12.07.2015 Views

Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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Two-Dimensional <strong>Systems</strong> 1230-100-10max V(t)membrane voltage, V-20-30-40-50-60-70stableAndronov-Hopf bifurcationis somewhere hereunstableI (V)membrane voltage, V-20-30-40-50-60-70supercriticalAndronov-Hopfbifurcationrestm<strong>in</strong> V(t)periodic orbitsunstable equilibrium-800 20 40 60 80 100<strong>in</strong>jected dc-current, I-800 20 40 60 80 100<strong>in</strong>jected dc-current, IabFigure 4.35: Andronov-Hopf bifurcation diagram <strong>in</strong> the I Na,p +I K -model with lowthresholdK + current. a. Equilibria of the model (solution of (4.13)). b. Equilibriaand limit cycles of the model.In Fig. 4.35a we plot the solution of (4.13) as an attempt to determ<strong>in</strong>e the bifurcationdiagram for the Andronov-Hopf bifurcation <strong>in</strong> the I Na,p +I K -model. However,all we can see is that the equilibrium persists as I <strong>in</strong>creases, but there is no <strong>in</strong>formationon its stability or on the existence of a limit cycle attractor. To study the limitcycle attractor, we need to simulate the model with various values of parameter I.For each I, we disregard the transient period and plot m<strong>in</strong> V (t) and max V (t) on the(I, V )-plane, as <strong>in</strong> Fig. 4.35b. When I is small, the solutions converge to the stableequilibrium, and both m<strong>in</strong> V (t) and max V (t) are equal to the rest<strong>in</strong>g voltage. WhenI <strong>in</strong>creases past I = 12, the m<strong>in</strong> V (t) and max V (t) values start to diverge, mean<strong>in</strong>gthat there is a limit cycle attractor whose amplitude <strong>in</strong>creases as I does. This methodis appropriate for analysis of supercritical Andronov-Hopf bifurcations but it fails forsubcritical Andronov-Hopf bifurcations. Why?Figure 4.36 depicts an <strong>in</strong>terest<strong>in</strong>g phenomenon observed <strong>in</strong> many biological neurons— excitation block. Spik<strong>in</strong>g activity of the layer 5 pyramidal neuron of rat’s visualcortex is blocked by strong excitation, i.e., <strong>in</strong>jection of strong depolariz<strong>in</strong>g current. Thegeometry of this phenomenon is illustrated <strong>in</strong> Fig. 4.37, bottom. As the magnitude ofthe <strong>in</strong>jected current <strong>in</strong>creases, the unstable equilibrium, which is the <strong>in</strong>tersection po<strong>in</strong>tof the nullcl<strong>in</strong>es, moves to right branch of the cubic V -nullcl<strong>in</strong>e and becomes stable.The limit cycle shr<strong>in</strong>ks and the spik<strong>in</strong>g activity disappears, typically but not necessarilyvia supercritical Andronov-Hopf type. Thus, the I Na,p +I K -model with low-thresholdK + current can exhibit two such bifurcations <strong>in</strong> response to ramp<strong>in</strong>g up of the <strong>in</strong>jectedcurrent, one lead<strong>in</strong>g to the appearance of periodic spik<strong>in</strong>g activity (Fig. 4.34), and thenone lead<strong>in</strong>g to its disappearance (Fig. 4.37).

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