Past exam of the further mathematical biology course of the University of Oxford / 2023 / 2 / a / ii

At an equilibrium $(v_{\ast },k{v}_{\ast })$, set $p=f^{\prime }(v_{\ast })$. The Jacobian matrix, its trace and its determinant areThe eigenvalues and their discriminant areThus the complete local linear stability test isIn the last two cases, $\mathcal{D}>0$ gives a node, while $\mathcal{D}<0$ gives a focus; the sign of $T$ distinguishes inward and outward trajectories. At $\mathcal{D}=0$, a nonzero repeated eigenvalue gives a degenerate node. At $p=\gamma <k$ the eigenvalues are purely imaginary, so linear stability analysis alone does not decide asymptotic stability; with the required crossing and nondegeneracy this is a Hopf bifurcation. At $p=k$ there is a zero eigenvalue, requiring a nonlinear test and generically marking the saddle-node bifurcation in the preceding answer.

For a unique simple equilibrium, $p<k$, so it can be a sink equilibrium or a source equilibrium. The number of equilibria alone does not decide stability. If $k\le \gamma$, the unique simple equilibrium is always a sink equilibrium. If $\gamma <k$, it can become a source equilibrium where $\gamma <p<k$. The same conclusion follows particularly simply if $\gamma >M$: the trace is negative at every nonsaddle equilibrium.

For three simple equilibria ordered by their $v$ coordinates, the middle one has $p>k$ and is a saddle equilibrium; the two outer ones have $p<k$. When both outer equilibria are sink equilibria, each has an attracting neighborhood, giving bistability. In the usual two-basin portrait, the stable manifold of the saddle equilibrium separates their basins. Each branch of its unstable manifold initially leaves towards a different side; its eventual destination requires a global argument. If $k\le \gamma$, both outer equilibria must be sink equilibria. If $\gamma <k$, either or both outer equilibria can instead be source equilibria; the same test $p\lessgtr \gamma$ applies separately to each. The three lower panels illustrate all three possibilities up to exchanging left and right.

The arrows in the phase plane follow directly from the nullclines: below $w=f(v)+I$ the horizontal component points right, and above it left; below $w=kv$ the vertical component points up, and above it down. Near a stable focus trajectories spiral inward; near an unstable focus they spiral outward. Near a saddle equilibrium they approach along its stable manifold and leave along its unstable manifold. These directions, together with the local tests, determine the sketches.

There is also a useful global conclusion when the unique equilibrium is a source equilibrium. Withthe leading term in $v$ is $-\beta v^4$, so sufficiently large ellipses are trapping regions. A small repelling neighborhood of the hyperbolic source equilibrium can be removed to form an invariant annulus. The Poincaré-Bendixson theorem then guarantees at least one periodic orbit in that annulus. The numerical sketch illustrates the resulting sustained oscillation; this argument does not assert uniqueness of a limit cycle. A stable equilibrium can instead represent a resting membrane potential, with perturbations producing damped relaxation; near a threshold, a sufficiently large perturbation can produce a large excursion before recovery. Global basin boundaries and coexistence of periodic orbits need more information than the number and local stability of equilibria.