12.07.2015 Views

Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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350 Burst<strong>in</strong>gslow <strong>in</strong>activation of <strong>in</strong>ward current120slow activation of outward current100V(t)100V(t)806040806020400200 500 1000 1500 2000 0 500 1000 1500 2000 2500 3000 3500abFigure 9.9: Hodgk<strong>in</strong>-Huxley (1952) model with three gat<strong>in</strong>g variables is m<strong>in</strong>imal forburst<strong>in</strong>g (modified from Fig. 1.10 <strong>in</strong> Izhikevich 2001).0uncoupledhalf-centeroscillator-mutual <strong>in</strong>hibition-Figure 9.10: Central pattern generation by mutually <strong>in</strong>hibitory oscillators.may argue that the model <strong>in</strong> the figure is not Hodgk<strong>in</strong>-Huxley at all, s<strong>in</strong>ce we changedthe k<strong>in</strong>etics of some currents by an order of magnitude.Th<strong>in</strong>k<strong>in</strong>g <strong>in</strong> terms of m<strong>in</strong>imal models, we can understand what is essential forspik<strong>in</strong>g and burst<strong>in</strong>g and what is not. In addition, we can clearly see that some wellknownconductance-based models form partially-ordered set. For example, the cha<strong>in</strong>of neuronal models Morris-Lecar (I Ca +I K ) ≺ Hodgk<strong>in</strong>-Huxley (I Na,t +I K ) ≺ Butera-R<strong>in</strong>zel-Smith (I Na,t +I K +I K,slow ) is obta<strong>in</strong>ed by add<strong>in</strong>g a conductance or gat<strong>in</strong>g variableto one model to get the next one. Here, A ≺ B means A is a subsystem of B.Understand<strong>in</strong>g the ionic bases of burst<strong>in</strong>g is an important step <strong>in</strong> analysis of burst<strong>in</strong>gdynamics. However, such an understand<strong>in</strong>g may not provide sufficient <strong>in</strong>formationon why the burst<strong>in</strong>g pattern looks as it does, what the neuro-computational propertiesof the neuron are, and how they depend on the parameters of the system. Indeed, weshowed <strong>in</strong> Chap. 5 that spik<strong>in</strong>g models based on quite different ionic mechanisms canhave identical dynamics and vice versa. This is true for burst<strong>in</strong>g models as well.

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