Dynamical Systems in Neuroscience:
Dynamical Systems in Neuroscience: Dynamical Systems in Neuroscience:
434 Solutions to Exercises, Chap. 90.5|x|yx1x2Syy0.5x 1Supercritical Andronov-Hopf BifurcationtSlow PassageEffectFigure 10.35: Hopf/Hopf bursting without co-existence of attractors; see Ex. 6 (modified fromHoppensteadt and Izhikevich 1997).Here, u is the deviation from the slow equilibrium u 0 . The slow subsystem˙u = µg(ze iωt + complex-conjugate, u)can be averaged and transformed into the canonical form.8. (Bursting in the I Na,t +I Na,slow -model) First, determine the parameters of the I Na,t -model correspondingto the subcritical Andronov-Hopf bifurcation, and hence the co-existence of theresting and spiking states. Then, add a slow high-threshold persistent Na + current that activatesduring spiking, depolarizes the membrane potential and stops the spiking. During resting,the current deactivates, the membrane potential hyperpolarizes and the neuron starts to fireagain.9. Substitute the slow Na + current in the exercise above with a slow dendritic compartmentwith dendritic resting potential far below the somatic resting potential. As the dendriticcompartment hyperpolarizes the somatic compartment, the soma starts to fire (due to theinhibition-induced firing described in Sect. 7.2.8). As the somatic compartment fires, dendriticcompartment slowly depolarizes, removes the hyperpolarization and stops firing.10. (Bursting in the I Na,p +I K +I Na,slow -model) The time constant τ slow (V ) is relatively small inthe voltage range corresponding to the spike after-hyperpolarization (AHP). Deactivation ofthe Na + current during each AHP is much stronger than its activation during the spike peak.As a result, Na + current deactivates (turns off) during the burst, and then slowly reactivatesto its baseline level during the resting period, as one can see in Fig. 10.36.11. The mechanism of spiking, illustrated in Fig. 10.37, is closely related to the phenomenon ofaccommodation and anodal break excitation. The key feature is that this bursting is notfast-slow.
Solutions to Exercises, Chap. 9 435membrane potential,V (mV)0-20-40-600 50 100 150 200 250 300 350 400 450Na + activationgate, mslow0.50.4deactivationreactivation0.30 50 100 150 200 250 300 350 400 450time (ms)Figure 10.36: Bursting in the I Na,p +I K +I Na,slow -model. See Ex. 10.slow increase of Ifast increase of I2.52.5221.51.5110.50.50.50Rest0.50Spiking12.5 2 1.5 1 0.5 0 0.5 1 1.5 2 2.512.5 2 1.5 1 0.5 0 0.5 1 1.5 2 2.5Figure 10.37: The system has a unique attractor — equilibrium, yet it can exhibit repetitive spikingactivity when the N-shaped nullcline is moved up not very slow.
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- Page 452 and 453: 442 Referencesterneurons mediated b
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- Page 456 and 457: 446 ReferencesGuckenheimer J. (1975
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- Page 462 and 463: 452 ReferencesRosenblum M.G. and Pi
- Page 464 and 465: 454 ReferencesTuckwell H.C. (1988)
- Page 466 and 467: 456 References9
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434 Solutions to Exercises, Chap. 90.5|x|yx1x2Syy0.5x 1Supercritical Andronov-Hopf BifurcationtSlow PassageEffectFigure 10.35: Hopf/Hopf burst<strong>in</strong>g without co-existence of attractors; see Ex. 6 (modified fromHoppensteadt and Izhikevich 1997).Here, u is the deviation from the slow equilibrium u 0 . The slow subsystem˙u = µg(ze iωt + complex-conjugate, u)can be averaged and transformed <strong>in</strong>to the canonical form.8. (Burst<strong>in</strong>g <strong>in</strong> the I Na,t +I Na,slow -model) First, determ<strong>in</strong>e the parameters of the I Na,t -model correspond<strong>in</strong>gto the subcritical Andronov-Hopf bifurcation, and hence the co-existence of therest<strong>in</strong>g and spik<strong>in</strong>g states. Then, add a slow high-threshold persistent Na + current that activatesdur<strong>in</strong>g spik<strong>in</strong>g, depolarizes the membrane potential and stops the spik<strong>in</strong>g. Dur<strong>in</strong>g rest<strong>in</strong>g,the current deactivates, the membrane potential hyperpolarizes and the neuron starts to fireaga<strong>in</strong>.9. Substitute the slow Na + current <strong>in</strong> the exercise above with a slow dendritic compartmentwith dendritic rest<strong>in</strong>g potential far below the somatic rest<strong>in</strong>g potential. As the dendriticcompartment hyperpolarizes the somatic compartment, the soma starts to fire (due to the<strong>in</strong>hibition-<strong>in</strong>duced fir<strong>in</strong>g described <strong>in</strong> Sect. 7.2.8). As the somatic compartment fires, dendriticcompartment slowly depolarizes, removes the hyperpolarization and stops fir<strong>in</strong>g.10. (Burst<strong>in</strong>g <strong>in</strong> the I Na,p +I K +I Na,slow -model) The time constant τ slow (V ) is relatively small <strong>in</strong>the voltage range correspond<strong>in</strong>g to the spike after-hyperpolarization (AHP). Deactivation ofthe Na + current dur<strong>in</strong>g each AHP is much stronger than its activation dur<strong>in</strong>g the spike peak.As a result, Na + current deactivates (turns off) dur<strong>in</strong>g the burst, and then slowly reactivatesto its basel<strong>in</strong>e level dur<strong>in</strong>g the rest<strong>in</strong>g period, as one can see <strong>in</strong> Fig. 10.36.11. The mechanism of spik<strong>in</strong>g, illustrated <strong>in</strong> Fig. 10.37, is closely related to the phenomenon ofaccommodation and anodal break excitation. The key feature is that this burst<strong>in</strong>g is notfast-slow.