Past exam of the mathematics course of the University of Cambridge / 2015 / ii / Paper 3 / 11E (Mathematical Biology)

Let $N=x+y$. Uninfected births and density limitation give $x(1-N)$, while horizontal transmission transfers individuals to the infected class at rate $\beta xy$. Infected births are infected by vertical transmission, and their net demographic growth is $y(1-N)-dy$. ConsequentlyThese give the stated equations. Adding them gives $\dot{N}=N(1-N)-dy$, displaying the additional mortality due to disease.

At the disease-free equilibrium point $(1,0)$, the linearization of a dynamical system of the infected equation is $\dot{y}=(\beta -d)y$. Thus exponential invasion occurs exactly when $\boxed{\beta >d}$. At equality there is no positive linear growth rate; the nonlinear boundary dynamics described below do not give sustained invasion.

Assume first $\beta >0$ and $d\ge 0$. The nonnegative equilibrium points areand, when both coordinates are nonnegative,The Jacobian matrix at $O$ has eigenvalues $1,1-d$, so $O$ cannot attract an initially positive uninfected population. At $U$ the eigenvalues are $-1,\beta -d$. At $I$, for $d<1$, they are $d(1+\beta )-\beta$ and $-(1-d)$. At an interior $C$, the equilibrium equations reduce the Jacobian matrix toThus the coexistence equilibrium is asymptotically stable precisely when $\beta /(1+\beta )<d<\beta$.

For positive initial populations these local results identify the global outcome. The relation $\dot{N}=N(1-N)-dy$ bounds all trajectories. With the Dulac function $1/(xy)$, the divergence of the rescaled vector field is $-1/y-1/x<0$, excluding interior periodic orbits. Boundary saddles cannot attract an interior trajectory except at the merging thresholds. The resulting long-term outcomes areAt $d=\beta /(1+\beta )$, $C$ merges into $I$; at $d=\beta$, it merges into $U$. The merged equilibria still attract interior trajectories, but convergence along the zero-eigenvalue direction is algebraic. For example at $d=\beta$, put $x=1+u$, $y=v$. The local centre manifold has $u=-(1+\beta )v+O(v^2)$, and $\dot{v}=-{\beta }^2{v}^2+O(v^3)$. At the other threshold, using $y=(1-d)+v$, $x=u$, gives $v=(\beta -1)u+O(u^2)$ and $\dot{u}=-{\beta }^2{u}^2+O(u^3)$.

The invariant axes also matter: $y=0$ leads to $U$ whenever $x(0)>0$; $x=0$ leads to $I$ for $d<1$ and to extinction for $d\ge 1$. At $d=1$, $I$ coincides with $O$. If $\beta =0$, disease declines for $d>0$ and any initially positive $x$ tends to $U$. If also $d=0$, every point $x+y=1$ is an equilibrium and the initial population proportions are preserved.

For $\beta >0$ and positive initial populations, summing the coordinates of the attracting equilibrium point gives the steady total populationIts minimum is $1/(1+\beta )$ at $d=\beta /(1+\beta )$. Small mortality leaves a fully infected population with only modest demographic loss. Raising mortality first reduces that population; after the threshold it suppresses infection sufficiently for the uninfected population to recover. Beyond the invasion threshold the disease cannot persist.