12.07.2015 Views

Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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Chapter 4Two-Dimensional <strong>Systems</strong>In this chapter we <strong>in</strong>troduce methods of phase plane analysis of two-dimensional systems.Most concepts will be illustrated us<strong>in</strong>g the I Na,p +I K -model <strong>in</strong> Fig. 4.1:leak I L<strong>in</strong>stantaneous I Na,p IC ˙V{ }} { { }} { { }} K{= I − g L (V −E L ) − g Na m ∞ (V ) (V −E Na ) − g K n (V −E K ) , (4.1)ṅ = (n ∞ (V ) − n)/τ(V ) , (4.2)hav<strong>in</strong>g leak current I L , persistent Na + current I Na,p with <strong>in</strong>stantaneous activation k<strong>in</strong>eticand a relatively slower persistent K + current I K with either high (Fig. 4.1a) orlow (Fig. 4.1b) threshold (the two choices result <strong>in</strong> fundamentally different dynamics).The state of the I Na,p +I K -model is a two-dimensional vector (V, n) ∈ R 2 on the phaseplane R 2 . New types of equilibria, orbits, and bifurcations can exist on the phase planethat cannot exist on the phase l<strong>in</strong>e R. Many <strong>in</strong>terest<strong>in</strong>g features of s<strong>in</strong>gle neuron dynamicscan be illustrated or expla<strong>in</strong>ed us<strong>in</strong>g two-dimensional systems. Even neuronalburst<strong>in</strong>g, which occurs <strong>in</strong> multi-dimensional systems, can be understood via bifurcationanalysis of two-dimensional systems.This model is equivalent <strong>in</strong> many respects to the well-known and widely usedI Ca +I K -model proposed by Morris and Lecar (1981) to describe voltage oscillations<strong>in</strong> the barnacle giant muscle fiber.4.1 Planar Vector FieldsTwo-dimensional dynamical systems, also called planar systems, are often written <strong>in</strong>the formẋ = f(x, y) ,ẏ = g(x, y) ,where the functions f and g describe the evolution of the two-dimensional state variable(x(t), y(t)). For any po<strong>in</strong>t (x 0 , y 0 ) on the phase plane the vector (f(x 0 , y 0 ), g(x 0 , y 0 ))93

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