12.07.2015 Views

Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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Excitability 235(a)(b)frequencypulsesrampfrequency-current (F-I) relation3.88subcriticalAndronov-Hopfbifurcationhomocl<strong>in</strong>ic5.25Andronov-Hopfcurrent, I(c)(d)0.7I=5.25near subcriticalAndronov-Hopfbifurcation(e)0.7I=3.8866near saddlehomocl<strong>in</strong>ic orbitbifurcationK+ gat<strong>in</strong>g variable, nK+ gat<strong>in</strong>g variable, n0.1-80 membrane potential, V (mV) 00.1-80 membrane potential, V (mV) 0Figure 7.10: The I Na,p +I K -model undergoes subcritical Andronov-Hopf bifurcation yetcan exhibit low-frequency fir<strong>in</strong>g when pulses (but not ramps) of current are <strong>in</strong>jected.Parameters: C = 1, I = 0, E L = −66.2, g L = 2, g Na = 5, g K = 4.5, m ∞ (V ) hasV 1/2 = −30 and k = 10, n ∞ (V ) has V 1/2 = −34 and k = 13, and τ(V ) = 1, E Na = 60mV and E K = −90 mV. The shaded region denotes the attraction doma<strong>in</strong> of the rest<strong>in</strong>gstate. The <strong>in</strong>set shows a distorted draw<strong>in</strong>g of the phase portrait.need to consider its dynamics at I = I 1 with the <strong>in</strong>itial condition set to the locationof the old equilibrium marked by the white square <strong>in</strong> the figure. Such a step evokes aspike response even though the new equilibrium is stable. F<strong>in</strong>ally, shock<strong>in</strong>g the neuronresults <strong>in</strong> an <strong>in</strong>stantaneous <strong>in</strong>crease of its membrane potential to a new value. As anexercise, prove that the magnitude of the <strong>in</strong>crease equals the product of pulse widthand pulse height divided by the membrane capacitance. This shifts the <strong>in</strong>itial conditionhorizontally to a new po<strong>in</strong>t, marked by the white square <strong>in</strong> Fig. 7.9c, and results <strong>in</strong> aspike response.Now, let us revisit the Hodgk<strong>in</strong> experiments and demonstrate the fundamental differencebetween the stimulation protocols. In Fig. 7.10a,b,c we simulate the I Na,p +I K -model and show that it is Class 2 excitable <strong>in</strong> response to ramps of current but Class1 excitable <strong>in</strong> response to steps of current. The apparent contradiction is resolved <strong>in</strong>

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