Dynamical Systems in Neuroscience:
Dynamical Systems in Neuroscience: Dynamical Systems in Neuroscience:
194 Bifurcationshomoclinic orbithomoclinic orbitsaddle homoclinic orbitbifurcationsaddle-node on invariant circlebifurcationFigure 6.26: Two bifurcations involving homoclinic trajectories to an equilibrium.6.2.4 HomoclinicA limit cycle can appear or disappear via a saddle homoclinic orbit bifurcation, asdepicted in Fig. 6.25. As the bifurcation parameter changes, the cycle becomes ahomoclinic orbit to the saddle equilibrium, and its period becomes infinite. After thebifurcation, the cycle no longer exists. A necessary condition for such a bifurcation isthat the steady-state I-V relation is not monotonic.One should be careful to distinguish the saddle homoclinic orbit bifurcation fromthe saddle-node on invariant circle bifurcation depicted in Fig. 6.26. Indeed, it mightbe easy to confuse these bifurcations, since both involve an equilibrium and a largeamplitudehomoclinic trajectory that becomes a limit cycle. The key difference is thatthe equilibrium is a saddle in the former and a saddle-node in the latter. The saddleequilibrium persists as the bifurcation parameter changes, whereas the saddle-nodeequilibrium disappears or bifurcates into two points, depending on the direction ofchange of the bifurcation parameter.Recall that a saddle on a plane has two real eigenvalues of opposite signs. Theirsum, λ 1 + λ 2 , is called the saddle quantity.• If λ 1 + λ 2 < 0, then the saddle homoclinic orbit bifurcation is supercritical, whichcorresponds to the (dis)appearance of a stable limit cycle.• If λ 1 + λ 2 > 0, then the saddle homoclinic orbit bifurcation is subcritical, whichcorresponds to the (dis)appearance of an unstable limit cycle.Thus, the saddle quantity plays the same role as the parameter a in the Andronov-Hopfbifurcation. The supercritical saddle homoclinic orbit bifurcation is more common inneuronal models than the subcritical one due to the reason explained in Sect. 6.3.6.Hence, we consider only the supercritical case below, and we drop the word “supercritical”for the sake of brevity.
Bifurcations 195stable manifoldoutin?saddleunstable manifoldFigure 6.27: Saddle homoclinic orbit bifurcationoccurs when the stable and unstable submanifolds ofthe saddle make a loop.A useful way to look at the bifurcation is to note that the saddle has one stableand one unstable direction on a phase plane. There are two orbits associated withthese directions, called the stable and unstable submanifolds, depicted in Fig. 6.27.Typically, the submanifolds miss each other, that is, the unstable submanifold goeseither inside or outside the stable one. This could happen for two different values ofthe bifurcation parameter. One can image that as the bifurcation parameter changescontinuously from one value to the other, the submanifolds join at some point and forma single homoclinic trajectory that starts and ends at the saddle.The saddle homoclinic orbit bifurcation is ubiquitous in neuronal models, and it caneasily be observed in the I Na,p +I K -model with fast K + conductance, as we illustratein Fig. 6.28. Let us start with I = 7 (top of Fig. 6.28) and decrease the bifurcationparameter I. First, there is only a stable limit cycle corresponding to periodic spikingactivity. When I decreases, a stable and an unstable equilibrium appear via saddlenodebifurcation (not shown in the figure), but the state of the model is still on thelimit cycle attractor. Further decrease of I moves the saddle equilibrium closer tothe limit cycle (case I = 4 in the figure), until the cycle becomes an infinite periodhomoclinic orbit to the saddle (case I ≈ 3.08), and then disappears (case I = 1). Atthis moment, the state of the system approaches the stable equilibrium, and the tonicspiking stops.Similarly to the fold limit cycle bifurcation, the saddle homoclinic orbit bifurcationexplains how the limit cycle attractor corresponding to periodic spiking behavior appearsand disappears. However, it does not explain the transition to periodic spikingbehavior. Indeed, when I = 4 in Fig. 6.28, the limit cycle attractor exists, yet theneuron may still be quiescent because its state may be at the stable node. The periodicspiking behavior appears only after external perturbations push the state of thesystem into the attraction domain of the limit cycle attractor, or I increases furtherand the stable node disappears via a saddle-node bifurcation.We can use linear theory to estimate the frequency of the limit cycle attractornear saddle homoclinic orbit bifurcation. Because the vector field is small near theequilibrium, the periodic trajectory slowly passes through a small neighborhood of theequilibrium, then quickly makes a rotation and returns to the neighborhood, as weillustrate in Fig. 6.29. Let T 1 denote the time required to make one rotation (dashed
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194 Bifurcationshomocl<strong>in</strong>ic orbithomocl<strong>in</strong>ic orbitsaddle homocl<strong>in</strong>ic orbitbifurcationsaddle-node on <strong>in</strong>variant circlebifurcationFigure 6.26: Two bifurcations <strong>in</strong>volv<strong>in</strong>g homocl<strong>in</strong>ic trajectories to an equilibrium.6.2.4 Homocl<strong>in</strong>icA limit cycle can appear or disappear via a saddle homocl<strong>in</strong>ic orbit bifurcation, asdepicted <strong>in</strong> Fig. 6.25. As the bifurcation parameter changes, the cycle becomes ahomocl<strong>in</strong>ic orbit to the saddle equilibrium, and its period becomes <strong>in</strong>f<strong>in</strong>ite. After thebifurcation, the cycle no longer exists. A necessary condition for such a bifurcation isthat the steady-state I-V relation is not monotonic.One should be careful to dist<strong>in</strong>guish the saddle homocl<strong>in</strong>ic orbit bifurcation fromthe saddle-node on <strong>in</strong>variant circle bifurcation depicted <strong>in</strong> Fig. 6.26. Indeed, it mightbe easy to confuse these bifurcations, s<strong>in</strong>ce both <strong>in</strong>volve an equilibrium and a largeamplitudehomocl<strong>in</strong>ic trajectory that becomes a limit cycle. The key difference is thatthe equilibrium is a saddle <strong>in</strong> the former and a saddle-node <strong>in</strong> the latter. The saddleequilibrium persists as the bifurcation parameter changes, whereas the saddle-nodeequilibrium disappears or bifurcates <strong>in</strong>to two po<strong>in</strong>ts, depend<strong>in</strong>g on the direction ofchange of the bifurcation parameter.Recall that a saddle on a plane has two real eigenvalues of opposite signs. Theirsum, λ 1 + λ 2 , is called the saddle quantity.• If λ 1 + λ 2 < 0, then the saddle homocl<strong>in</strong>ic orbit bifurcation is supercritical, whichcorresponds to the (dis)appearance of a stable limit cycle.• If λ 1 + λ 2 > 0, then the saddle homocl<strong>in</strong>ic orbit bifurcation is subcritical, whichcorresponds to the (dis)appearance of an unstable limit cycle.Thus, the saddle quantity plays the same role as the parameter a <strong>in</strong> the Andronov-Hopfbifurcation. The supercritical saddle homocl<strong>in</strong>ic orbit bifurcation is more common <strong>in</strong>neuronal models than the subcritical one due to the reason expla<strong>in</strong>ed <strong>in</strong> Sect. 6.3.6.Hence, we consider only the supercritical case below, and we drop the word “supercritical”for the sake of brevity.