Dynamical Systems in Neuroscience:
Dynamical Systems in Neuroscience: Dynamical Systems in Neuroscience:
392 BurstingReview of Important Concepts• A burst of spikes is a train of action potentials followed by a periodof quiescence.• Bursting activity typically involves two time scales: fast spiking andslow modulation via a resonant current.• Many mathematical models of bursters have fast-slow formẋ = f(x, u) (fast spiking),˙u = µg(x, u) (slow modulation).• To dissect a burster, one freezes its slow subsystem (i.e., sets µ = 0)and uses the slow variable u as a bifurcation parameter to study thefast subsystem.• The fast subsystem undergoes two important bifurcations during aburst: (1) bifurcation of an equilibrium resulting in transition to spikingstate, and (2) bifurcation of a limit cycle attractor resulting intransition to resting state.• Different types of bifurcations result in different topological types ofbursting.• There are 16 basic types of bursting, summarized in Fig. 9.23.• Different topological types of bursters have different neurocomputationalproperties.Bibliographical NotesThe history of formal classification of bursting starts with the seminal paper by Rinzel(1987), who contrasted the bifurcation mechanism of the “square-wave”, “parabolic”,and “elliptic” bursters. Then, Bertram et al. (1995) followed Rinzel’s suggestionand referred to the bursters using Roman numbers, adding a new, Type IV burster.Another, “tapered” type of bursting was studied simultaneously and independently byHolden and Erneux (1993a,b), Smolen et al. (1993), and Pernarowski (1994). Laterde Vries (1998) suggested to refer to it as Type V burster. Yet another, “triangular”type of bursting was studied by Rush and Rinzel (1994), making the total number ofidentified bursters to be 6. To honor these pioneers, we described these six classicalbursters in the order consisted with the numbering nomenclature of Bertram et al.(1995). Their bifurcation mechanisms are summarized in Fig. 9.53.
Bursting 393Saddle-Node Saddle Supercritical FoldBifurcations on Invariant Homoclinic Andronov- LimitCircle Orbit Hopf Cycletriangular square-wave taperedFold Type I Type V Type IVSaddle-Nodeon InvariantCircleparabolicType IISupercriticalAndronov-HopfSubcriticalAndronov-HopfellipticType IIIFigure 9.53: Bifurcation mechanisms and classical nomenclature of the 6 burstersknown in the XX century. Compare with Fig. 9.23 and Fig. 9.24.The complete classification of bursters was provided by Izhikevich (2000), and itwas actually motivated by the paper of Guckenheimer et al. (1997). There is a drasticdifference between Izhikevich’s approach, and that of the scientists mentioned above.They used a bottom-up approach; that is, they considered biophysically plausible conductancebased models describing experimentally observable cellular behavior and thenthey determined the types of bursting these models exhibited. In contrast, Izhikevich(2000) used the top-down approach and considered all possible pairs of co-dimension-1bifurcations of rest and spiking states, which resulted in different types of bursting. Itwas an easy task to provide a conductance-based model exhibiting each bursting type.Thus, many of the bursters are “theoretical” in the sense that they have yet to be seenin experiments.Interestingly, a challenging problem was to suggest a naming scheme for the bursters.The names should be self-explanatory and easy to remember and understand. Thus,the numbering scheme suggested by Bertram et al. (1995) would lead, e.g., to burstersof Type XXVII, Type LXIII, Type LCXVI, etc. We cannot use descriptions such as“elliptic”, “parabolic”, “hyperbolic”, “triangular”, “rectangular”, etc., since they aremisleading. In this book we follow Izhikevich (2000) and name the bursters accordingto the two bifurcations involved, as in Fig. 9.23.Not all bursters can be represented in the fast-slow form with a clear separation
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Burst<strong>in</strong>g 393Saddle-Node Saddle Supercritical FoldBifurcations on Invariant Homocl<strong>in</strong>ic Andronov- LimitCircle Orbit Hopf Cycletriangular square-wave taperedFold Type I Type V Type IVSaddle-Nodeon InvariantCircleparabolicType IISupercriticalAndronov-HopfSubcriticalAndronov-HopfellipticType IIIFigure 9.53: Bifurcation mechanisms and classical nomenclature of the 6 burstersknown <strong>in</strong> the XX century. Compare with Fig. 9.23 and Fig. 9.24.The complete classification of bursters was provided by Izhikevich (2000), and itwas actually motivated by the paper of Guckenheimer et al. (1997). There is a drasticdifference between Izhikevich’s approach, and that of the scientists mentioned above.They used a bottom-up approach; that is, they considered biophysically plausible conductancebased models describ<strong>in</strong>g experimentally observable cellular behavior and thenthey determ<strong>in</strong>ed the types of burst<strong>in</strong>g these models exhibited. In contrast, Izhikevich(2000) used the top-down approach and considered all possible pairs of co-dimension-1bifurcations of rest and spik<strong>in</strong>g states, which resulted <strong>in</strong> different types of burst<strong>in</strong>g. Itwas an easy task to provide a conductance-based model exhibit<strong>in</strong>g each burst<strong>in</strong>g type.Thus, many of the bursters are “theoretical” <strong>in</strong> the sense that they have yet to be seen<strong>in</strong> experiments.Interest<strong>in</strong>gly, a challeng<strong>in</strong>g problem was to suggest a nam<strong>in</strong>g scheme for the bursters.The names should be self-explanatory and easy to remember and understand. Thus,the number<strong>in</strong>g scheme suggested by Bertram et al. (1995) would lead, e.g., to burstersof Type XXVII, Type LXIII, Type LCXVI, etc. We cannot use descriptions such as“elliptic”, “parabolic”, “hyperbolic”, “triangular”, “rectangular”, etc., s<strong>in</strong>ce they aremislead<strong>in</strong>g. In this book we follow Izhikevich (2000) and name the bursters accord<strong>in</strong>gto the two bifurcations <strong>in</strong>volved, as <strong>in</strong> Fig. 9.23.Not all bursters can be represented <strong>in</strong> the fast-slow form with a clear separation