Paper 3, Section II, C

Mathematical Biology
Part II, 2018

Consider fluctuations of a population described by the vector x=(x1,x2,,xN)\mathbf{x}=\left(x_{1}, x_{2}, \ldots, x_{N}\right). The probability of the state x\mathbf{x} at time t,P(x,t)t, P(\mathbf{x}, t), obeys the multivariate Fokker-Planck equation

Pt=xi(Ai(x)P)+122xixj(Bij(x)P),\frac{\partial P}{\partial t}=-\frac{\partial}{\partial x_{i}}\left(A_{i}(\mathbf{x}) P\right)+\frac{1}{2} \frac{\partial^{2}}{\partial x_{i} \partial x_{j}}\left(B_{i j}(\mathbf{x}) P\right),

where P=P(x,t),AiP=P(\mathbf{x}, t), A_{i} is a drift vector and BijB_{i j} is a symmetric positive-definite diffusion matrix, and the summation convention is used throughout.

(a) Show that the Fokker-Planck equation can be expressed as a continuity equation

Pt+J=0\frac{\partial P}{\partial t}+\nabla \cdot \mathbf{J}=0

for some choice of probability flux J\mathbf{J} which you should determine explicitly. Here, =(x1,x2,,xN)\nabla=\left(\frac{\partial}{\partial x_{1}}, \frac{\partial}{\partial x_{2}}, \ldots, \frac{\partial}{\partial x_{N}}\right) denotes the gradient operator.

(b) Show that the above implies that an initially normalised probability distribution remains normalised,

P(x,t)dV=1\int P(\mathbf{x}, t) d V=1

at all times, where the volume element dV=dx1dx2dxNd V=d x_{1} d x_{2} \ldots d x_{N}.

(c) Show that the first two moments of the probability distribution obey

ddtxk=Akddtxkxl=xlAk+xkAl+Bkl\begin{aligned} \frac{d}{d t}\left\langle x_{k}\right\rangle &=\left\langle A_{k}\right\rangle \\ \frac{d}{d t}\left\langle x_{k} x_{l}\right\rangle &=\left\langle x_{l} A_{k}+x_{k} A_{l}+B_{k l}\right\rangle \end{aligned}

(d) Now consider small fluctuations with zero mean, and assume that it is possible to linearise the drift vector and the diffusion matrix as Ai(x)=aijxjA_{i}(\mathbf{x})=a_{i j} x_{j} and Bij(x)=bijB_{i j}(\mathbf{x})=b_{i j} where aija_{i j} has real negative eigenvalues and bijb_{i j} is a symmetric positive-definite matrix. Express the probability flux in terms of the matrices aija_{i j} and bijb_{i j} and assume that it vanishes in the stationary state.

(e) Hence show that the multivariate normal distribution,

P(x)=1Zexp(12Dijxixj)P(\mathbf{x})=\frac{1}{Z} \exp \left(-\frac{1}{2} D_{i j} x_{i} x_{j}\right)

where ZZ is a normalisation and DijD_{i j} is symmetric, is a solution of the linearised FokkerPlanck equation in the stationary state, and obtain an equation that relates DijD_{i j} to the matrices aija_{i j} and bijb_{i j}.

(f) Show that the inverse of the matrix DijD_{i j} is the matrix of covariances Cij=xixjC_{i j}=\left\langle x_{i} x_{j}\right\rangle and obtain an equation relating CijC_{i j} to the matrices aija_{i j} and bijb_{i j}.