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Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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382 Burst<strong>in</strong>gspik<strong>in</strong>gsupercriticalAndronov-Hopfbifurcationfoldbifurcationfoldbifurcationxurest<strong>in</strong>gfoldsupercriticalAndronov-HopffoldFigure 9.44: “Fold/Hopf” burst<strong>in</strong>g: The rest<strong>in</strong>g state disappears via saddle-node (fold)bifurcation and the spik<strong>in</strong>g limit cycle shr<strong>in</strong>ks to a po<strong>in</strong>t via supercritical Andronov-Hopf bifurcation (modified from Izhikevich 2000).thalamo-cortical relay neuron by Rush and R<strong>in</strong>zel (1994), and it was called “triangular”<strong>in</strong> earlier studies (Wang and R<strong>in</strong>zel 1995) because of the shape of the voltage envelope.As one can see <strong>in</strong> the figure, the fast subsystem can have five equilibria, two ofwhich are stable nodes. This is a consequence of the qu<strong>in</strong>tic shape of the V -nullcl<strong>in</strong>eof the fast subsystem. While the trajectory is at the lower equilibrium, the V -nullcl<strong>in</strong>emoves up, the equilibrium disappears via fold bifurcation, and the fast subsystemstarts to fire spikes. Dur<strong>in</strong>g this active period, the V -nullcl<strong>in</strong>e slowly moves down,and the spik<strong>in</strong>g limit cycle disappears via saddle-node on <strong>in</strong>variant circle bifurcation.The fast subsystem, however, is at the second stable equilibrium correspond<strong>in</strong>g to adepolarized state. The slow V -nullcl<strong>in</strong>e cont<strong>in</strong>ues to move down, and this equilibriumdisappears via another fold bifurcation, thereby clos<strong>in</strong>g the “fold/fold” hysteresis loop.Alternatively, “fold/circle” burst<strong>in</strong>g can be of the slow-wave type depicted <strong>in</strong> Fig. 9.28hav<strong>in</strong>g only three equilibria. The slow subsystem needs to be at least two-dimensional<strong>in</strong> this case, though.

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