12.07.2015 Views

Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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Burst<strong>in</strong>g 385bifurcations of limit cyclessaddle-nodeon <strong>in</strong>variantcirclesaddlehomocl<strong>in</strong>icorbitsupercriticalAndronov-Hopffoldlimitcyclesaddle-node(fold)fold/circlefold/homocl<strong>in</strong>icfold/Hopffold/fold cyclebifurcations of equilibriasaddle-nodeon <strong>in</strong>variantcirclesupercriticalAndronov-HopfsubcriticalAndronov-Hopfcircle/circleHopf/circlesubHopf/circlecircle/homocl<strong>in</strong>icHopf/homocl<strong>in</strong>icsubHopf/homocl<strong>in</strong>iccircle/HopfHopf/HopfsubHopf/Hopfcircle/fold cycleHopf/fold cyclesubHopf/fold cyclebistabilitybeforethe burstbistabilityat the endof the burstFigure 9.47: Bistability, i.e., co-existence of rest<strong>in</strong>g and spik<strong>in</strong>g states, depends on thetopological type of burst<strong>in</strong>g.the amplitude of emerg<strong>in</strong>g oscillations, which could be tricky. In models, slow passagethrough supercritical Andronov-Hopf bifurcations often results <strong>in</strong> a delayed transitionto oscillations with an <strong>in</strong>termediate or large amplitude, hence such a bifurcation maylook like subcritical. In record<strong>in</strong>gs, like the one <strong>in</strong> Fig. 9.39a or <strong>in</strong> Fig. 9.46, noisedestabilizes the focus equilibrium before the subcritical Andronov-Hopf bifurcationtakes place and gives the impression that the amplitude <strong>in</strong>creases gradually, i.e., as ifthe bifurcation were supercritical.The existence of fast subthreshold oscillations <strong>in</strong>dicates that the burst<strong>in</strong>g neuronacts as a resonator, at least right before the onset of a burst. In Sect. 7.2.2 we showedthat such neurons prefer a certa<strong>in</strong> resonant frequency of stimulation that matches thefrequency of subthreshold oscillations. A resonant <strong>in</strong>put may excite the neuron and<strong>in</strong>itiate a burst or it may delay the transition to the burst, depend<strong>in</strong>g on its phaserelative to the phase of subthreshold oscillations.In contrast, all bursters <strong>in</strong> the upper half of the table <strong>in</strong> Fig. 9.23, i.e., “fold/*” and“circle/*” types, do not have fast subthreshold oscillations, at least before the onsetof each burst (see Ex. 5). The fast subsystem of such bursters acts as an <strong>in</strong>tegrator:It prefers high-frequency <strong>in</strong>puts; the higher the frequency, the sooner the transition tothe spik<strong>in</strong>g state. The phase of the <strong>in</strong>put does not play any role here.

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