A detailed probabilistic treatment is given of a birth-and-death process proposed by Williams (1969) in which the elements of the process bear up to s inanimate marks. Equations for the second-order moments of the process, and approximate marginal univariate solutions, are derived. The exact bivariate solution is given for the case s = 1. For general s the variance of the mark population is also derived. MARKED BIRTH-AND-DEATH PROCESS; MULTIVARIATE LAGRANGE DIFFERENTIAL EQUATIONS; MULTITYPE BRANCHING PROCESS Introduction In a recent paper Williams (1969) has considered the following process. We have a population of elements developing as a linear birth-and-death process and each element bears a number of inanimate marks. When a marked element dies then it and its marks are removed from the population; when a marked element divides, its marks segregate independently and without loss, each with probability I into one of the two daughter elements. It is assumed that marking an element does not affect its birth and death rates (cf. Clifford and Sudbury (1972)). Williams' process was proposed as a model of an experiment conducted on bacteria (the elements) infected by phage (the inanimate marks), by Meynell (1959). (The bacteria are lysogenic with respect to the phage, which means that the phage do not interfere with the behaviour of the bacteria.) His object was to use the marking process in order to separate the birth and death rates of the bacteria; if we denote birth and death rates by A and p respectively, then the mean-value measures on the bacterial population (assumed to develop as a linear birth-and-death process) will estimate the difference (A p). As the marks are inanimate, however, they will only be affected by p and so mean-value measures on the mark population enable us to estimate p (see Williams (1969)). The papers of Meynell and Williams taken together provide a most interesting example of a virological experiment and the probability model of that experiment. Received in revised form 10 August 1973 Research sponsored by a Science Research Council grant and a World Health Organisation research traineeship. 423 This content downloaded from 207.46.13.180 on Thu, 08 Sep 2016 05:01:40 UTC All use subject to http://about.jstor.org/terms 424 BYRON J. T. MORGAN Williams considered the mean-value solution of his model; this was adequate for Meynell's experiment in which population sizes were large. Here we shall give a fuller probabilistic statement of the model and make some contributions to its solution. Two differences from Williams' work lie in the adoption of an upper bound, s, to the number of marks any element can have, and the assumption (except where stated) of birth and death rates independent of time. We 'shall assume general initial conditions which, in Meynell's experiment, correspond to the distribution of phage over bacteria once these have been initially exposed to a phage population. Williams took this distribution to be Poisson; the distribution of phage over bacteria has, however, been the subject of much probabilistic study (e.g., see Gani (1965) and Morgan (1971)). Finally, let us note that the value s = I, on which we shall particularly concentrate, probably gives the most realistic description of Meynell's experiment. 2. The probability model The model is a continuous-time Markov process with discrete states. Let N,(t) be a random variable denoting the number of elements with i marks at time t, i = 0, 1,2, ..., s. Let n = (n0,n1, l , n.., ), Iu = (Uo, u1, *,us), N(t) = (No(t), N(t), --,N,(t)), and suppose N,(O) = ai, i = 0, 1,2, --, s. Let P(n; t) = P(N(t) = n), then corresponding to death we have the transitions, (No, ..., Ni, ---, N,) (No, --, Ni 1, , N,) at probability rate pNi, i = 0, 1,2, ,s, while corresponding to birth we have the following transitions (No, N,) ,(No + 1, .., N,) at probability rate, 2 =1 N 2-' +,ANo, (No, , Ni ,N2i,..., X (No, Ni + 2, , N2iI, at probability rate, This content downloaded from 207.46.13.180 on Thu, 08 Sep 2016 05:01:40 UTC All use subject to http://about.jstor.org/terms On the distribution of inanimate marks over a linear birth-and-death process 425 where we use [is] to denote the integral part of is, and (No, " , Ni, * * *, Nk i Nk, "', Ns) -*(No, ..., Nj + 1, ..., Nk-i + 1, ..., Nj 1, ..., Nl) at probability rate 2N 2-k' i = 1,2,...,C[(k 1)], k = 3,4,...,s (smaller k give transitions already considered above). It is readily verified that the sum of the transition rates is (A + yp) X = oN,, as required in the overall birth-and-death process. Introducing the operator Di, which increases the argument ni of a function of n by 1, we then deduce that the Kolmogorov forward equations (for all possible n) are, d S S (1) +(n; t) P(n; t)(A+ t)1 (n2 + D,P(n;t)(n +1) dt ==0 1=0