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Dynamical Systems in Neuroscience:

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214 BifurcationsIn contrast, if the equilibrium loses stability via subcritical Andronov-Hopf bifurcation,the state of the system diverges from it, which results <strong>in</strong> an immediate spike orsome k<strong>in</strong>d of a large-amplitude jump. Such a loss of stability is called hard: neither theequilibrium nor its neighborhood are attractive. The hard loss of stability usually leadsto noticeable or catastrophic changes <strong>in</strong> systems behavior, and the stability boundaryis called dangerous (Baut<strong>in</strong> 1949). Chang<strong>in</strong>g the bifurcation parameter <strong>in</strong> the oppositedirection will make the equilibrium stable aga<strong>in</strong>, but may not br<strong>in</strong>g the state of thesystem back to it. Saddle-node bifurcation is hard unless it is on an <strong>in</strong>variant circle.In this case, the loss of stability is catastrophic, i.e., lead<strong>in</strong>g to noticeable spikes, butreversible. Saddle homocl<strong>in</strong>ic orbit bifurcation is hard, regardless of whether it is subcriticalor supercritical. In general, most bifurcations <strong>in</strong> neurons or at least <strong>in</strong> neuronalmodels are hard.Review of Important Concepts• Stable equilibrium (rest<strong>in</strong>g state) <strong>in</strong> a typical neuronal model caneither– disappear via saddle-node bifurcation, which can be off or on<strong>in</strong>variant circle, or– lose stability via Andronov-Hopf bifurcation, which can be supercriticalor subcritical.These four cases are summarized <strong>in</strong> Fig. 6.46.• Stable limit cycle (periodic spik<strong>in</strong>g state) <strong>in</strong> a typical twodimensionalneuronal model can either– be cut by saddle-node on <strong>in</strong>variant circle bifurcation,– shr<strong>in</strong>k to a po<strong>in</strong>t via supercritical Andronov-Hopf bifurcation,– disappear via fold limit cycle bifurcation, or– disappear via saddle homocl<strong>in</strong>ic orbit bifurcation.These four cases are summarized <strong>in</strong> Fig. 6.47.• Some atypical (codimension-2) bifurcations may play importantroles <strong>in</strong> neuronal dynamics.• Bogdanov-Takens bifurcation separates <strong>in</strong>tegrators from resonators.Bibliographical NotesThough bifurcation theory can be traced back to Po<strong>in</strong>care and Andronov, it is a relativelynew branch of mathematics. The first attempt to apply it to neuroscience was as

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