Past exam of the mathematical physiology course of the University of Oxford / 2011 / 1 / b / ii / Solution

The voltage nullcline is $g(v,n)=I^{\ast }$ and the recovery nullcline is $n=n_{\infty }(v)$. In the relevant region $g_n>0$, so below the voltage curve $\dot{v}>0$ and above it $\dot{v}<0$. Below the recovery curve $\dot{n}>0$, while above it $\dot{n}<0$.

For small $ϵ$, the outer voltage branches with $g_v>0$ attract fast voltage motion; the middle branch with $g_v<0$ repels it. Arrange the intersections so the equilibrium is on that middle branch, with recovery increasing along the high-voltage attracting branch and decreasing along the low-voltage attracting branch. The trajectory then drifts towards a fold, jumps rapidly to the other attracting branch at nearly constant $n$, and repeats. This is a relaxation oscillation.

The local linear stability calculation supports the construction. At an equilibrium,A sufficiently steep recovery curve gives positive determinant, while $g_v<-ϵ$ gives positive trace. Hence the equilibrium is a repeller. The bounded physical rectangle can be chosen forward invariant, and a small neighborhood of the repeller removed to give a trapping annulus. The Poincaré-Bendixson theorem then supplies a periodic orbit. These conditions can be realized because the recovery function and the applied current control the intersection and its slope; oscillation is possible, not forced for every choice.

For the illustration, ${\gamma }_K=0.1$, $H=0.8$, $v_K=0.2$, $ϵ=0.01$, $I^{\ast }=0$, $M(v)=(1+e^{-25(v-0.2)}{)}^{-1}$ and $n_{\infty }(v)=0.8/(1+e^{-20v})$. The rectangle $[-0.2,1]\times [0,0.8]$ has inward-pointing flow, and the displayed numerical trajectory realizes the folded-branch mechanism.