Past exam of the mathematics course of the University of Cambridge / 2015 / ii / Paper 4 / 5E (Mathematical Biology)

Put $x=1+y$ and retain linear terms. The delay differential equation is $\dot{y}(t)=-\alpha y(t-T)$, with characteristic equation of a delay differential equation $\lambda +\alpha e^{-\lambda T}=0$. Write $z=\lambda T$ and $q=\alpha T$. If a root has $\mathrm{Re}z\ge 0$, then $|z|=q{e}^{-\mathrm{Re}z}\le q$. Writing $z=a+ib$ gives $a=-q{e}^{-a}\cos b$. For $q<\pi /2$, $|b|<\pi /2$ and the right side is negative, contradicting $a\ge 0$. All characteristic roots therefore lie in the left half-plane.

An imaginary root $z=ib$ requires $\cos b=0$ and $b=q\sin b$. The first crossing is $q=\pi /2$, $z=\pm i\pi /2$. Implicit differentiation gives $dz/dq=-e^{-z}/(1+z)$, whose real part at the positive crossing is $(\pi /2)/(1+{\pi }^2/4)>0$. The later imaginary crossings occur at $q=\pi /2+2\pi k$ and also enter the right half-plane. Thus the equilibrium is locally exponentially asymptotically stable whenand unstable when $\alpha T>\pi /2$. At equality the linearized equation has neutral oscillatory modes; it is the Hopf bifurcation threshold. If stability is meant for the nonlinear equation itself, the borderline can also be resolved. In dimensionless time $s=t/T$, let $q_0=\pi /2$. A small center oscillation has the form $y=R\cos \theta +R^2(\cos 2\theta /5+\sin 2\theta /10)+O(R^3)$. Substitution at quadratic order gives these second-harmonic coefficients. At cubic order the fundamental forcing is $q_0{R}^3(\cos \theta /20+3\sin \theta /20)$. Projection onto the center mode divides the complex forcing by the characteristic derivative $1+i{q}_0$, yielding the Hopf normal form amplitude equationIts cubic coefficient is negative, so the equilibrium at equality is locally asymptotically stable, with algebraic rather than exponential decay. Thus the usual strictly decaying linear condition is $\alpha T<\pi /2$, while nonlinear local asymptotic stability includes $\alpha T=\pi /2$. The purely imaginary linear modes alone would not justify that borderline conclusion.

The delayed positive term models recruitment depending on the population a time $T$ earlier; the quadratic negative term represents instantaneous density-dependent losses. Thus this delay differential equation delays recruitment rather than the strength of crowding acting on the present population.

With $x=1+y$, the linearized equation is $\dot{y}=\alpha y(t-T)-2\alpha y(t)$. Its characteristic equation of a delay differential equation is $\lambda +2\alpha =\alpha e^{-\lambda T}$. A root with nonnegative real part would satisfy $|\lambda +2\alpha |\ge 2\alpha$, whereas $|\alpha e^{-\lambda T}|\le \alpha$. This is impossible. Consequently the equilibrium is locally exponentially asymptotically stable for every $\alpha >0$ and $T>0$. There is no upper bound on $\alpha T$ in this model.