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Dynamical Systems in Neuroscience:

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Excitability 269(a)current, I3002001000-100-200I (V)I fast (V)I 0 (V)-300-100 -50 0 50membrane potential, V (mV)(b)current, I2000150010005000-500current response to voltage-clamp stepsI 0 (V)I fast (V)I (V)-2 -1 0 1 2 3time, (logarithmic scale, log 10 ms)(c)current, I10005000<strong>in</strong>stantaneous I-V (d) fast I-V relation (e) steady-state I-V15002000I 0 (V)1000current, I5000I fast (V)current, I150010005000I (V)-5000 0.1 0.2time (ms)-5000 10 20time (ms)-5000 500 1000time (ms)Figure 7.46: (a) The I Na,p +I K +I K(M) -model <strong>in</strong> Fig. 7.45a has three I-V relations: InstantaneousI 0 (V ) = I leak (V ) + I Na,p (V ) describes spike upstroke dynamics. The curveI fast (V ) = I 0 (V ) + I K (V ) is the I-V relation of the fast I Na,p +I K -subsystem responsiblefor spike-generat<strong>in</strong>g mechanism. The curve I ∞ (V ) = I fast (V ) + I K(M) (V ) is thesteady-state (asymptotic) I-V relation of the full model. Dots denote values obta<strong>in</strong>edfrom a simulated voltage-clamp experiment <strong>in</strong> (b); notice the logarithmic time scale.Magnifications of current responses are shown <strong>in</strong> (c,d,e). Simulated time constants:τ Na,p (V ) = 0.01 ms, τ K (V ) = 1 ms, τ M (V ) = 100 ms.experiment, depicted <strong>in</strong> Fig. 7.46b. We hold the model at V = −70 mV and stepthe command voltage to various values. The values of the current, taken at t = 0.05ms, t = 5 ms, and t = 500 ms <strong>in</strong> Fig. 7.46b, result <strong>in</strong> the <strong>in</strong>stantaneous, fast, andsteady-state I-V curves, respectively. Notice that the data <strong>in</strong> Fig. 7.46b are plottedon the logarithmic time scale. Various magnifications us<strong>in</strong>g the l<strong>in</strong>ear time scale aredepicted <strong>in</strong> Fig. 7.46c,d, and e. Numerically obta<strong>in</strong>ed values of the three I-V relationsare depicted as dots <strong>in</strong> Fig. 7.46a. They approximate the theoretical values quite wellbecause there is a 100-fold separation of time scales <strong>in</strong> the model.

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