + τ), (x(t), y(t)) T satisfies (5.5.6). Moreover, x(t) = (1 + σ)έΤ('-'5) - 1, y(t) = ((l+a)F(ß)

-

+ 1,

x(t6 + r) = -1 + and y(te + τ) = 1 '

e

F(ß)

.

(II) For t e (te + r, ίη + τ), where ίη = ts + ln(2e r - (1 + a)F(ß))

- ln(l - σ),

(x(t), y(t)) T satisfies (5.5.7). Moreover, x ( 0 = (1 + σ - 2 F ( ß ) e T ) e - ( t ~ ^ + 1, y(t) = ((1 + a)F(ß)

x

- 2 e

ritn-^τΛ2F(ß)e*-α+σ) n l Χ(ίη + τ) - 11 - 2e*-{\+a)F(ß)i

_

) e + 1, °>, e

T

and y(i 7 + T) = l - ( l - a ) e - r . 3T 2T /τττ^ Tf 8e4T+(—2σ2 +8σ+18)e2*+(1 +σ)2 (1 —σ ) ^ R . 4g +(5-aV ,· ^ l+q+2e 2 Ρ < 2τ 2 τ (20+4σ)^+(1+σ)(-σ -2σ+7)^ 8ί +1-σ ' (3+σ> F(yS) < eT, then x(tj + τ) < σ. Moreover, xtl+T € C~ and e C+. Therefore, by Case 1, we have (x(0, y(t))T —> (— 1, l ) r as ί —> oo. See Figure 5.5.5.

ilV) If 1 < Β < 8^+(-2σ2+8σ+18>2Γ+(1+σ)2(1-σ) · UVj Π 1 < ρ < (2ο+4<τ)^+(1+σ)(-σ2-2σ+7)^ ' then jc(?7 + τ) > σ and tl

1ι <

Ff ^ ^ \Ρ>

+ τ > ί 8 = ί5 + ln(2F(^)e r - (1 + σ)) - 1η(1 - σ).

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<

1+σ+2^2τ (3+σ)β* '

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

Figure 5.5.5. The trajectory of (5.5.3) with |σ I < 1 and the initial value (φ,ψ)τ +

Xp

x

T

with 1 < β < e and

< F(ß) < e , where β denotes the ratio

trntt! > pi represents the point (*(*,· + τ), y^ + τ)) for i = 1, 2, . . . , 7.

For ί g (i7 4- r,

+ r ) , (x(t), y ( i ) ) r satisfies (5.5.6). Moreover, _ A f - t F W

+ l - . * ^

_ ,

1—σ y(0

e

= [(1 + σ)Γ(β)

jc(r 8 + τ ) = - 1 +

- 2eT]e~(t~t5)

+ 1,

4e2T - 4F(ß)eT + 1 - σ 2 _ — —
and y(t% + τ ) = 1 - — — — T (1 - a)e 2 F ( ß ) e - (1 + σ )

\

(V) For t g (?8 + τ, ^ + τ ) , where t9 = t5 + l n ( 4 e 2 r - 4 F ( / 3 ) e r + 1 - σ1) - l n ( l - σ 2 ) , Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

132

5 Signal transmission delays

(jt(i), y ( t ) ) T satisfies (5.5.5). Moreover,

x(t)

=

y(t)

=

4elT - 4F(ß)eT

+ 1-

σ2

1—σ

χ(ί9 + τ) = - 1 + (1

_ 11

(t-h)

1-σ zt „2r „τx +ι / ι(1 ——2\ 4F(ß)e — λAe az)F(ß)

e

9

_ . v (f•h)

1,

+a)e~\

and y(t9 + T) = -l where F2(ß) r V | , Tf V J

=

(l+a)F2(ß)e-z,

+

F(F(ß)).

16gSr+8(7—a2)g3T+(l—σ2)(9—g2)gT < a 48e4r +16(2—σ2)β2τ +(1 -σ 2 ) 2 - P

8e 4 r +(-2a 2 +8g+18)e 2 r +(l+a) 2 (l-g) (20+4σ)β3τ+(1+σ)(-<τ2-2σ+7)βτ '

< 1 5 4 $ ^ ' then + T ) - σ ' m 0 r e 0 V e r ' *'9+τ ^ C " a n d y i 9 + r e C + . Therefore, by Case 1, we have (x(t), y(t))T — ( — 1 , l ) r a s i —• oo. See Figure 5.5.6.

(-1,1)

(
(l

l) ß

Figure 5.5.6. The trajectory of (5.5.3) with |σ| < 1 and the initial value (φ, ψ)Τ e X+'+ with 1 < β < e* and denotes the ratio

< Πβ) <

where β

' ^ represents the point (jc(f,· + r), y(ti + τ)) for i = 1,

2,...,9.

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133

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

3T 3 2 τ rvmTf 1 Λ · 1, < F(ß) W(R\ < 4e *+(5-σ )β ' (VII) If 1 < β < 48ί + 16(2 σ2)βίτ+(1_σ2)ί + then + τ > ho = tg + In F 2 (/3)

and y(tg + τ) < σ. For this case, we again repeat the above procedure. We now claim that for any β e (1, 0 such that xt*+TeC~

and

yt*+TeC+.

(5.5.18)

Therefore, from Case 1, we have (x(t), y(t)) T (—1, l ) r as t ->· oo. Suppose, by way of contradiction, that this claim is not true. Then the above procedure can be repeated inductively to generate a sequence {tn}^iLi such that tn is the n-th zero of (χ — o)(y — σ) in [0, oo). Moreover, ti = ln(
Un- 2 =

t4n-1 = t4n-3 + ln(2eT - (1 + a)Fn~\ß)) n l

T

t4n = t4n-3+ln(2F - (ß)e t4n+l

= Un-3 + ln(4e

2r

- ln(l - σ),

- ( 1 + σ ) ) - ln(l - σ), n

- AF ~\ß)eT

+ 1 - σ 2 ) - ln(l - σ 2 );

x(ti + τ) = - 1 + (1 + σ ) β ~ τ ,

y(fi+T)

-l+ß(\+a)e~T-

=

x(t4n-2 + τ) = -1 + „ , , . y(t4n-2 + t ) = 1

1+σ Fn~Hß)'

2e r — (1 + a)Fn~^{ß)

*

_τ ;

2Fn~l(ß)eT (1+σ) (1-σ)β~τ, x(t4n-1 + τ) = 1 T n 2e - (1 +a)F ~Hß) yifAn-1 + T ) = 1 - (1 -a)e~T· Ae JC (t4n + τ) = - 1 +

— AF (ß)e + 1 —
J X(t4n+i + τ) = - 1 + (1 + σ)ε~τ, \y(t4n+ι+τ) = — 1 + (1 + a)Fn(ß)e~T; ( x(t) =
=

(l+a)ße-«-^-l, Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

r

\

134 x(t)

5 Signal transmission delays = (1 +a)e~(f_,l)

- 1,

y(f) = [(1 + σ)β - 2e*]e-«-V

+ 1,

x(t) = [1 + σ - 2Fn-liß)eT]e-^-^ y(t)

'

= [(1 + a)F»~Hß)

€

'

=

+

τ];

+ 1,

^ f e

~

+

_ 4g2r — 4Fn~l(ß)eT + 1 — σ2 1—σ n l y(i) = [(1 + a)F ~ (ß) - 2eT]e~^-t4"-3)

1

^ +

1>

^„-2

τ

^

+ r, ί4»-ι + τ],

^ _ t € [ίφ,-ι + τ, t4n + τ]; + ι,

~σ - 4eT + (1 -
t e [Un + τ, f4«+i + τ]; _ h

1—σ

\ x{t) = (1 + I y(i) = [(1 + < r)F»(/J) -

- l, + 1,

'

G [ 4w+1

'

+ τ ' '4<M+1>-2 +

where η = 1 , 2 , F ° 0 8 ) = F1^) = and F n (/3) = F(Fn~\ß)) η >2. Furthermore, it is necessary that, for all η = 1 , 2 , . . . , x(t4n-i +τ) > σ,

i.e.,

τ];

for

ί4«-ΐ + τ > ί 4η

(5.5.19)

ί4η+ι + τ > ί 4 (η+ΐ)-2·

(5.5.20)

and y(Un+\ + τ) < σ,

i.e.,

Otherwise, if «ο is the first positive integer such that (5.5.19) or (5.5.20) does not hold, then a similar argument as above shows that (5.5.18) holds for t* = i 4 « 0 -i or t* = i 4 „ 0 +i, respectively, which means the above claim is true. From the above expressions of tn and jc (tn + τ), it is clear that (5.5.19) is equivalent to „ , 1 + σ + 2e2z n F ~\ß)< , η = 1,2,... (5.5.21) w (3 + σ )ετ and that (5.5.20) is equivalent to Fn(ß)<eT,

n = 1,2,....

(5.5.22)

On the other hand, after some simple calculations, we have d

dß

rrm (ß)

[4g2r ~ (1 + σ)2] [4e2r — (1 — σ)2] " (4e2r — 4ßeT + 1 — σ 2 ) 2 > °

(5 5 23)

· ·

and d .. 32ße2*(e* - β) + \6e2*{ß2 - 1) + 8(1 -(F(ß)-ß) = —γ 2^2 aß (4elr — 4ßex + 1 — σιΥ Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

a2)ße*

>0

(5·5·24)

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

135

for all 1 < β < eT. (5.5.23) and (5.5.24) imply that F(ß) and F(ß) - β are strictly increasing on (1, eT). Since lim (F(ß) - β) = 0, β—* ι we have F(ß) — β > 0 for all β e (1, e r ) which, together with the monotonicity of F(ß) and F(ß) - β, implies that F"(ß) > Fn~\ß)

and Fn+l(ß)

for all β 6 (1, eT) and « = 1,2,

> Fn(ß) -

Fn~\ß)

Therefore, we have

F"(ß) = ß+ > =

- Fn(ß)

ß + n

Σ

(Fi(ß)-Fi~1(ß))

ß+n~(F(ß)-ß) 2 Mß ~l)e* T T ' Ae (e -ß)+l-a?

for all β € (1, eT) and η = 1 , 2 , . . . . Note that It follows that lim Fn(ß) = oo

> 0 for β e

(l,eT).

n—> oo T

for all jÖ € (1, e ), which contradicts (5.5.21) and (5.5.22), and hence the above claim is true. Case 5. We finally consider the case where φ, ψ £ C~ and ψ(0) > φ(0) > eT ψ(0) — eT + 1. For this case, the proof is similar to that of Case 4, and thus is omitted. This completes the proof of Theorem 5.5.2. The essential part in the above proof is the connection between (5.5.3) and a onedimensional map. This will be used later in our discussions on transient behaviors. Similar arguments are also applied to show that delay-induced oscillation can be prevented by increasing the threshold or by decreasing the synaptic connection in the following frustrated network x = -x + f(y(t _y = - y - f(x(t

- r)), - τ)),

with / given in (5.5.4). In particular, we establish the existence of stable limit cycle in (5.5.25) if |σ| < 1: the existence of such a cycle is independent of the size of τ though the period and the amplitude of such a cycle depend on τ. We also show that the important parameter for nonlinear oscillations is not τ but σ, and the critical values of σ are ±1 in the sense that if |σ | > 1 then each trajectory of (5.5.25) converges to a single equilibrium. More precisely, we obtain Theorem 5.5.6. Let \σ\ < 1. Then there exist Φο = (φο, Ψο)τ € Χσ and Tq > 0 such that the solution ( χ φ ° ( ί ) , y®0(t))T

t > 7o. Moreover, lim f ^ oo [x



for every solution ( χ φ ( ί ) , y*(t))T

of (5.5.25) with initial value Φο is periodic for φ

(0 - χ °(ί)] = 0 and l i m ^ o o t / ^ O - * φ ° ( 0 ] = 0 of (5.5.25) with Φ = (φ, ψ)τ € Χσ. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

136

5 Signal transmission delays

The proof is very similar to that given above for (5.5.3). The basic idea is to show that a typical trajectory of (5.5.25) is spiraling and rotates around the point (σ,σ). Figure 5.5.7 shows the periodic orbit in case |σ| < 1, this orbit can be obtained as follows:

Figure 5.5.7. The periodic trajectory of (5.5.25) with |σ| < 1, where (φ, ψ ) τ e X a ' + satisfies PQ =

= λο for a critical value λο· The trajectory starts at

(
(φ, Ψ)

The value

is chosen so that P4 coincides with PQ to obtain a periodic orbit. Note

a = -1 + (1 + o)e~~T, b = 1 - (1 -

a)e~x.

Choose a point Po = {φ{ϋ), ψ(0)) in the 2-dimensional Euclidean space (not the phase space) suchthat (φ(0) — σ, ·ψ(0) — σ) is in the first quadrant and (φ, ψ) e The trajectory moves first along the line connecting Po to (—1, 1) until Pi = (x(t\ + τ), y(t\ + τ)), where t\ is the first time when the trajectory meets the vertical line χ = a . Then the trajectory changes its direction and moves along the line connecting Pi to (—1, —1) until P2 = (xfa + τ), y(t2 + τ)), where ti is the first time when the trajectory meets the horizontal line y = σ. In the same way, the trajectory moves to P3 = (x(t3 + r), y(t3 + τ)) and then to P4 = (x{U + τ), y(t^ + τ)) along the line connecting P2 to (1, — 1) and then the line connecting P3 to (1,1), with t^ and t4 being the next zeros of x(t) — σ and y(t) — a, respectively. The periodic orbit is obtained by choosing (φ(ϋ), ψ(0)) so that Po coincides with P4. The existence of such (^(0), ψ(0)) is guaranteed by the existence of a fixed point to a 1-dimensional increasing map. Figures 5.5.8 and 5.5.9 show that a typical trajectory spirals towards Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.5 A network of two neurons: McCulloch-Pitts nonlinearity

137

the above limit cycle from outside or from inside.

Figure 5.5.8. A typical trajectory of (5.5.25) with |σ| < 1, where the initial value {φ, ψ ) τ € is chosen so that > λο· The trajectory spirals towards the periodic orbit along broken lines: from PQ = (
Note that the existence of a limit cycle is independent of the size of r provided |σ I < 1. The next result shows that increasing \σ | is the key to prevent the occurrence of a limit cycle. Theorem 5.5.7. (i) Let \σ\ > 1 and Φ = (φ,ψ)τ € Χσ. Then, as t 00, Φ Φ τ φ T T (χ (ί),γ (ί)) ( 1 , - 1 f if a > 1, and (χ (t), y* (t)) ( - 1 , l) if σ < —I; (ii) Leta = 1. Then, as t 00, (χφ(t), y*(t))T -> ( 1 , - l ) r if either Φ e X+'+ U X~'+ U X~- or Φ € Xp- and eXp-and(p(0)^l,· (iii) Leta = - 1 . Then, as t 00, ( * φ ( ί ) , y * ( t ) ) T ( - 1 , l)7" if either Φ € X+'+ υ X+- υ χ-or Φ e X~>+ and ψ(0) = - 1 , and (χφ(ί), y<s>(t))T -* + ( - 1 , - I f if Φ € x~' and\Jr(0) Φ - 1 .

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138

5 Signal transmission delays

y

Figure 5.5.9. A typical trajectory of (5.5.25) with |σ| < 1, where the initial value (ιφ, ψ ) ) τ 6 X a ' + satisfies < λο- The trajectory spirals towards the periodic orbit along broken lines in the same way as in Figure 5.5.8, but from inside the periodic orbit.

5.6 Delay-induced transient oscillations Numerical investigations indicate that even when precautions are taken to avoid delayinduced instabilities in the Hopfield's network of neurons, oscillations may arise in the presence of delays. These are delay-induced transient oscillations (DITOs), and cannot be predicted by the analysis of the asymptotic dynamics of neural networks. Examples of DITOs were first reported in two mutually exciting neurons and in rings by Babcock and Westervelt [ 1987] and Baldi and Attyia [1994]. In Pakdaman, GrottaRagazzo, Malta and Vibert [ 1997a] and Pakdaman, Grotta-Ragazzo and Malta [ 1998a], it was observed that when DITOs appear, the duration of the transients, that is, the time required for the system to stabilize at an equilibrium, increases as the characteristic charge-discharge time of the neurons tends to zero, or, equivalently, as the delay is increased to infinity. In fact, DITOs can last so long that for practical purposes they are indistinguishable from sustained oscillations. Therefore, understanding the origin of DITOs in neural networks constitutes an important complement to the analysis of delay induced instabilities in order to avoid long lasting oscillations that deteriorate network performance.

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5.6 Delay-induced transient oscillations

139

It seems that one can attribute the presence of DITOs to the fact that solutions of (5.2.1) transiently follow those of the difference system defined by

System (5.6.1) can display attracting oscillations even when almost all solutions of the corresponding delay equation (5.2.1) are convergent. The fact that some solutions display transient oscillations before stabilizing at equilibrium is the result of the difference between the asymptotic dynamics of the continuous dynamical system (5.4.1) and the discrete network (5.6.1). The main purpose of this section is to confirm the above observation through the analytical calculation of the transient regime duration (TRD) for the following twoneuron network x(t)

= -x(t)

+ wf(y(t

- τ)),

y(t)

= -y(t)

+ wf(x(t

- τ)),

(5.6.2)

where / is the step function, i.e., f(x) = 1 if χ > 0 and —1 otherwise. Following Pakdaman, Grotta-Ragazzo and Malta [ 1998a], we show that the dynamics of system (5.6.2) can be understood in terms of the iterations of a one-dimensional map. We first provide a geometrical description of the construction of this map, and then state the general results. Assuming that the delay r is set to zero, system (5.6.2) defines a two-dimensional ordinary differential equation, the trajectories of which in the (JC, plane can be easily constructed. We denote by r\ = — (w, w), ri -- (0, 0), = (w, w), r[ = (—w, w), and rj = (w, —w). r\ and rj are the stable equilibria of the system to which most trajectories converge. Trajectories with initial conditions (w, ν) such that u > 0 and ν > 0 are the segments of the straight lines connecting the initial point to r3. An example of such a trajectory in the two-dimensional phase space is shown in Figure 5.6.1. For an initial condition (u, ν) with 0 < — u < v, the trajectory coincides with the straight line connecting the initial point to r3, until reaching the y-axis, that is, as long as x(t) < 0. From this point on, the trajectory is the segment connecting the intersection point with the y-axis to the equilibrium point r3. Thus, such trajectories are broken lines. One example is also shown in Figure 5.6.1. The trajectories of other initial conditions can be constructed in the same way.

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Figure 5.6.1. Two typical trajectories of the network (5.6.2) with delay τ = 0.

In the following, we will mainly concentrate on the influence of delays on the trajectories of initial conditions which are constant mappings with constant values (u, ν) satisfying either u > 0 and ν > 0 or 0 < — u < v. These two regions of initial conditions are referred to as regions I and II, respectively. Trajectories of other constant initial conditions can be derived from these through symmetry considerations. Since the input-output function of the neurons is a step function, delays do not alter the trajectories with initial conditions in I. These are the same as for the system in the absence of delay, that is, they are segments connecting the initial condition to Γ3, as exemplified in Figure 5.6.2. The situation is different for initial conditions in II. In the case without delay, the effect of the sign change in χ appeared instantaneously, as the trajectory switched to the straight line connecting r3 right upon crossing the y-axis. When interunit signal transmission is not instantaneous, this effect is delayed, that is, the trajectory continues along the same straight line towards the point r'3, during a time interval equal to the delay r, before switching to the segment that takes it towards the point For all trajectories starting in region II, the break points have exactly the same abscissae. Thus, there is a critical value of v, denoted by v\, such that the break point is exactly on the x-axis. It is easy to verify that υι = w(eT — 1). This point plays an important role, as all initial conditions in II whose trajectories cross the _y-axis above vi reach the stable point ri after exactly one break point, such as the one shown in Figure 5.6.2. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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Figure 5.6.2. Some typical trajectories of the network (5.6.2) with τ > 0: a trajectory starting in region I is the segment connecting the initial condition to r3; a trajectory starting from region II moves towards r3 first. If the intersection of the trajectory with the y-axis is above v\ then the trajectory reaches r^ after one break point.

Trajectories of other initial conditions will display two or more break points before they enter the region where y < 0, that is, at some point, the second variable changes sign. This sign change will affect the trajectory after a time delay, and provoke a second break point, where the trajectory switches to the straight line leading towards r\. Other break points follow this, so that eventually the broken line representing the trajectory will end up at the equilibrium point An example of a trajectory with three break points is shown in Figure 5.6.3. The analysis hinges upon the construction of a map that describes the trajectories after the second break point. Let us first note that the break point and the trajectory with an initial condition in II are entirely determined by the ordinate ν at which the trajectory first intersects the y-axis. So in the following we only take this value into consideration. Then, for a given value ν the construction of the map relies on the observation that we can find a point on the Jt-axis, denoted by g(v) in Figure 5.6.3, whose trajectory coincides exactly with that of the original initial condition after the second break point. Geometrically, this point is easily obtained as the intersection point between the jc-axis and the straight line connecting r\ to the second break point of the trajectory. Thanks to this simple Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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Figure 5.6.3. A trajectory with 3 break points.

geometrical construction and after elementary calculations about the values of χ (τ +t*) and y(r + 1 * ) where t* is the first time where the trajectory intersects the y-axis, we can obtain g(v) analytically as g(v

) =

w

(2 +

e

~T)v

2 w — (v + w)e~T

(5 6

3)

Once the point (g(v), 0) on the jc-axis is determined, we can reiterate the same analysis and construction assuming that the initial condition is in fact this new point. Thus, we can easily see that if g(v) > vi, the trajectory will have only one more break point before tending to the equilibrium r j along a straight line. This situation is represented in Figure 5.6.3. Conversely, when g(v) < v\, the trajectory will have at least two more breaking points, since it will cross the y-axis again. For such points, we can construct a point (0, s) on the y-axis such that the trajectory after the second break point, i.e., the fourth break point along the trajectory starting from the initial condition, coincides with that of (0, s). Thanks to the symmetries of the system, we have s = g(g(v)) = g2(v). If g2(v) > vi, we stop the process because after the fifth break point, the trajectory will converge to the stable equilibrium point along a straight line, otherwise, we continue to iterate g until there is η such that gn(v) > See Figure 5.6.4. Such a value η exists because g(v) > ν for all υ > 0. Figure 5.6.5 shows an example of the function g and the cobweb construction of the sequence of points g" (ν), until it becomes larger than v\. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.6 Delay-induced transient oscillations

Figure 5.6.4. The sign change of y for a trajectory starting in region Π provokes a second break point where the trajectory switches to the straight line leading to r\.

Figure 5.6.5. An example of the iteration of the map g. The curve is the graph of the function g, the horizontal line is at v\ and the straight line is the diagonal. The cobweb construction shows the successive iterates g(v) until they become larger than ν ι. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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We can formalize the above construction as follows: Theorem 5.6.1. For r = (u, υ) e R 2 such that ν > —u > 0, let V(r) = w(v + u)/(w — u) and η be the integer such that < v \ < gn(V(r))y where t^ = w(eT — 1). Then, there exist Τ > ητ and θ > 0 such that for t > T, z(t, r) = z(t — Θ, rn), where z(t, ξ) is the solution of (5.6.2) through the initial condition ξ (ξ — r or rn), and rn represents the point (0, gn( V(r))) on the y-axis for η even, and the point (gn(V(r)), 0) on the x-axis for η odd. Our interest is in the transient oscillations of the solutions, and the incidence of these on the TRD, that is, the time it takes the trajectory to reach a small neighborhood of the equilibrium point. Using g and its iterates we can characterize both aspects of the dynamics of the system. Oscillations for a system like (5.6.2) can be defined as points where either one of the variables χ or y take the value zero, in other words, a solution oscillates as long as there are times t such that x(t)y(t) = 0. We refer to such points as the zeros of the solution, and we denote by N(r, r) the number of zeros of the solution of (5.6.2) with delay r , going through the initial condition r. The larger the value of N, the more the solution oscillates. The following result shows that solutions of initial conditions r — (u, v) in region II close to the straight line u + υ = 0 can display arbitrarily large numbers of zeros: Theorem 5.6.2. Let vn = g~n(vi), then Vn

~ 2(2 + e~x)n

2w(2-e~T)n(eT - 1) x n + [(2 + e~ ) - (2 - e~T)n](er

- 1)'

We have νι(τ) > ι>2(τ) > · · · > υ^(τ) > · · · > 0, and vn tends to zero as η —• oo. For r = (u, v) e R 2 , with 0 < — u < v, we have N(r, τ) = 1, 2 ρ andlp + 1 (ρ > 1) for υ > ι>ι — (1 + v\/w)u, ν = vp — (1 + vp/w)u and vp+\ — [1 + (vp+i)/w]u < ν < vp — (1 + vp/w)u, respectively. Recall that the trajectory through (0, υ) is a broken line, and it converges to a stable equilibrium along a straight line after several break points. We call the time required for the trajectory to reach the starting point of the final straight line along which the trajectory converges to an equilibrium the transient region duration(TKD). The number of zeros of a solution informs us about the TRD, because successive zeros cannot be arbitrarily close to one another, and as long as the solution is oscillating, it is obviously not stabilizing at the equilibrium point. Thus, any measure of the length of transient oscillations provides a lower bound for the TRD. We have the following result: Theorem 5.6.3. We denote by T(r, x ) the transient regime duration of a solution of (5.6.2) with delay τ going through the initial condition r = (u, ν) e R 2 with 0 < —u < v. Then, T(r, τ) > ρτ for vp+\ — [1 + (vp+\)/w]u < ν < vp — (1 + vp/w)u. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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This result indicates that for fixed delay r, the TRD grows indefinitely as initial conditions get closer to the line u + ν = 0. In Pakdaman, Grotta-Ragazzo and Vibert [1998a], a numerical calculation is carried out that shows the TRD, as a function of the ordinate ν for initial conditions (—10 -3 , υ), increases abruptly as ν approaches ΙΟ" 3 . All the previous analyses were carried out for a system with fixed delay values. Our main concern is to see how the results are modified when the delay is increased. This is done in the following theorem which stems from the fact that the values νη(τ) of the bounds delimiting the regions with a given number of zeros, and hence with a given range of TRD, increase exponentially with the delay. Theorem 5.6.4. For a fixed initial condition r = (u,v) with 0 < —u < v, there is a strictly increasing sequence 0 < τ\ < T2 < · • · < Xk < · · ·, such that z(t, r), the solution of (5.6.2) with delay τ going through r, displays exactly 1, 2p, or 2p + 1 zero(s) for τ < τ\, τ = τρ, or τρ < τ < τρ+ι, respectively. In other words, N(r, τ) = 1, 2ρ, and 2ρ + I for τ < τ\, τ = τρ, and τρ < τ < τρ+\, respectively. Thus, for fixed r, Ν(r, τ) and consequently T(r, r) are increasing functions of r, when r is large enough. The speed with which these quantities increase can then be determined by remarking that ν„(τ) ^ 2weT/(n + 2) as r — o o , and by using the lower bound on the TRD provided above. We have the following result on exponential increase of transient regime duration. Theorem 5.6.5. Let r = (u, v) e R 2 with 0 < — u < v, and let ρ = [2we T /{w + V)] — 1 where the brackets indicate the integer part, and V = w(u + v)/(w — u). Then, we have T(r, τ) > ρτ or T(r, τ) > 2wreT/(w + V) as τ οο. Numerical simulations carried out in Pakdaman, Grotta-Ragazzo and Malta [1998a] confirmed that the above lower bound for TRD is in good agreement with the true value of TRD. All the analysis and results presented until this point were carried out for a steplike input-output function g. This assumption allowed us to carry out a detailed analytical description of the system dynamics. Numerical investigations show that similar results hold for the system J i(t) = -x(t) + wg(y(t - r)), Ρ ί y(t) = -y(t) + wg(x(t - τ)), ·°· ; where the signal functions are g(x) = tanh(ax) with aw > 1 (Pakdaman, GrottaRagazzo and Malta [1998a]) and for an excitatory ring (Pakdaman, Malta, GrottaRagazzo, Arino and Vibert [1997c]).

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5.7 Effect of delay on the basin of attraction We now consider the following system of delay differential equations = -yi*i(0 xi(t) = -yix2(0

+ /l + U)if(x2(t - τι)),

(5.7.1)

+ h + W2f(x\(t - τ 2 ))

with (5.7.2) We note that all results presented here can be extended to general smooth signal functions, but to describe some numerical simulations it is more convenient to specify the signal function involved. As illustrated before, many networks are designed so that stable equilibrium points represent stored information and transmission delay may render some equilibria unstable, thus deteriorating information retrieval. Much progress has been made to derive sufficient conditions on network parameters such as connection weights, network connection topology and signal delays that ensure that the behaviors of networks with or without delay are similar. In particular, in Sections 5.3 and 5.2, we provide two types of conditions: (i) Constraints that ensure that the delay does not induce the loss of information stored in the stable equilibrium points. This is achieved when the system with delay has exactly the same stable equilibria as the one without delay, that is, the delay does not alter the local stability of the stable equilibrium points. (ii) Constraints that ensure that the delay does not induce spurious stable undamped oscillations. This is achieved when both the system without delay and the one implementing delay are (almost) convergent, that is, the delay does not alter the global stability of the system. Constraints (i) and (ii) avoid delay induced changes in the asymptotic behaviors of the system. However, even under such constraints, the delay may influence important features in the system's dynamics. For instance, changing the delay can alter the boundary of the basins of attraction of the stable equilibrium points. This can be of prime importance in associative memory networks in which the position of the basin boundaries determines which information is retrieved for a given initial condition. In this section, we are going to introduce the work of Pakdaman, Grotta-Ragazzo, Malta, Arino and Vibert [1998b] for the network (5.7.1) to show that constraints (i) and (ii) may not be sufficient to preserve the shape of the basin boundaries. Namely, we consider the dynamics of a network of two neurons connected through delayed excitatory connections. This system has been chosen because it is simple enough to allow thorough theoretical and numerical analyses, while it retains essential features of large networks. Pakdaman et al. showed that this two neuron network satisfies both of the conditions stated above, that is, the delay does not affect the Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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147

local stability of its stable equilibria, and, no matter what value the delays take, the two-neuron network remains almost convergent. Yet, they showed that the boundary separating the basins of attraction of two stable equilibria depends on the delays. Clearly, we can choose the phase space X = C([—τ2, 0]; Μ) χ C([—τι, 0]; R) equipped with the supremum norm. For an initial condition φ = (φι, 02) e X, there exists a unique solution of (5.7.1), denoted by z(t, φ) = (*i (t, φ), χι(t, φ)), such that χι(θ, φ) = φχ(θ) for - τ 2 < θ < 0, χ2(θ, φ) = 02(0) for - τ ι < θ < 0 and z(t, 0) satisfies (5.7.1) for t > 0. Also for such a solution of (5.7.1), we denote by = ((*i)f(0), (*2),(0)) the element of X defined by (x\)t(&) = xx(t + θ) for Ζί(φ) θ e [ - τ 2 , 0 ] and (xi)t(ß) — x2(t + Θ) for θ e [ - τ ι , 0]. We then obtain a semiflow {zr(^)}f>o on X. In spite of the differences between (5.7.1) and their related ODEs mentioned above, there are cases in which their dynamics are, in some sense, very similar. For (5.7.1), this happens when w\W2 > 0 (positive feedback condition). In the rest of this section, we will assume that both connection weights are strictly positive, that is, wi > 0 and w 2 > 0. 7 For 0 = (0i, 02) and ψ = (ψ\, fa) in X, we say that φ > ψ (resp. φ > ψ) if 01 (0') > Ψι(θ') (resp. 0ι(0') > ψλ(θ')) for all Θ' e [ - τ 2 , 0 ] and 0 2 ( 0 ) > ψ2iß) (resp. 02(0) > ψ2(θ)) for all θ e [ - τ ι , 0]. Note that if φ > xj/, the set defined by {χ e X; 0 > χ > ψ} is an open subset of X. The positive feedback condition and Theorem 2.5 in Smith [1987] (see also discussions in Section 5.2) imply that (5.7.1) generates an eventually strongly monotone semiflow, that is, if 0 > φ' and φ φ φ' then z,(0) > Ζί(φ') for t > 2max{Tj, τ2}. Throughout this section, constant functions in X are identified with the value they take in M2. Note that z{t) = C*i (t), X2(t)) taking the value (a, b), that is, x\(t') = a for t' > — X2 and X2(t) = b for t > —x\, is a solution of (5.7.1) if and only if (a, b) satisfies the system: ί -για + Ii + wif(b) = 0, [ - n b + I 2 + w 2 f ( a ) = 0. The number and the value of the solutions of system (5.7.3) depend on the values of the parameters (γι, γ2, wi, W2, h , h ) · Generally, the parameter set can be separated into two regions, one in which the system has a unique solution and another in which it has three solutions. These solutions do not depend on the delays τ\ and Τ2· Geometrically, they are the intersection points between the curves —για + Ιι + w\f(b) = 0 and — Yib + h + w 2 / ( a ) = 0 in the (a, ft)-plane. An example of three equilibria is shown in Figure 5.7.1, and another example of one equilibrium is shown in Figure 5.7.2. 7

We point out that if u>i < 0 and u>2 < 0 then a simple change of variables y\ = x\ and y2 = — leads to a new network equation for (_vi, >2) for which the connection weights are strictly positive.

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5 Signal transmission delays

Figure 5.7.1. The case of three equilibrium points of the two-neuron system. The equilibrium points of (5.7.1) are the intersection points between the curves Γι : -na + /ι + W1 f(b) = 0 and Γ 2 : ~nb + h + f(a) = 0.

b

Figure 5.7.2. The case of one equilibrium point of the two-neuron system. The equilibrium point of (5.7.1) is the intersection point between the curves Γι : -γιa + Ii + wif{b) = 0 and Γ 2 : + I2 + f{a) = 0. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5.7 Effect of delay on the basin of attraction

In the rest of this section we will restrict our attention to the case where system (5.7.3) has three distinct solutions. The constant functions associated with these solutions constitute the equilibria of (5.7.1) and will be denoted by r\, Γ2 and r^, so that, when considered as constant functions in X, they are ordered as r\ < r2 < rj. The eventually strong monotonicity property and the results in Smith [1987] imply that the asymptotic dynamics of (5.7.1) and its related ODE (obtained by setting Π = T2 = 0) is essentially the same in the following sense: (PI) The equilibrium n , k = 1,2, 3, of (5.7.1) is locally asymptotically stable if and only if the same is true of the related ODE; (P2) The union of the basins of attraction of the stable equilibria of (5.7.1) is an open and dense set in the phase space X, the same being true for the corresponding ODE. Recall that the basin of attraction of a stable equilibrium is the set of initial conditions in X whose trajectories converge to the equilibrium point. From the above results it can be deduced that r\ and ri are locally asymptotically stable equilibrium points, ri is unstable, and the union of the basins of attraction of r\ and r3 is an open and dense set in X. So that neither (5.7.1) nor its related ODE has stable undamped oscillatory solutions. Our focus in this section is the effect of delay on the boundary Β separating the basin of attraction of r\ from that of r3. Any neighborhood of a point in Β intersects the basins of attraction of both ri and rj. The following description of the boundary separating the two basins of attraction provides the basis for a method of numerical approximations. Theorem 5.7.1. Let u be a given element in X such that u > 0. There exists a continuous, strictly decreasing map8 bu : X —> Μ such that (i) For all φ G Χ, φ + ί>«(0) u is the unique intersection between the line going through φ and directed by u (i.e., the set {φ + Xu , λ e with the boundary separating the two basins of attraction. (ii) The set {φ € X; bu (φ) > 0} is exactly the basin of attraction

ofr\.

(iii) The set {φ e X; bu (φ) < 0} is exactly the basin of attraction ofr3. (iv) The set {φ € X; bu (φ) — 0} is exactly the boundary separating the two basins of attraction. Proof. We first prove the following claim: The solution going through an initial condition smaller (resp. larger) than Γ2 and different from Γ2 converges to r\ (resp. r?,), that is, for φ Ε X, if φ < ri (resp. φ > rj) and φ φ Γ2 then ζΛΦ) converges to r\ (resp. r$) as t —»• 00 (see Figure 5.7.3J. 8

We say that bu : X —y R is strictly decreasing if φ > φ' and φ φ φ' in Χ implies bu (φ) < bu (φ')

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5 Signal transmission delays

Figure 5.7.3. The convergence of a typical trajectory starting from a point below r-i or above Γ2 in the order relation of X.

To prove the claim, let φ e X be such that φ 0. Therefore, we must have Z/(V0 r\ as t —»• 00. In the same way, it can be shown that there is ψ' smaller than φ such that zt {ψ') tends to r\. Therefore, for t sufficiently large, Zt (Φ) is bounded by two solutions converging to r\, so that Zt (Φ) tends to π . In a similar way, it can be shown that for φ > r2 with Φ φ r2-> ζάΦ) converges to r^. This proves the claim. Let u G X be such that u > 0. For a given φ e X and a real number c, we define the translated function 0C given by 0C = φ+cu. Given φ e X, there exist real number c' and c" such that φα < Γ2 if c < c' and φα > ri if c > c". From the claim we deduce that, as t 00, Zf(c) —• π if c < c' and Z/(0C) —> if c > c". Now, let r us denote by b'{φ) the supremum of the set of values of c such that Zt($c) \ as t —> 00 and by b" (φ) the infimum of the set of values of c such that zf (0C) — r j as t oo. Clearly, we have 0'(φ) < We claim that &(φ) = 0"(φ). Indeed, suppose that this is false. Then, there is an open set Q of initial conditions w e X such that φ + < w < φ + //'()«. According to the monotonicity property, solutions going through initial conditions in Q converges neither to r\ nor r3. This implies that the open set Q does not intersect the union of the basins of attraction of Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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r\ and 7-3, which contradicts property (P2). So, b'(φ) = b"(φ) and we denote this value by bu(φ). As the basins of attraction of the two stable equilibria r\ and are open sets, φ + bu(4>)u does not belong to either of the basins and is necessarily in the boundary. The characterization of the basins and the boundary described in the theorem are derived from the construction of bu. The fact that the map bu from X to R is continuous and strictly decreasing stems from the continuous dependence of the solutions of (5.7.1) on initial conditions, and from the monotonicity of the semiflow. This completes the proof. Corollary 5.7.2. Let φ belong to the boundary B, then Ztü0 converges to r\ (resp. r$) as t —> 00 for all ψ < φ (resp. ψ > φ) such that φ φ φ. Proof. Let φ be in the boundary. For φ e X such that φ < φ and φ φ φ, we have ζτ(Φ) < ζτ(Φ) for Τ sufficiently large. Let u be defined by u = ζτ(Φ) — ζτ(Φ) > 0. Then we have ζτ(Φ) = ζτ(Φ) + u. As ζτ(Φ) belongs to the boundary we have buizriif)) = 1 > 0, so ζτ(Φ), and therefore, φ belongs to the basin of attraction of r\. In a similar way, it can be shown that Ζί(φ) converges to r3 for φ > φ and φ φ φ. This completes the proof. From Theorem 5.7.1, we can deduce that the boundary has a regular structure. Indeed, let μ > 0 be in X, and Η be a hyperplane supplementary to u. Therefore, for all φ € X, we can write in a unique way φ = h+ ku, where h e Η and λ e R. ku is the projection of φ onto the line Ru along the direction H. We denote by pu (φ) — —λ, the unique real number such that φ + pu (0)w belongs to Η. Let φ belong to the boundary Β, we can write φ — h — pu{4>)u. From the definition of bu, we know that bu(h) is the unique real number such that h + bu(h)u belongs to B, so that we have necessarily ρΗ{φ) = —bu(h). From this we can deduce that the boundary is homeomorphic to the linear hyperplane Η. Corollary 5.7.3. The map φ φ + pu($)u from Β to Η is a homeomorphism with inverse: h —> h + bu (h)u from Η to Β. Corollary 5.7.4. The boundary Β is the graph of a map from the hyperplane Η to the line Rm. Proof. As the map (h, ku) —»• h + ku from Η χ Rm to X is an isomorphism, we identify these two spaces. From Corollary 5.7.3, we obtain that the boundary Β is the subset of Η χ Rm defined by {(h, bu(h)u)·, h e H}. In other words, Β is the graph of the map L from Η to Rm defined by L(h) — bu(h)u. The basin boundary can also be characterized in terms of the dynamics of the solutions. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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Theorem 5.7.5. The boundary Β between the basins of attraction of the two locally stable equilibria r\ and r3 of (5.7.1) is constituted by the initial value φ such that the solution Zt (Φ) either converges to r2 or is asymptotically periodic. A sketch of the above theorem is presented in Pakdaman, Grotta-Ragazzo, Malta, Arino and Vibert [1998b], see also the discussions in the next section. The above results show that the space X is partitioned into three disjoint subsets that are positively invariant by the semiflow generated by (5.7.1): the two basins of attraction of the stable equilibrium points and the boundary separating them. In terms of the temporal evolution of solutions of (5.7.1), this means that there are only three types of asymptotical behaviors: solutions tend to r\ or or oscillate around r^. Solutions oscillating indefinitely around r j are not likely to be observed as they are unstable. Nevertheless, the presence of these oscillating solutions, as will be shown in later sections, is important for describing the global attractor and for describing transient oscillations which can be easily obtained in numerical investigations. Theorem 5.7.1 provides a method to locate numerically some points on the boundary for a given initial condition. A given initial condition can be shifted "up" or "down", along a "positive" direction, until it crosses the boundary (Figure 5.7.4). However, φ+ bu(
Ψ

Figure 5.7.4. Schematic representation of the map bu in a plane, where r\, and r3 are the equilibria and u is a strictly positive element in X. The decreasing curve is the basin boundary. All points below the boundary are in the basin of attraction of r\, whereas all points above the boundary are in the basin of attraction of r$. A given initial point has to be translated in the same direction as u to reach the boundary if it is in the basin of attraction of r\, and in the opposite direction if it is in the basin of attraction of r3. The amount by which the initial point has to be translated to reach the boundary gives the value of bu.

carrying such a task requires extensive computations because solutions close to the boundary tend to have oscillatory transients (Babcock and Westervelt [1987]) whose duration increases exponentially with the delay (Pakdaman, Grotta-Ragazzo, Malta and Vibert [1997c]), as shown in the last section. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5.7 Effect of delay on the basin of attraction

In what follows, we present some discussions about comparison of basin boundaries. One method to compare the boundaries separating the basins of attraction of the equilibria r\ and 7-3 of (5.7.1) for different values of τ\ and T2 is to consider a restricted set of initial conditions such that there exists a one-to-one correspondence between initial conditions for different values of τι and Τ2· One method for doing this is to consider constant functions as initial conditions. In the following, we will restrict our attention to this class. For a given real number c\, let β (ci) be the unique real number such that (c\, ß(c\ )) is on the basin boundary. From the characterization of the basin boundary described above, we know that the function β is a decreasing continuous function defined on the real line. The graph of β divides the (ci, C2)-plane into two regions. Points "below" this graph correspond exactly to the constant initial conditions lying in the basin of attraction of r\, and those "above" the graph to the ones belonging to the basin of attraction of r3. Thus Bc, the graph of β in the (c\, C2)-plane, can be considered as the boundary separating the basins of attraction of the two stable equilibrium points. To illustrate the effect of the delays on Bc, we consider identical neurons connected with either symmetrical weights or symmetrical delays. In both cases, we set I\ = — ^ and I2 = ψ . For these parameters, the coordinates of the equilibria r\, V2 and η are Π

=

(-a,

-b),

r2

= ( x \ u , xi«) = (0, 0),

r3 =

(a,

b),

where (a, b) is the strictly positive solution of (5.7.3). Note that in this case, the system is invariant under the change (jq, X2) — ( ~ x \ , —xi). This implies that Bc is also symmetric under the same transformation. Symmetrical weights. For identical neurons, receiving the same inputs and connected through symmetrical weights and delays, the neurons are indistinguishable from each other. Therefore, the basin boundary Bc is the straight line defined by c\ -

x\u +C2-

X2u =

0.

Hence, in this situation, varying both delays together does not bring any change to the basin boundary. However, when the delays are no longer equal, Bc undergoes changes depending on the values of the delays. Numerical simulations in Pakdaman, Grotta-Ragazzo, Malta, Arino and Vibert [1998b] show that when T2 decreases from τι to 0, the slope of the boundary at each point increases from — 1 to a strictly negative limit value which depends on ri. Symmetrical delays. For neurons connected through symmetrical delays but nonsymmetrical weights, changes in the delays may modify the graph Bc. Again, numerical simulations in Pakdaman, Grotta-Ragazzo, Malta, Arino and Vibert [1998b] show that the boundary Bc changes for nonsymmetric weights, when the delay is changed. In particular, the slope of the boundary decreases as the delays ri = %2 are increased from 0 to 2, except at the unstable point V2. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5 Signal transmission delays

The two-neuron network studied here is representative for irreducible cooperative networks, since the theoretical description of the basin boundary, and more generally, the global picture of the phase portrait derived for the two-neuron network can be extended to irreducible cooperative networks of arbitrary size (Pakdaman, Malta, Grotta-Ragazzo and Vibert [1996]).

5.8

Synchronized activities

Assuming the network considered in (5.7.1) consists of two identical neurons without external inputs, we obtain the following: *l(0

-

-μχι(ί)

+ f(X2(t

-

τ)),

X2(0

=

-μΧ2(0

+ f(xi(t

-

τ)).

(5.8.1)

In what follows, we will assume that / : R —»• R is C2-smooth with / ( 0 ) = 0, l i m ^ ± 0 0 f{x) = / ± 0 0 e R , / ' ( 0 ) > μ and / ' ( * ) > Oforall* € R, a n d / " ( * ) * < 0 for Λ: Φ 0. It is easy to see that (5.8.1) has three equilibria ( ξ _ , £_), ( 0 , 0 ) and (£+, £+) with ξ~ and ξ+ denoting the negative and positive zeros of (5.8.2)

μξ = / ( ξ ) ,

and ξ± denoting the constant mapping on [—1,0] with the constant value also that / ' ( f ± ) < μ. See Figure 5.8.1.

/(*)

ξ

Figure 5.8.1. The graphs of y = / ( ξ ) and y = ξ intersect at (ξ~,ξ~), and ( $ + , £ + ) with /'(ξ±)

< μ.

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(0,0)

Note

5.8 Synchronized activities

155

Then results in Section 5.7 shows that almost every trajectory is convergent to (ξ-, £_) or (£+, More precisely, there exists a closed subspace Η of codimension 1 in the phase space C([—r, 0]; R 2 ) and a homeomorphism g : Η -> g(H) = Β c C([—r, 0]; R 2 ) such that the graph Β = g(H) is exactly the boundary separating the basins of attraction of (£_, £_) and (£+, £+). Moreover, this boundary consists of the initial value (φ\,φι) e C([—r, 0]; R 2 ) whose trajectories converge either to (0,0) or to a periodic orbit. In this section, we describe the detailed dynamics of the semiflow of (5.8.1) restricted to the boundary Β and depict the geometric and topological structures of Β. Note also that the above discussions show that, due to the nature of identical decay rate μ and synaptic delay r of the two neurons, almost every trajectory is eventually synchronized (convergent to the synchronous equilibria (£_, £_) or (£+, £+)X here a solution (JCI, X2) of (5.8.1) is synchronous if χχ = χι on the common domains of definition. Clearly, if we let ζ denote the common value of xi = X2, then a synchronous solution is completely characterized by the scalar delay differential equation z(t) = -μζ(ί)

+ f(z(t - τ)).

(5.8.3)

If we further normalize the delay τ by x(t) = z(rt) then we get an equivalent system x(t) = -τμχ(ί)

+ zf(x(t

- 1)).

Renaming the parameter and the signal function, we arrive at x(0 = -μχ(0

+ f(x(t - 1))

(5.8.4)

with parameter μ > 0 and a C2-smooth bounded nonlinearity / : Μ —»· Μ satisfying all conditions stated in the above discussion. We now summarize the results from the monograph of Krisztin, Walther and Wu [1999] and explain the major steps towards these results. Every element φ e C([—1,0]; M) determines a solution χφ : [—1, oo) —>· R of equation (5.8.4). The relations F(t,) — xf

define a continuous semiflow F : R + χ C —> C. Using the associated variation-ofconstants formula jc(f) = β-μ'χ(0)

+ f e-^-s)f(x(s Jo

- 1)) ds,

one can easily show that all maps F(t, •) with t > 0 are injective and continuously differentiable and F is monotone with respect to the pointwise ordering < on C. The positively invariant set S = {φ eC·, (χφ)~Ηθ) is unbounded} Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5 Signal transmission delays

separates the domain of absorption into the interior of the positively invariant cone Κ = {φ e C ; 0 ( j ) > 0 for all 5 € [ - 1 , 0 ] } from the domain of absorption into the interior of —AT. Such a separatrix has the following nonordering property: if φ, ψ e C and φ < ψ then either φ e C \ S or ψ e C \ S. Or equivalently, S does not contain two distinct elements which are ordered. In order to derive a graph representation of 5, we linearize equation (5.8.4) at the equilibrium 0. The derivatives {D2F(t, 0)}f>o form a strongly continuous semigroup and satisfy D2F{t,m

=

yt,

with the solution y^ : [—1, oo) —• R of the linearized equation y{t) = - ß y ( t ) + f'(0)y(t

- 1)

(5.8.5)

given by — <j>. As discussed in Section 5.4, the spectrum σ of the generator of the semigroup consists of simple eigenvalues which coincide with the zeros of the characteristic function + μ — /'(0)e~k

ϋΒλ\->λ

G C.

There is one real eigenvalue λο· The other eigenvalues form a sequence of complex conjugate pairs (kj, λ7 ) with Reλ 7 + ι < Rely < λο

and

(2j — \)π < ImXj < 2 j n

for all integers j > 1, and Re kj — oo as j —• oo. The number of eigenvalues in the open right halfplane depends on μ and /'(Ο). Any odd integer 2j + 1, j € N, can be achieved. Let P, L, and Q denote the realified generalized eigenspaces of the generator associated with the spectral sets {λο}, {λχ, λι}, and σ \ {λο, λι, λι}, respectively. Then C = β φ Ζ , φ P. It is easy to see that Ρ = Μχ 0 with χ ο (0) = ek°9 for θ e [ - 1 , 0], We assume that /'(Ο) > μ/cos6ß,

where θμ e (t",27t) and θμ = -μίαηθμ.

(5.8.6)

As shown in Section 5.4, this is equivalent to Ο^βλι.

(5.8.7)

Then the unstable space of the generator of the semigroup {D2F(t, 0)},>o is at least 3-dimensional. In Krisztin, Walther and Wu [1999], we proved the following graph representation theorem. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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157

Theorem 5.8.1. There exists a map Sep : Q φ L ->· Ρ such that S = {χ + Sep(x); χ € Q φ L) and || SepOc) - Sep(x)|| < β λ °||χ - χ||

/or χ, χ e β φ L .

For each φ € C, let φρ, φ®, φL denote the projections of φ onto P, Q, L, respectively. If φ e C \ S, then φρ Φ Sep( 0, we obtain φρ < Sep( Sep( (ξ-, ξ-) as t -»• oo.

as t —> oo, and if φ is below S

A local approximation of S near zero is possible by the linearization (5.8.5). To be more precise, we recall that there exists a 3-dimensional C' -submanifold Wioc of C which is tangent at 0 to the realified eigenspace L Θ Ρ of the spectral set {λο, λι, λ ι} and has the property that for every φ e Wioc there exist a solution x : R —> R and i e l such that xq = Φ and xs e W\oc for all s < Τ. The forward extension W = F(R+ χ Wioc) of Wioc is an invariant subset of C. In the following, for a given Banach space E, we use E e to denote the open ball in Ε centered at the zero and of radius ε > 0. Then we have the following local approximation theorem for Wioc Π S. Theorem 5.8.3. There exist η > 0 and ε > 0 and a Lipschitz continuous map Sep^ : Q Φ Ρ such that Sep„(0) = 0, (Wioc Π S ) η

S e p ^ L ^ c Q@P E , ( 0

+

Lr,

+

Ρ

ε

) =

{ χ +

Sep „(χ); χ

€

L„}.

We refer to Figure 5.8.2 for illustration. Recall that W c C is invariant. Later we will see there exists a close relation between W and the global attractor of (5.8.4), and thus it is important to describe the structure of W. The following is the main result in Krisztin, Walther and Wu [1999]. Theorem 5.8.4. W is invariant, and the semiflow defines a continuous flow Fw '• R χ W —> W. For W and for the part W H S of the separatrix S in W, we have the following graph representations (see Figure 5.8.3 for illustration): Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5 Signal transmission delays Q

Ε

Gι

(codim 0 = 3)

(codim Ε = 3)

(dimGj = 1)

Figure 5.8.3. Graph representations of W and W f l 5 .

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5 . 8 Synchronized activities

159

There exist subspaces G2 C G3 of C of dimensions 2 and 3, respectively, a complementary space Gi 0/G2 in G3, a closed complementary space Ε of Gz in C, a compact set Ds C G2 and a compact set Dw C G3, and continuous mappings w : Dw —> Ε and ws : Ds G\® Ε such that9 W = {χ + w(x); χ e Dw},

W Π 5 = {χ + ws(x)·, χ e Ds}.

We have Dw = dDwU Dw,

W = {X + w(x); χ

eDw},

ο

and the restriction of w to Dw is C1 -smooth. The restriction of Fw to R χ W is C1 -smooth. The domain Ds is homeomorphic to the closed unit disk in ]R2, and consists of the trace of a simple closed C 1 -curve and its interior. The map ws is C1 -smooth in the sense that ws | ° is Ds Cl-smooth, and for each χ Ε d Ds there is an open neighborhood Ν of χ in G2 so that ws \nhDs extends to a C1-map on N. C o n c e r n i n g t h e d y n a m i c s , w e h a v e that

the set 0

= wns\(wns)

=

{x +

«;s(x);/

e

dDs}

is a periodic orbit, and there is no other periodic orbit in W. The open annulus (W il S) \ {0} consists of heteroclinic connections from 0 to Θ. (Namely, F(t, φ) 0 as t -> —00 and F(t, φ) ^ Θ as t ^ 00 for every φ in the open annulus). For every φ e W, Fw(t, φ) 0 as t —> —00. W is compact and contains the equilibria (£_, ξ-) and (£+, §+) in C. For every φ e W. (ξ-,ξ-)<φ<(ξ+,ξ+). There exist homeomorphisms from W and Dw onto the closed unit ball in R3, which send bdW = W \

W =

{ x +

w(x)\

χ e

dDw}

and

8DW

onto the unit sphere S2 C M3. If we define χ_ and χ+ by (ξ-, ξ~) = χ _ + ι υ ( χ - ) a n d ( ξ + , £ + ) = χ + + νυ(χ+), respectively, then the set dDw \ { χ - , χ+} is a 2-dimensional C 1 -submanifold of Gi, and the restriction of w to Dw \ {χ~, X+} is C1 -smooth (in the sense explained Q

0

'For a given set Μ of a topological space, we use M, W and 3Μ to denote the interior, closure and boundary of M, respectively.

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5 Signal transmission delays

above for ws). The points φ e W \ (W Π S) above the separatrix S form a connected set and satisfy Fw(t, φ) —>• (ξ+, £+) as t —> oo, and all φ G W \ (W Π S) below the separatrix S form a connected set and satisfy Fw(t, φ) —> (£_, £_) as t —> oo. Finally, for every φ in the set bd W and different from £_) and £+)> Fw(t, φ) Θ as t —*• — oo. Combining all the results stated so far, one may visualize the invariant set W as a smooth solid spindle which is split by an invariant disk into the basins of attraction towards the tips (ξ-, ξ-) and (£+, £+), see Figure 5.8.4.

Figure 5.8.4. A smooth solid spindle in the attractor of a scalar delay positive feedback equation.

The first steps toward these results exploit the monotonicity of the semiflow. Also important is the observation that the set W is contained in the order interval between the stationary points (£_, ) and (£+ and that there are heteroclinic connections from 0 to (£_, ξ - ) and (£+, £+)> given by monotone solutions χ : R —>• R without zeros. An important tool for the investigation of finer structures is a version of the discrete Liapunov functional V : C \ {0} ->· Ν counting sign changes, which was introduced by Mallet-Paret [1988]. Related are α-priori estimates for the growth and decay of solutions with segments in (sub-)level sets of V, which go back to Mallet-Paret [1988] and in a special case to Walther [1977,1979]. These tools are first used to characterize the invariant sets W \ {0} and W Π 5 \ {0} as the sets of segments xt of solutions χ :R-+R with α-limit set {0} which satisfy V(xt) < 2 for all ί e R and V(xt)_ = 2 for all f e 1 , respectively. Moreover, nontrivial differences of segments in W and Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.8 Synchronized activities

161

W H S belong to V _ 1 ({0, 2}) and V _ 1 (2), respectively. The last facts permit to introduce global coordinates on W and W Π S. It is not difficult to show that the continuous linear evaluation map Π2 : C 9 0 ^

(0(0), 0 ( - l ) ) r G R 2

is injective on W Π S, and the continuous linear evaluation map Π3 : C —• R 3 given by n 3 0 = (0(O),0(-l),cp(0))r and (Pr/>0)(O =

-

^c P (d>)e k o t ,

where Prp is the spectral projection onto Ρ along Q φ L, is injective on W. The inverse maps of the restrictions of Π2 and Π3 to W Π S and W, respectively, turn out to be locally Lipschitz continuous. The next step leads to the desired graph representation. We embed R 3 d Π3 W and 2 R D Π 2 (W Π S) in a simple way as subspaces G 3 D G2 into C, so that representations by maps w and ws with domains in G3 and G2 and ranges in complements Ε of G3 in C and Ε Θ G\ of Gi in C, Gi c G3, become obvious. It is not hard to deduce that W is given by the restriction of w to an open set, and that this restriction is C 1 -smooth. On W, the semiflow extends to a flow Fw : 1 x F W, and Fw is C 1 -smooth on the C 1 -manifold R χ W. Phase plane techniques apply to the coordinate curves in R 2 (or G2) of the flowlines t (-> Fw(t, 0) in the invariant set W H S , and yield the periodic orbit 0 = (Wns)\(wn5) as well as the identification Π 2 (W Π S) = int(n 2 <9), which implies that also W Π S is given by the restriction of u>s to an open subset of G2.

The investigation of the smoothness of the part W Π 5 of the separatrix 5 in W and of the manifold boundary bd W = W\W starts with a study of the stability of the periodic orbit Θ. We use the fact that there is a heteroclinic flowline in W Π S from the stationary point 0 e C to the orbit Θ, i.e., in the level set V - 1 (2), in order to show that precisely one Floquet multiplier lies outside the unit circle. It also follows that the center space of the linearized period map, or monodromy operator Μ = D2F(CO,

po),

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5 Signal transmission delays

given by po g Θ and the minimal period ω > 0, is at most 2-dimensional. The study of the linearized stability is closely related to earlier work in Lani-Wayda and Walther [1995] and to α-priori results on Floquet multipliers and eigenspaces for general monotone cyclic feedback systems with delay due to Mallet-Paret and Sell [1996a], To show that the graph WHS C V~l(2) U {0} is C 1 -smooth. We identify pieces of W Π S in a hyperplane Y transversal to Θ, as open sets in the smooth transversal intersection of W with the center-stable manifold of the Poincare return map which is associated with Y and a point po g Θ. Then we use the C'-flow Fw to obtain the smoothness of the set W Π S \ {0}. (Smoothness close to 0 and the relation T0(W nS) = L follow by other arguments.) Of course, this approach relies on a result on the existence of C 1 -smooth center-stable manifolds at fixed points of C J -maps in Banach spaces. A technical aspect of the smoothness proof for W Π 5 is that close to a fixed point of a C 1 -map with one-dimensional linear center space we have to construct open and positively invariant subsets of the center-stable manifold Wcs provided a single trajectory in Wcs but not in the strong stable manifold converges to the fixed point. Having established the smoothness of W Π S we show in a series of propositions that the manifold boundary bd W = W \ W (without the stationary points (ξ-, ξ_) and (ξ + , £+)) coincides with the forward extensions of a local unstable manifold of the period map F(a>, ·) at a fixed point po e Θ. The long proof of this fact involves the charts Π2 and Π3, and uses most of the results obtained before. The next step achieves the smoothness of a piece of W in a hyperplane Η of C transversal to the periodic orbit Θ. We construct a C 1 -smooth graph over an open set in a plane X12 C Η which extends such a piece of WC\ Η close to a point po e Θ Π Η beyond the boundary. Using the flow Fw we then derive that W and W Π S are C 1 -smooth up to their manifold boundaries, in the sense stated before. The final steps lead to the topological description of W. First we use the characterization of bd W mentioned above to define homeomorphisms from bd W onto the unit sphere S2 C M3. Then a generalization due to Bing [1983] of the Schoenfliess theorem from planar topology is employed to obtain homeomorphisms from W onto the closed unit ball in R 3 . The application of Bing's theorem requires to identify the bounded component of the complement of the set n 3 ( b d W) = S2 in R 3 as the set Π3 W, and to verify that Π3 W is uniformly locally 1-connected. This means that for every ε > 0 there exists S > 0 so that every closed curve in a subset of Π3 W with diameter less than δ can be continuously deformed to a point in a subset of Π3 W with diameter less than ε. We point that the proof of this topological property relies on the smoothness of the set

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5.8 Synchronized activities

and involves subsets of boundaries of neighborhoods of 0, (£_, £_), (£+, ξ+) in W which are transversal to the flow Fw • In order to construct these smooth boundaries we have to go back to the variation-of-constants formula for retarded functional differential equations in the framework of sun-dual and sun-star dual semigroups (Diekmann, van Gils, Verduyn Lunel and Walther [1995]). We note that for some general sigmoid functions (including / ( £ ) = tanh(a£) with some a > 0) under the condition equivalent to Re λ 2 < 0, Krisztin and Walther [ 1999] proved that W is indeed the global attractor of the semiflow generated by (5.8.4). Note also that the solid spindle W loses its smoothness at the two tips ( ξ - , £_) and (ξ+, ξ+), the singularity at these two points have been completely characterized by Walther [1998], As a final remark, we note that system (5.8.3) can be rewritten as ε ώ ( ΐ ) = - ß w ( t ) + f(w(t

- 1))

with ε = τ - 1 . Therefore, the case of large delay corresponds to the situation where ε —>• 0. This motivates the study of Chen [1999] on the limiting profiles of the periodic solutions for the equation e i ( i ) = —x(t) + f(x(t

- 1), λ)

(5.8.8)

with f € C3(R χ M; R), / ( 0 , λ) = 0 and f{x, λ) = (1 + λ)χ + αχ2 + bx3 + o(x3) as χ

Ο.10

It was shown that in case where a = 0, there is a neighborhood U of (0,0) in the (λ, ε) plane and a sectorial region S in U such that if (λ, ε) e U, then there is a unique periodic solution of (5.8.8) with period 1 + ε + 0 ( | ε | ( | λ | + |ε|)) as (λ, ε) —> (0, 0) if and only if (λ, ε) € S. Moreover, if b < 0 then for small and fixed λ = λο > 0, the set {ε; (λο, ε) 6 5} is an interval (0, ε(λο)). At the point (λο, ε(λο)), there is a Hopf bifurcation of sine waves and the periodic solution approaches a square wave as ε —> 0. That is, the periodic solution χ χ 0 , ε ( 0 c u 0 (respectively, C2x0) as ε —> 0 uniformly on compact subsets of (0, (respectively, l)), where c u 0 and C2x0 are the two distinct nontrivial fixed points of / ( · , λο). If b > 0 then for small and fixed λ = λο > 0, the set (ε; (λο, ε) e S} is an interval (ε(λο), β(λο))· At the point (λο, ε (λο)), there is a Hopf bifurcation. For small and fixed λ = λο < 0, the set {ε; (λο, ε) e S} is an interval (0, α(λο)). As ε 0, the unique solution becomes pulse-like in the following sense: the periodic solution has the property that χχ0ι£(ί) —• 0 as ε —• 0 uniformly on compact subsets of (0, U l). The magnitude of the pulse exceeds max{|cu 0 |, k2x0l}· See Figures 5.8.5 and 5.8.6. 10

The case where

f(x, λ) = - ( 1 + λ)χ + αχ2 + bx3 -I- o(x3) as x ->• 0 was discussed by Chow, Hale and Huang [1992] and Hale and Huang [1994],

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5 Signal transmission delays

Figure 5.8.5. Limiting profiles of periodic solutions when a = 0 and b < 0: square waves.

Figure 5.8.6. Limiting profiles of periodic solutions when a = 0 and b > 0: pulses. In the case where αφ 0, there exist a neighborhood U of ( 0 , 0 ) and a sectorial region S in U such that if (λ, ε) e U, then there is a unique periodic orbit, determined by the solution * λ > 6 of (5.8.8) with period 1 + ε + 0(|ε|(|λ| + |ε|)) as (λ, ε) (0,0) if and only if (λ, ε) e S. Moreover, for small and fixed λ = λο > 0, the set {ε; (λο, ε) € S} is an interval (ε(λο), β(λο)). At the point (λο, ε(λο)), there is a Hopf bifurcation. For small and fixed λ = λο < 0, the set {ε; (λο, ε) € 5} is an interval (0, α(λο)). As ε —>· 0, the unique periodic solution becomes pulse-like with the magnitude of the pulse exceeding |coa0I> where cox0 is the nontrivial fixed point of / ( · , λο) in a small neighborhood of 0. See Figure 5.8.7 for the limiting profiles in

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5.9 Desynchronization, phase-locked oscillation and connecting orbits

such a case.

ε

χ ε(λ)

. ΛΠΜΙ - λ

Figure 5.8.7. Limiting profiles for the case where α φ 0.

5.9 Desynchronization, phase-locked oscillation and connecting orbits We continue our discussion about the following system of delay differential equations J ii(0 = - μ κ ( ί ) + f(v(t - τ)), [ v(t) = -μν(ί)

+ f(u(t

- τ)),

or equivalently (after rescaling the time variable) ü{t) = - μ τ Μ ( 0 + τ/(υ(ί - 1)), v{t) = -μτν(ί)

+ rf(u(t

- 1))

(5.9.1)

for a network of two identical neurons, where τ and μ are given positive constants and / : R Ε is a C'-map satisfying the following set of conditions: / ( 0 ) = 0, / ' ( ξ ) > 0 for all £ € R (Monotone Positive Feedback); /(§) = has exactly three zeros ξ ~ < 0 < £ + andmax{/'(£~), /'(£+)} < μ < /'(0) (Dissipativeness and Instability of 0); / ( § ) = - / ( - £ ) for all $ € R (Symmetry); The function (0, oo) 3 ξ (Concavity).

€ l i s monotonically decreasing

It should be mentioned that some results described in this section (obtained in Chen, Krisztin and Wu [1999]) do not require the symmetry and concavity conditions on / , and that the function / ( $ ) = arctan(y£), ξ e R Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

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5 Signal transmission delays

or /(ξ)

= —

ζ-,

ξεΚ

satisfies the above set of conditions with γ > μ. Our ultimate goal is to describe the global attractor of system (5.9.1), and our focus here is on the existence of connecting orbits from a synchronous periodic solution to a phase-locked periodic solution, here a phase-locked 71-periodic solution of (5.9.1) is a solution satisfying u(t) = v(t — - j ) for all ί e R. The existence of such connecting orbits suggests a mechanism for desynchronization of the network. As shown in the last section, a synchronous solution of (5.9.1) is completely governed by the scalar delay differential equation for w(= u = v): + r f ( w ( t - 1)).

w(t) = -μτιν(ί)

(5.9.2)

The zeros of the characteristic equation λ + μτ-

τ/'(0)e~k = 0

(5.9.3)

form the spectrum of the generator As of the solution semiflow of the linear equation Z(f) = - μ τ Ζ ( ί ) + τ/'(0)Ζ(ί - 1).

(5.9.4)

All zeros of (5.9.3) are simple eigenvalues of As. There is one real eigenvalue, the other eigenvalues form complex conjugate pairs with real parts less than the value of the real eigenvalue. Defining 2π — arccos y j j ^

Απ — arccos y^gj

f

V [ / ' ( 0 ) ] 2 - μ2 '

Τί

~

VU'm2

- μ1 '

one finds that for τ > xs the realified generalized eigenspace of As associated with the spectral set of the eigenvalues with positive real part is at least 3-dimensional, while for rs < τ < x's it is exactly 3-dimensional. When τ > ts, there exists a A

3-dimensional local unstable manifold W ^ j o c of the solution semiflow generated by equation (5.9.2), which is, at zero, tangent to the 3-dimensional realified generalized eigenspace of the generator As associated with the positive real eigenvalue and the pair of complex conjugate eigenvalues with the greatest real part. We know that W3tS is a 3-dimensional submanifold of C which contains a smooth invariant disk bordered by a periodic orbit. The disk separates W34i into halves: one half contains connecting orbits from the stationary point 0 or the periodic orbit to a positive stationary point, the other half contains connecting orbits from the stationary point 0 or the periodic orbit to a negative stationary point. Moreover, if τ e (τΛ, τ5') then W ^ j is indeed the global attractor of the solution semiflow of equation (5.9.2). Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

167

5.9 Desynchronization, phase-locked oscillation and connecting orbits

It is interesting to note that the characteristic equation of the linearization at 0 of the full system (5.9.1), namely, ί X(t) = -μτΧ(ί) I Y(t) = -μτΥ{t)

+ Tf'(0)Y(t + zf'(0)X(t

- 1), - 1)

1

""

;

can be decoupled as [λ + μτ - τ/'(0)
= 0,

(5.9.6)

where the first factor corresponds to the characteristic equation (5.9.3). Defining td =

it — arccos j f j ^ Arm

1

-!*

2

'

one sees that for τ > the zeros of the equation λ + μτ + τ f'(0)e~k = 0 are simple and occur in complex conjugate pairs, at least one pair of complex conjugate zeros has positive real part, while for 0 < τ < r^ all zeros have nonpositive real parts. In Chen and Wu [1998], it was proved that when r > r j , there exists a complete analogue W o f W3,i for the full system (5.9.1). In particular, this is separated by a smooth invariant disk borded by a phase-locked periodic orbit OiConsequently, when τ > τ^(> %d), the unstable space of the generator of the Co-semigroup generated by (5.9.5) is at least 5-dimensional and there exists a 5dimensional local unstable manifold \V5j0c tangent, at zero, to the realified generalized eigenspace of the generator associated with the positive real and the two leading pairs of complex conjugate eigenvalues with positive real parts. The global forward extension Λ

Λ

Λ

Λ

Λ

Λ

of W5,ioc contains Ψτ,,ά and W-i<s = {(ψ, φ)\ φ € Wß.j}, and in particular, W5 A

A

contains a phase-locked periodic orbit O2 and a synchronous periodic orbit O4. Chen, Krisztin and Wu [1999] described completely the global dynamics of the semiflow of system (5.9.1) restricted to W5 and established the existence of heteroclinic orbits from the synchronous periodic orbit O4 to the phase-locked periodic orbit 02· The purpose of this section is to briefly describe the main results, technical tools and approach of the above work. First of all, we make the following change of variables

I

x (t) = u (2i), y(t) = v(2t - 1)

to get an equivalent cyclic system of delay differential equations ί x(t) = -2μτχ(ί) + 2τ/(γ(ΐ)), I y(t) = —2μτγ(ί) + 2rf(x{t -

f4- Q

7)

1)).

Powerful technical tools and general results have been developed in the recent work of Mallet-Paret and Sell [1996a, b]. Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

168

5 Signal transmission delays

In what follows, we are going to describe the results only for the transformed system (5.9.7), with Οa, 02, W5 and denoting the analogues of O4, Ö2, W^^, W5 and W3;ii, respectively. The fundamental technical tool for (5.9.7) is an integer-valued Liapunov functional V : C(K) \ {0} -> {0, 2 , . . . , 00}, where Κ = [ - 1 , 0] U {1}, C(K) = {φ : Κ ->• Ε; φ is continuous } is the phase space of system (5.9.7) and V(
W5 \ {0} =

φ e C(K) \ {0};

there exists a solution ζφ : Μ Ε2 v of (5.9.7) with z Q = φ, V ( z f ) < 4 for all t e Ε and zf —> 0 as t ——00

Also (T2)

V(

if φ, f € W5 and φ φ ψ.

Another important observation we make is that every non-constant periodic solution of (5.9.1) is either synchronous or phase locked. This, together with the concavity and symmetry conditions, enables us to apply the results of Krisztin and Walther [1999] about the uniqueness of periodic solutions in each level set of V to show that (

there exists at most one periodic orbit in each level set V~l(2k), k = 1 , 2 , . . . , of V.

'

One can also apply this Liapunov functional to show that (T5)

there exists no homoclinic connections of O2 and O4.

The most important application of V is perhaps the proof of the Poincare-Bendixson Theorem for cyclic systems (Mallet-Paret and Sell [1996b]) which, together with the existence of a heteroclinic connection from 0 to non-trivial equilibria of (5.9.7) (Hirsch [1988] or Smith [1987]), enables us to completely describe the dynamics of the semiflow on W5. To be more precise, we introduce the separatrix S — {φ € C(K); φ = 0 or V ( z f ) > 0 for all t > 0}. Then we show that (T6) if φ e (W5 r\S)\

{0}, then α (φ) = {0}, ω(φ) = 02 or O4,

if φ G W5 \ S, then α (φ) = {0}, ω (φ) = {χ+} or {χ_}, if φ G bd ήφζ

\ (S U {χ-, χ+}), then α (φ) = 0 4 or 02, ω{φ) = {χ+} or {χ_},

(bd W5 (Ί S) \ { 0 2 U O4}, then α (φ) = 0 4 and ω (φ) = 02, Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5.9 Desynchronization, phase-locked oscillation and connecting orbits

where bd W5 =

169

W5\W5.

To show that there is indeed a connecting orbit from Ο4 to O2, we need further information about the Floquet multipliers of O4. We apply the general theory of Mallet-Paret and Sell [1996a] and some type of homotopy argument to show that

(T7)

there exist exactly 3 Roquet multipliers of O4, counting multiplicities, outside the unit circle, and the associated realified generalized eigenspace is contained in V - 1 ({0, 2}) U {0}.

An immediate consequence of (T7) implies that there exists a 3-dimensional C 1 smooth local unstable manifold W ^ i P ) of a Poincare-map Ρ associated with a certain hyperplane. This manifold contains at least one point of a connecting orbit from O4 to χ+ and at least one point of a connecting orbit from O4 to χ _ . Since W^iP) is 3dimensional, there exists a continuous curve in W ^ i P ) \ Ο4 connecting these points, and a continuity argument shows that the curve contains a point φ e W^iP) Π S, which gives that (T8)

there exists a connecting orbit from Ο4 to O2·

The above results can be depicted in Figure 5.9.1.

Figure 5.9.1. Two solid spindles and their connecting orbits in a system of two identical neurons. The left spindle is separated by a disk bordered by a phaselocked oscillation, and the right spindle is separated by a synchronous periodic orbit. There exist connecting orbits from the synchronized to the phase-locked orbit.

Unauthenticated | 80.253.17.194 Download Date | 11/12/12 3:34 PM

5 Signal transmission delays

170

There are several questions remaining to be answered. Denoting by W u (04) the forward extension of we conjecture that (bd

η S) \ {02 υ o4} = (wu(04) η S) \ {o2 υ oA} = {φ I α(φ) = and ω(φ) = O2}.

Also, note that ψ e (W5 Π S) \ {0}

ω(φ) = 02 or 0 4 .

It is thus important to describe Hi = {
i = 2,4

(the basins of attraction of 0,· in W5). Finally, we note that the next critical value of 3π—arccos x when (5.9.6) has a pair of purely imaginary zeros is t2 j = τ^). It is then reasonable to expect that W5 is the global attractor of (5.9.1) if r e (Ts, Limiting profiles of the synchronous periodic orbit and the phase-locked periodic solution have been described in Chen and Wu [1999c], pulses and square waves are both observed.

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P. van den Driessche and X. F. Zou [1998], Global attractivity in delayed Hopfiled neural network models. SI AM J. Appl. Math. 58, 1878-1890. H.-O. Walther [1977], Über Ejektivität und periodische Lösungen bei Funktionaldifferentialgleichungeng mit verteilter Verzögerung. Habilitationsschrift, Universität München. H.-O. Waither [1979], On instability, ω-limit sets and periodic solutions of nonlinear autonomous differential delay equations. In: Functional Differential Equations and Approximation of Fixed Points (H.-O. Peitgen and H.-O. Walther, eds.). Lecture Notes in Math. 730, Springer-Verlag, New York, 489-503. H.-O. Walther [1998], The singularities of an attractor of a delay differential equations, Funct. Differential Equations 5, 513-548. H. R. Wilson and J. D. Cowan [1972], Excitatory and inhibitory interactions in localized populations of model neurons. Biophys. J. 12, 1-24. J. Wu [1998], Symmetric functional differential equations and neural networks with memory. Trans. Amer. Math. Soc. 350, 4799-4838. J. Wu [1999], Dynamics of neural networks with delay: attraction and content-addressable memory. In: Proc. Internat. Conference on Dynamical Systems and Nonlinear Mechanics (K. Vajravelu, ed.), Florida, 1999. J. Wu, T. Faria and Y. Huang [1999], Synchronization and stable phase-locking in a network of neurons with memory, Math. Comput. Modelling 30, 117-138. S. Yoshizawa, M. Morita and S. I. Amari [1993], Capacity of associative memory using a nonmonotonic neural model. Neural Networks 6, 167-176.

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Index

α-limit set, 94 5 1 -equivariant degree, 111 Τ-periodic orbit, 94 ω-limit point, 62 ω-limit set, 62, 94 0-1 distribution, 43 Boltzman machine, 70 Brain-State-in-a-Box (BSB), 69 CS, 27, 59 Cohen-Grossberg Convergence Theorem, 66 Cohen-Grossberg system, 64 CAM, 61 Eigen and Schuster equation, 69 Floquet multiplier, 94, 161, 169 GA, 51, 53 Gilpin and Ayala system, 69 Gronwall's inequality, 27 Grossberg's Learning Theorem, 57, 58 Hartline-Ratliff-Miller equation, 69 Hebb's law, 11 Hodgkin-Huxley model, 11 Hopf bifurcation, 89, 105 Kirchhoff's law, 78 LaSalle's Invariance Principle, 62, 63 Liapunov (energy) function, 62,63,70,72, 74, 75 Liapunov functional, 97, 168 May and Leonard's model, 71 McCulloch-Pitts model, 15, 69, 117 Pavlovian conditioning, 27 Perron property, 103 Perron-Firobenius theorem, 84 Poincare-Bendixson, 168 Poincare-map, 169 Ranvier node, 5 UCR, 27 UCS, 59 Wilson-Cowan equation, 18 Volterra-Lotka equation, 68 absolute refractory period, 4

absolutely globally attractive, 98 action potential, 2, 3, 9 activation dynamics, 23 activation function, 16 activity, 17 additive STM, 11 admissible, 66 amplification function, 16 amplifier, 77 artificial neural network, 8 associative memory, 146 asymptotically autonomous, 46 asymptotically stable, 61, 83, 94, 99,107 asynchronous equilibria, 120 asynchronous, 89, 119 attractivity, 95, 98 axon hillock, 2 axon, 1 backward propagation, 53 basin boundaries, 146 basin of attraction, 83, 94,146,149,170 bias, 17 brightness contrast, 36 canonical form, 80 capacitor, 77 cascade of neural nets, 52 cascade of neurons, 19 cell body, 1 cell, 8 cellular neural network, 15 center manifold, 108 center-stable manifold, 162 central nervous system, 7 chaos, 89 characteristic equation, 99, 167 characteristic function, 156 characteristic value, 99 chemical synapse, 2 chord cycle, 72 chord, 72

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180

Index fully connected, 20 fully frustrated, 104

circuit diagram, 77 circuit, 7 combinatorial matrix theory, 71 combinatorially symmetric, 71 command neuron, 24 computational performance, 90 concavity, 165 conditional stimulus, 27, 57 connected, 72 content, 61 content-addressable memory, 67 contour enhancement, 40,49 contour enhancing, 49 contraction, 96 contractive neural network, 88 converge, 82 current-voltage characteristics, 3 cycle, 104 cyclic system, 168 cytoplasm, 2

generator, 156 generic algorithm (GA), 51, 53 generic convergence, 98 global Hopf bifurcation theorem, 111 global attractor, 88, 95, 157 global continuation, 110 global coordinate, 161 global existence, 110 global pattern formation, 68 gradient-descent, 53 graph representation, 156, 161 graph theoretic test, 82 heteroclinic connection, 160 hidden neuron, 19 homotopy, 169 hyperpolarization, 3

delay differential equation, 90, 91 delay distribution, 91 delay-induced instability, 101 dendrite, 1 depolarization, 3 desynchronization, 89, 119, 166 digraph, 71 directed graph (digraph), 103 direction, 108, 109 discrete Liapunov functional, 160 discrete, 90 dissipativeness, 165 domain of absorption, 156 domain of attraction, 89 energy, 70 equilibrium, 2, 61, 94 eventually strongly monotone, 89, 98, 147 excitatory, 6 fair, 40, 42,49 feature detector, 51 feedback network, 18, 21 feedforward network, 18 frustrated network, 89, 104 frustration, 89

impedance, 77 impulse, 2, 3 inhibitory signal threshold, 68 inhibitory, 6 initial time, 92 initial value problem, 92 initial value, 92 input layer, 18 input neuron, 18, 24 input-output function, 16 instar, 30 invariant disk, 160 invariant, 62, 94 ion channel, 3 ion pump, 3 irreducible, 72, 82 joint activation-weight dynamics, 22 labeled graph, 17 learning, 30, 70 limiting equation, 30, 46 limiting profile, 163 linearization principle, 83 linearization, 99 linearized equation, 156 linearized stability, 162 linearly unstable, 83

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Index local approximation, 157 locally Lipschitz continuous, 92 locally connected network, 20 locally uniform, 40 logistic function, 15 long-term memory, 9 luminance, 35

path, 71 pattern class, 33 pattern discrimiation, 60 pattern recognition, 32 pattern variable, 38 payoff function, 54 periodic solution, 88 peripheral nervous system, 7 persistent, 38 phase-locked, 88, 89, 166, 167 phase space, 91 piecewise linear function, 15 plasticity, 7 point dissipative, 95 population, 8 positive orbit, 94 positively invariant, 62 postsynaptic, 2 prediction, 60 presynaptic inhibition, 6 presynaptic, 2 propagation time delay, 11 pulse-like, 163

macromolecular evolution, 69 manifold boundary, 161 masking field model, 70 mathematical classification, 68 maximal interval of existence, 92 membrane equation, 11 modified reflectance coefficient, 32 monodromy operator, 95, 161 monotone semiflow, 98 monotone system, 62, 80 myelin sheath, 5 nearest neighbor interaction, 86 negative orbit, 94 nerve cell, 1 neural gain, 15, 99 neural network, 17 neuron, 1, 8 neurotransmitter, 2, 5 node, 8 noise-saturation dilemma, 34 nonlinear amplifier, 77 normal form, 108 normalized, 40

quenching, 40, 49, 50

on-center off-surround network, 34, 64 orbitally (asymptotically) stable, 94, 109 order-preserving, 40, 98 orientation, 51 output layer, 18 output neuron, 18 output variable, 17 outstar learning, 26 outstar, 24 overshooting, 4 pairing, 57 partial ordering, 79 partially connected, 20 passive decay LTM, 11

realified generalized eigenspace, 156 recall, 26 receptive field, 20 recognization, 33, 70 recurrent network, 21 reflectance coefficients, 26, 31 relative activity, 35 relative refractory period, 4 relaxation time, 79 resistor, 77 resting potential, 3 saltatory conduction, 5 semiflow, 94 separatrix, 156 short-term memory, 9 shunting STM equation, 12 shunting equation, 11 shunting network, 34 sigmoid function, 15, 16 sign stability, 81

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182 sign symmetric, 71, 86 sign symmetry, 81 signal function, 10, 17 signaling mechanism, 2 signed digraph, 104 simplex signal set, 76 single-layer network, 18 slowly oscillating, 113 solution, 92 soma, 1 spanning tree, 72 spatial pattern, 25, 26, 31 spectrum, 156 spikes, 3 spindle, 160, 169 spine, 2 square wave, 163 stability modulus, 83 stable matrix, 84 stable, 83, 94, 99 steady-state, 52, 61 step function, 15 strength, 17 strongly continuous semigroup, 156 strongly order-preserving, 82, 98 subcritical bifurcation, 108 submanifold, 157 subthreshold, 68 sun-dual, 163 sun-star, 163 supercritical bifurcation, 108 suprathreshold, 68 sustained oscillation, 138 switching speed, 90 symmetry, 165 synapse, 2, 17 synaptic coupling coefficient, 9 synaptic coupling, 50 synaptic ending, 2 synaptic gap, 2 synaptic transmission, 5 synaptic vesicle, 2

Index synchronization, 88, 89, 120 synchronized, 105, 116, 118 synchronous equilibria, 120 synchronous, 118, 155 system, 7 threshold, 11, 17, 116 total LTM trace, 26 total activity normalization, 35 total activity, 26, 38 total input, 26 trajectory, 62, 94 transcendental equation, 100 transient oscillaiton, 89, 138 transient region duration (TRD), 144 transient, 38 transition, 120 transmission velocity, 5 transversality condition, 106 tree, 72 tridiagonal, 73, 86 trivalent, 40 ultimate periodicity, 120 ultimately periodic, 116 unconditional response, 27 unconditional stimulus, 27, 57 undirected graph, 71 uniform, 40, 50 uniform distribution, 43 unit, 8 unstable dimension, 108 unstable space, 156 unstable, 83, 94, 99 voltage amplification function, 77 voting paradox, 71 weight dynamics, 23 weight, 9, 17 weighted sum, 31

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