Dynamical Systems in Neuroscience:
Dynamical Systems in Neuroscience: Dynamical Systems in Neuroscience:
398 Bursting0membrane potential, V (mV)-10-20-30-40-50-60-700 50 100 150 200 250 300 350time (ms)Figure 9.59: Bursting in theI Na,p +I K +I Na,slow -model; see Ex. 10.y21.510.5y-nullcline10x-1-20-0.5-1-3 -2 -1 0 1 2 3xx-nullcline1I(t)0-10 200 400 600timeFigure 9.60: The phase portrait of the system in Ex. 11 shows that there is only onestable equilibrium for any value of I. Yet, the system bursts when I is periodicallymodulated.10. (Bursting in the I Na,p +I K +I Na,slow -model) Explore numerically this model withthe fast subsystem as in Fig. 6.16 and a slow Na + current with parameters:g Na,slow = 0.5, m ∞,slow (V ) with V 1/2 = −50 mV and k = 10 mV, and τ slow (V ) =5 + 100 exp(−(V + 20) 2 /25 2 ). Explain the origin of bursting oscillations whenI = 27 in Fig. 9.59.11. The Bonhoeffer–van der Pol oscillatorẋ = I + x − x 3 /3 − y ,ẏ = 0.2(1.05 + x) ,with nullclines as in Fig. 9.60, is Class 3 excitable. It has a unique stable equilibriumfor any value of I (check). Periodic modulations of I shift the x-nullclineup and down but do not change the stability of the equilibrium. Why does thesystem burst in Fig. 9.60? Explore the phenomenon numerically and explain theexistence of repetitive spikes without a limit cycle.12. Prove that the fast-slow “2+2” systemż = (1 + u + iω)z − z|z| 2 , z ∈ C ,
Bursting 399˙u = µ(u − u 3 − w) ,ẇ = µ(|z| 2 − 1) ,is a slow-wave burster, even though the slow subsystem cannot oscillate for anyfixed value of the fast subsystem z.13. (Ermentrout and Kopell 1986) Consider the system˙ϑ = 1 − cos ϑ + (1 + cos ϑ)r(ψ) ,˙ψ = ω ,with ϑ and ψ being phase variables on the unit circle S 1 and r(ψ) being anycontinuous function that changes sign. Show that this system exhibits burstingactivity when ω is sufficiently small but positive. What type of bursting is that?14. Prove that the canonical model for “subHopf/fold cycle” bursting (9.11) exhibitssustained bursting activity when 0 < a < 1. What happens when a approaches0 or 1?15. Show that the canonical model for “fold/homoclinic” bursting (9.7) is equivalentto a simpler model (eq.27 in Izhikevich 2000 and Chap. 8)˙v = v 2 + w ,ẇ = µ ,with after-spike (v = +∞) resetting v ← 1 and w ← w − d, when I is sufficientlylarge and µ and d are sufficiently small.16. Derive the canonical model for “fold/homoclinic” bursting (9.7) assuming thatthe fast subsystem is near a saddle-node homoclinic orbit bifurcation point atsome u = u 0 , which is an equilibrium of the slow subsystem.17. Derive the canonical models (9.9) and (9.10) for “circle/circle” bursting.18. Show that the averaged slow subsystems of the canonical models for “circle/circle”bursters (9.9) and (9.10) have the formandrespectively, where˙u 1 = −µ 1 u 1 + d 1 f(I + u 1 − u 2 ) ,˙u 2 = −µ 2 u 2 + d 2 f(I + u 1 − u 2 ) ,˙u 1 = −µ 1 u 2 + d 1 f(I + u 1 ) ,˙u 2 = −µ 2 (u 2 − u 1 ) + d 2 f(I + u 1 ) ,f(u) =√ uπ/2 + arcot √ uis the frequency of spiking of the fast subsystem (in Hz).
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- Page 418 and 419: 408 Solutions to Exercises, Chap. 3
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- Page 452 and 453: 442 Referencesterneurons mediated b
- Page 454 and 455: 444 ReferencesDickson C.T., Magistr
- Page 456 and 457: 446 ReferencesGuckenheimer J. (1975
398 Burst<strong>in</strong>g0membrane potential, V (mV)-10-20-30-40-50-60-700 50 100 150 200 250 300 350time (ms)Figure 9.59: Burst<strong>in</strong>g <strong>in</strong> theI Na,p +I K +I Na,slow -model; see Ex. 10.y21.510.5y-nullcl<strong>in</strong>e10x-1-20-0.5-1-3 -2 -1 0 1 2 3xx-nullcl<strong>in</strong>e1I(t)0-10 200 400 600timeFigure 9.60: The phase portrait of the system <strong>in</strong> Ex. 11 shows that there is only onestable equilibrium for any value of I. Yet, the system bursts when I is periodicallymodulated.10. (Burst<strong>in</strong>g <strong>in</strong> the I Na,p +I K +I Na,slow -model) Explore numerically this model withthe fast subsystem as <strong>in</strong> Fig. 6.16 and a slow Na + current with parameters:g Na,slow = 0.5, m ∞,slow (V ) with V 1/2 = −50 mV and k = 10 mV, and τ slow (V ) =5 + 100 exp(−(V + 20) 2 /25 2 ). Expla<strong>in</strong> the orig<strong>in</strong> of burst<strong>in</strong>g oscillations whenI = 27 <strong>in</strong> Fig. 9.59.11. The Bonhoeffer–van der Pol oscillatorẋ = I + x − x 3 /3 − y ,ẏ = 0.2(1.05 + x) ,with nullcl<strong>in</strong>es as <strong>in</strong> Fig. 9.60, is Class 3 excitable. It has a unique stable equilibriumfor any value of I (check). Periodic modulations of I shift the x-nullcl<strong>in</strong>eup and down but do not change the stability of the equilibrium. Why does thesystem burst <strong>in</strong> Fig. 9.60? Explore the phenomenon numerically and expla<strong>in</strong> theexistence of repetitive spikes without a limit cycle.12. Prove that the fast-slow “2+2” systemż = (1 + u + iω)z − z|z| 2 , z ∈ C ,