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Dynamical Systems in Neuroscience:

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Burst<strong>in</strong>g 349voltage-gatedCa2+-gated<strong>in</strong>activation of <strong>in</strong>ward currentamplify<strong>in</strong>g gateCa 2+ -gatedvoltage-gatedactivationof <strong>in</strong>wardcurrent<strong>in</strong>activationof outwardcurrentactivationof <strong>in</strong>wardcurrent<strong>in</strong>activationof outwardcurrentresonant gatevoltage-gatedCa 2+ -gatedactivation <strong>in</strong>activationof outward of <strong>in</strong>wardcurrent current<strong>in</strong>activationof <strong>in</strong>wardcurrentINa,t+IhINa,t(fast and slow)INa,t slow +IKINa,p+IK+Ihactivationof outwardcurrentamplify<strong>in</strong>g gateCa 2+ -gatedvoltage-gatedactivationof <strong>in</strong>wardcurrent<strong>in</strong>activationof outwardcurrentactivationof <strong>in</strong>wardcurrent<strong>in</strong>activationof outwardcurrentresonant gatevoltage-gatedCa 2+ -gatedactivation <strong>in</strong>activationof outward of <strong>in</strong>wardcurrent current<strong>in</strong>activationof <strong>in</strong>wardcurrentICa(N)ICa(L)+IKactivationof outwardcurrentactivation of outward currentamplify<strong>in</strong>g gateCa 2+ -gatedvoltage-gatedactivationof <strong>in</strong>wardcurrent<strong>in</strong>activationof outwardcurrentactivationof <strong>in</strong>wardcurrent<strong>in</strong>activationof outwardcurrentresonant gatevoltage-gatedCa 2+ -gatedactivation <strong>in</strong>activationof outward of <strong>in</strong>wardcurrent current<strong>in</strong>activationof <strong>in</strong>wardcurrentINa,t+IK(M)INa,p+IK+IK(M)activationof outwardcurrentamplify<strong>in</strong>g gateCa 2+ -gatedvoltage-gatedactivationof <strong>in</strong>wardcurrent<strong>in</strong>activationof outwardcurrentactivationof <strong>in</strong>wardcurrent<strong>in</strong>activationof outwardcurrentresonant gatevoltage-gatedCa 2+ -gatedactivation <strong>in</strong>activationof outward of <strong>in</strong>wardcurrent current<strong>in</strong>activationof <strong>in</strong>wardcurrentICa(T)+IAHPICa+IK+IAHPactivationof outwardcurrentFigure 9.8: Some m<strong>in</strong>imal models for burst<strong>in</strong>g.burst<strong>in</strong>g are conf<strong>in</strong>ed to the left column <strong>in</strong> Fig. 9.8. M<strong>in</strong>imal models <strong>in</strong> this columnwould provide testable hypotheses on the ionic basis of burst<strong>in</strong>g, and they could guidenovel experiments. If block abolishes burst<strong>in</strong>g, we cannot conclude that the blockedcurrent drives the burst<strong>in</strong>g — it may merely be necessary for provid<strong>in</strong>g backgroundstimulation.Notice that the I Na,tslow + I K -model and the I Na,t + I K(M) -model <strong>in</strong> the figure (seethe shaded rectangles) consist of the same gat<strong>in</strong>g variables: Na + activation gate m,<strong>in</strong>activation gate h, and K + activation gate n. Both models are equivalent to theHodgk<strong>in</strong>-Huxley model, the only difference be<strong>in</strong>g the choice of the slow gate. Thus, <strong>in</strong>contrast to the common biophysical folklore, the Hodgk<strong>in</strong>-Huxley model is a m<strong>in</strong>imalmodel for burst<strong>in</strong>g, and there are two fundamentally different ways <strong>in</strong> which one canmake it burst without any additional currents, as we show <strong>in</strong> Fig. 9.9. Of course, one

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