Dynamical Systems in Neuroscience:
Dynamical Systems in Neuroscience: Dynamical Systems in Neuroscience:
198 Bifurcationsfrequency (Hz)400300200200numericaltheoretical1001/ln10003.08 3.0903 3.1 3.2 3.3 3.4 3.5 3.6injected dc-current, IFigure 6.30: Frequency of spiking in the I Na,p +I K -model with parameters as inFig. 6.28 near a saddle homoclinic orbit bifurcation. Dots are numerical results; thecontinuous curve is ω(I) = 1000λ(I)/{− ln(0.2(I − 3.0814))}, where the eigenvalueλ(I) = 0.87 √ 4.51 − I was obtained from the normal form (6.3). The inset shows amagnified region near the bifurcation value I = 3.0814.big homoclinic orbitlimit cycleFigure 6.31: Big saddle homoclinic orbit bifurcation.Both, the saddle-node on invariant circle bifurcation and the saddle homoclinicorbit bifurcation result in spiking with decreasing frequency so that their frequencycurrent(F-I) curves go continuously to zero. The key difference is that the formerasymptotes as √ I − I b , whereas the latter as 1/ ln(I − I b ). The striking feature of thelogarithmic decay in Fig. 6.30 is that the frequency is greater than 100 Hz and thetheoretical curve does not seem to go to zero for all I except those in an infinitesimalneighborhood of the bifurcation value I b . Such a neighborhood is almost impossible tocatch numerically, let alone experimentally in real neurons.Many neuronal models, and even some cortical pyramidal neurons (see Fig. 7.42)exhibit a saddle homoclinic orbit bifurcation depicted in Fig. 6.31. Here, the unstablemanifold of a saddle returns to the saddle along the opposite side, thereby making abig loop, hence the name big saddle homoclinic orbit bifurcation. This kind of bifurcationoften occurs when an excitable system is near a codimension-2 Bogdanov-Takensbifurcation considered in Sect. 6.3.3, and it has the same properties as the “small”homoclinic orbit bifurcation considered above: it could be subcritical or supercritical,
Bifurcations 199heteroclinic orbitFigure 6.32: Heteroclinic orbit bifurcation does not change the existence of stability ofany equilibrium or periodic orbit.depending on the saddle quantity, it results in a logarithmic F-I curve, and it impliesthe co-existence of attractors. All methods of analysis of excitable systems near “small”saddle homoclinic orbit bifurcations can also be applied to the case in Fig. 6.31.6.3 Other Interesting CasesSaddle-node and Andronov-Hopf bifurcations of equilibria combined with fold limitcycle, homoclinic orbit bifurcation, and heteroclinic orbit bifurcation (see Fig. 6.32)exhaust all possible bifurcations of codimension-1 on a plane. These bifurcations canalso occur in higher-dimensional systems. Below we discuss additional codimension-1bifurcations in three-dimensional phase space, and then we consider some codimension-2 bifurcations that play an important role in neuronal dynamics. The first-time readermay read only Sect. 6.3.6 and skip the rest.6.3.1 Three-dimensional phase spaceSo far we considered four bifurcations of equilibria and four bifurcations of limit cycleson a phase plane. The same eight bifurcations can appear in multi-dimensional systems.Below we briefly discuss the new kinds of bifurcations that are possible in a threedimensionalphase space but cannot occur on a plane.First, there are no new bifurcations of equilibria in multi-dimensional phase space.Indeed, what could possibly happen with the Jacobian matrix of an equilibrium ofa multi-dimensional dynamical system? A simple zero eigenvalue would result ina saddle-node bifurcation, and a simple pair of purely imaginary complex-conjugateeigenvalues would result in an Andronov-Hopf bifurcation. Both are exactly the sameas in the lower-dimensional systems considered before. Thus, adding dimensions to adynamical system does not create new possibilities for bifurcations of equilibria.In contrast, adding the third dimension to a planar dynamical system creates newpossibilities for bifurcations of limit cycles, some of which are depicted in Fig. 6.33.Below we briefly describe these bifurcations.The saddle-focus homoclinic orbit bifurcation in Fig. 6.33 is similar to the saddlehomoclinic orbit bifurcation considered in Sect. 6.2.4 except that the equilibrium hasa pair of complex-conjugate eigenvalues and a non-zero real eigenvalue. The homoclinicorbit originates in the subspace spanned by the eigenvector corresponding to
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198 Bifurcationsfrequency (Hz)400300200200numericaltheoretical1001/ln10003.08 3.0903 3.1 3.2 3.3 3.4 3.5 3.6<strong>in</strong>jected dc-current, IFigure 6.30: Frequency of spik<strong>in</strong>g <strong>in</strong> the I Na,p +I K -model with parameters as <strong>in</strong>Fig. 6.28 near a saddle homocl<strong>in</strong>ic orbit bifurcation. Dots are numerical results; thecont<strong>in</strong>uous curve is ω(I) = 1000λ(I)/{− ln(0.2(I − 3.0814))}, where the eigenvalueλ(I) = 0.87 √ 4.51 − I was obta<strong>in</strong>ed from the normal form (6.3). The <strong>in</strong>set shows amagnified region near the bifurcation value I = 3.0814.big homocl<strong>in</strong>ic orbitlimit cycleFigure 6.31: Big saddle homocl<strong>in</strong>ic orbit bifurcation.Both, the saddle-node on <strong>in</strong>variant circle bifurcation and the saddle homocl<strong>in</strong>icorbit bifurcation result <strong>in</strong> spik<strong>in</strong>g with decreas<strong>in</strong>g frequency so that their frequencycurrent(F-I) curves go cont<strong>in</strong>uously to zero. The key difference is that the formerasymptotes as √ I − I b , whereas the latter as 1/ ln(I − I b ). The strik<strong>in</strong>g feature of thelogarithmic decay <strong>in</strong> Fig. 6.30 is that the frequency is greater than 100 Hz and thetheoretical curve does not seem to go to zero for all I except those <strong>in</strong> an <strong>in</strong>f<strong>in</strong>itesimalneighborhood of the bifurcation value I b . Such a neighborhood is almost impossible tocatch numerically, let alone experimentally <strong>in</strong> real neurons.Many neuronal models, and even some cortical pyramidal neurons (see Fig. 7.42)exhibit a saddle homocl<strong>in</strong>ic orbit bifurcation depicted <strong>in</strong> Fig. 6.31. Here, the unstablemanifold of a saddle returns to the saddle along the opposite side, thereby mak<strong>in</strong>g abig loop, hence the name big saddle homocl<strong>in</strong>ic orbit bifurcation. This k<strong>in</strong>d of bifurcationoften occurs when an excitable system is near a codimension-2 Bogdanov-Takensbifurcation considered <strong>in</strong> Sect. 6.3.3, and it has the same properties as the “small”homocl<strong>in</strong>ic orbit bifurcation considered above: it could be subcritical or supercritical,