12.07.2015 Views

Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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Two-Dimensional <strong>Systems</strong> 970.7K + activation variable, n0.60.50.40.30.20.1(d)relativerefractory(c)absoluterefractoryn-nullcl<strong>in</strong>e(b)spikedownstroke(a)spikeupstroke(regenerative)peakof spikeV-nullcl<strong>in</strong>e0rest<strong>in</strong>g-80 -70 -60 -50 -40 -30 -20 -10 0 10 20membrane voltage, V (mV)Figure 4.4: Nullcl<strong>in</strong>es of the I Na,p +I K -model (4.1, 4.2) with low-threshold K + current<strong>in</strong> Fig. 4.1b. (The vector field is slightly distorted for the sake of clarity of illustration).As another example, let us determ<strong>in</strong>e the nullcl<strong>in</strong>es of the I Na,p +I K -model (4.1,4.2). The V -nullcl<strong>in</strong>e is given by the equationwhich has the solutionI − g L (V −E L ) − g Na m ∞ (V ) (V −E Na ) − g K n (V −E K ) = 0 ,n = I − g L(V −E L ) − g Na m ∞ (V ) (V −E Na )g K (V − E K )(V -nullcl<strong>in</strong>e)depicted <strong>in</strong> Fig. 4.4. It typically has the form of a cubic parabola. The equationdef<strong>in</strong>es the n-nullcl<strong>in</strong>en = n ∞ (V )n ∞ (V ) − n = 0(n-nullcl<strong>in</strong>e),which co<strong>in</strong>cides with the K + steady-state activation function n ∞ (V ), though only an<strong>in</strong>itial segment of this curve fits <strong>in</strong> Fig. 4.4. It is easy to see how the V - and n-nullcl<strong>in</strong>es partition the phase plane <strong>in</strong>to four regions, <strong>in</strong> each of which the vector fieldhas a different direction:(a) Both V and n <strong>in</strong>crease: Both Na + and K + currents activate and lead to theupstroke of the action potential.(b) V decreases but n still <strong>in</strong>creases: Na + current deactivates but the slower K +current still activates and leads to the downstroke of the action potential.

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