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Dynamical Systems in Neuroscience:

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Two-Dimensional <strong>Systems</strong> 101stableunstable4.1.3 Limit cyclesFigure 4.9: Limit cycles (periodic orbits).A trajectory that forms a closed loop is called a periodic trajectory or a periodic orbit(the latter is usually reserved for mapp<strong>in</strong>gs, which we do not consider here). Sometimesperiodic trajectories are isolated, as <strong>in</strong> Fig. 4.9, sometimes they are part of a cont<strong>in</strong>uum,as <strong>in</strong> Fig. 4.13, left. An isolated periodic trajectory is called a limit cycle. The existenceof limit cycles is a major feature of two-dimensional systems that cannot exist <strong>in</strong> R 1 .If the <strong>in</strong>itial po<strong>in</strong>t is on a limit cycle, then the solution (x(t), y(t)) stays on the cycleforever, and the system exhibits periodic behavior; i.e.,x(t) = x(t + T ) and y(t) = y(t + T ) (for all t)for some T > 0. The m<strong>in</strong>imal T for which this equality holds is called the periodof the limit cycle. A limit cycle is said to be asymptotically stable if any trajectorywith the <strong>in</strong>itial po<strong>in</strong>t sufficiently near the cycle approaches the cycle as t → ∞. Suchasymptotically stable limit cycles are often called limit cycle attractors, s<strong>in</strong>ce they“attract” all nearby trajectories. The stable limit cycle <strong>in</strong> Fig. 4.9 is an attractor.The limit cycle <strong>in</strong> Fig. 4.10 is also an attractor; It corresponds to the periodic (tonic)spik<strong>in</strong>g of the I Na,p +I K -model (4.1, 4.2). The unstable limit cycle <strong>in</strong> Fig. 4.9 is oftencalled a repeller, s<strong>in</strong>ce it repels all nearby trajectories. Notice that there is always atleast one equilibrium <strong>in</strong>side any limit cycle on a plane.In Fig. 4.11 we depict limit cycles of three types of neurons recorded <strong>in</strong> vitro.S<strong>in</strong>ce we do not know the state of the <strong>in</strong>ternal variables, such as the magnitude of theactivation and <strong>in</strong>activation of Na + and K + currents, we plot the cycles on the (V, V ′ )-plane, where V ′ is the time derivative of V . The cycles look jerky because of the poordata sampl<strong>in</strong>g rate dur<strong>in</strong>g each spike.4.1.4 Relaxation OscillatorsMany models <strong>in</strong> science and eng<strong>in</strong>eer<strong>in</strong>g can be reduced to two-dimensional fast/slowsystems of the formẋ = f(x, y) (fast variable)

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