6. Birth and Death Processes 6.1 Pure Birth Process (Yule-Furry Process) 6.2 Generalizations 6.3 Birth and Death Processes 6.4 Relationship to Markov Chains 6.5 Linear Birth and Death Processes 230 6.1 Pure Birth Process (Yule-Furry Process) Example. Consider cells which reproduce according to the following rules: i. A cell present at time t has probability λh + o(h) of splitting in two in the interval (t, t + h) ii. This probability is independent of age. iii. Events between different cells are independent Time> 231 Non-Probablistic Analysis n(t) = no. of cells at time t ⇒ λn(t)∆(t) births occur in (t, t + ∆t) where λ = birth rate per single cell. n(t + ∆t) = n(t) + n(t)λ∆t n(t + ∆t) − n(t) ∆t → n (t) = n(t)λ or n (t) n(t) = d dt log n(t) = λ log n(t) = λt + c n(t) = Keλt , n(0) = n0 n(t) = n0eλt 232 Probabilistic Analysis N(t) = no. of cells at time t P{N(t) = n} = Pn(t) Prob. of birth in (t, t + h) if {N(t) = n} = nλh + o(h) Pn(t + h) = Pn(t)(1 − nλh + o(h)) + Pn−1(t)((n − 1)λh + o(h)) Pn(t + h) − Pn(t) = −nλhPn(t) + Pn−1(t)(n − 1)λh + o(h) Pn(t + h) − Pn(t) h = −nλPn(t) + Pn−1(t)(n − 1)λ + o(h) as h → 0 Pn(t) = −nλPn(t) + (n − 1)λPn−1(t) Initial condition Pn0 (0) = P{N(0) = n0} = 1 233 Pn(t) = −nλPn(t) + (n − 1)λPn−1(t); Pn0 (0) = 1 Solution: (1) Pn(t) = n − 1 n − n0 e−λn0t (1 − e−λt )n−n0 n = n0, n0 + 1, . . . Solution is negative binomial distribution; i.e. Probability of obtaining exactly n0 successes in n trials. Suppose p = prob. of success and q = 1 − p = prob. of failure. Then in first (n − 1) trials results in (n0 − 1) successes and (n − n0) failures followed by success on nth trial; i.e. (2) n − 1 n0 − 1 pn0−1 qn−n0 · p = n − 1 n − n0 pn0 qn−n0 n = n0, n0 + 1, ... If p = e−λt and q = 1 − e−λt ⇒ (2) is same as (1). 234 Yule studied this process in connection with theory of evolution; i.e. population consists of the species within a genus and creation of new element is due to mutations. Neglects probability of species dying out and size of species. Furry used same model for radioactive transmutations. Notes on Negative Binomial Distribution The geometric distribution is defined as the number of trials to achieve one success for a series of Bernoulli trials; i.e. Geometric Distribution: P{N = n} = pqn−1 , n = 1, 2, . . . N is number of trials for 1 success φN (s) = E(e−sN ) = p ∞ n=1 e−sn qn−1 = p q ∞ n=1 (e−s q)n . 235 But ∞ n=1 (e−s q)n = e−s q/(1 − e−s q) φN (s) = p q · e−s q 1 − e−sq = pe−s 1 − e−sq = pz 1 − qz if z = e−s φN (z) = pz(1 − qz)−1 φN (z) = p{(1 − qz)−1 + z(1 − qz)−2 q} φN (1) = p{(1 − q)−1 + q(1 − q)−2 } = 1 + q p = p + q p = 1 p Similarly φN (1) = 2 p2 − 2 p ⇒ V (n) = 1 p2 − 1 p 236 Theorem: If Ni (i = 1, 2, . . . , n0) are iid geometric random variables with parameter p, then N = N1 + N2 + . . . + Nn0 is a negative binomial distribution having generating function φN (z) = pz 1 − qz n0 , z = e−s ˙.. E(N) = n0/p, V (N) = n0 1 p2 − 1 p If p = e−λt and N(t) is a pure birth process E[N(t)] = n0eλt V [N(t)] = n0[e2λt − eλt ] 237 6.2 Generalization In Poisson Process, the prob. of a change during (t, t + h) is independent of number of changes in (0, t). Assume instead that if n changes occur in (0, t), the probability of new change to n + 1 in (t, t + h) is λnh + o(h). The probability of more than one change is o(h). Then Pn(t + h) = Pn(t)(1 − λnh) + Pn−1(t)λn−1h + o(h), n = 0 P0(t + h) = P0(t)(1 − λ0h) + o(h) ⇒ Pn(t) = −λnPn(t) + λn−1Pn−1(t) P0(t) = −λ0P0(t). Equations can be solved recursively with P0(t) = P0(0)e−λ0t . 238 If the initial condition is Pn0 (0) = 1, then the resulting equations are: Pn(t) = −λnPn(t) + λn−1Pn−1(t), n > n0 Pn0 (t) = −λn0 Pn0 (t) Pure birth process assumed λn = nλ. Change of Language If n transitions take place during (0, t), we may refer to the process as being in state En. Changes occur En → En+1 → En+2 → . . . for the pure birth process. 239 6.3 Birth and Death Processes Consider transitions En → En−1 as well as En → En+1 if n ≥ 1. If n = 0, we only allow E0 → E1. Assume that if the process at time t is in En, then during (t, t + h) the transitions En → En+1 have prob. λnh + o(h), En → En−1 have prob. µnh + o(h) and Prob. more than 1 change occurs = o(h) Pn(t + h) = Pn(t){1 − λnh − µnh} + Pn−1(t){λn−1h} + Pn+1(t){µn+1h} + o(h) ⇒ Pn(t) = −(λn + µn)Pn(t) + λn−1Pn−1(t) + µn+1Pn+1(t) 240 For n = 0 P0(t + h) = P0(t){1 − λ0h} + P1(t)µ1h + o(h) ⇒ P0(t) = −λ0P0(t) + µ1P1(t) If the initial conditions Pn0 (0) = 1 in which case 0 in above is replaced by n0. If λ0 = 0, then E0 → E1 is impossible and E0 is an absorbing state. If λ0 = 0, then P0(t) = µ1P1(t) ≥ 0 so that P0(t) increases monotonically. Note: lim t→∞ P0(t) = P0(∞) = Probability of being absorbed. 241 P0(t) = −λ0P0(t) + µ1P1(t) Pn(t) = −(λn + µn)Pn(t) + λn−1Pn−1(t) + µn+1Pn+1(t) As t → ∞, Pn(t) → Pn (limit) hence P0(t) = 0 for large t and Pn(t) = 0 for large t. Therefore 0 = −λ0P0 + µ1P1 ⇒ P1 = λ0 µ1 P0 0 = −(λ1 + µ1)P1 + λ0P0 + µ2P2 ⇒ P2 = λ0λ1 µ2µ1 P0 ⇒ P3 = λ0λ1λ2 µ1µ2µ3 P0 etc. Note that dependence on initial conditions has disappeared. 242 6.4 Relation to Markov Chains Define for t → ∞ P(En+1|En) = Prob. of transition En → En+1 = Prob. of going to En+1 conditional on being in En. Similarly define P(En−1|En). P(En+1|En) ∝ λn, P(En−1|En) ∝ µn ⇒ P(En+1|En) = λn λn + µn , P(En−1|En) = µn λn + µn Same conditional probabilities hold if it is given that a transition will take place during (t, t + h) conditional on being in En. 243 6.5 Linear birth and death processes λn = nλ , µn = nµ ⇒ P0(t) = µP1(t) Pn(t) = −(λ + µ)nPn(t) + λ(n − 1)Pn−1(t) + µ(n + 1)Pn+1(t) Steady state behavior is characterized by lim t→∞ P0(t) = 0 ⇒ P1(∞) = 0 Similarly as t → ∞ Pn(∞) = 0 If P0(∞) = 1 ⇒ Probability of ultimate extinction is 1. If P0(∞) = P0 < 1, the relations P1 = P2 = P3 . . . = 0 imply with prob. 1 − P0 the population can increase without bounds. The population must either die out or increase indefinitely. 244 Pn(t) = −(λ + µ)nPn(t) + λ(n − 1)P(n−1)(t) + µ(n + 1)Pn+1(t) Define Mean by M(t) = ∞ n=1 nPn(t) and consider M (t) = ∞ 1 nPn(t). M (t) = −(λ + µ) ∞ 1 n2 Pn(t) + λ ∞ 1 (n − 1)n P(n−1)(t) + µ ∞ 1 (n + 1)nPn+1(t) Write (n − 1)n = (n − 1)2 + (n − 1), (n + 1)n = (n + 1)2 − (n + 1) 245 M (t) = − (λ + µ) ∞ 1 n2 Pn(t) + λ ∞ 1 (n − 1)2 Pn−1(t) + µ ∞ 1 (n + 1)2 Pn+1(t) + 1 · P1(t) + λ ∞ 1 (n − 1)Pn−1(t) − µ ∞ 1 (n + 1)Pn+1(t) + P1(t) ⇒ M (t) = λ ∞ 1 nPn(t) − µ ∞ 1 nPn(t) = (λ − µ)M(t) M(t) = n0e(λ−µ)t if Pn0 (0) = 1 246 M(t) = n0e(λ−µ)t M(t) → 0 or ∞ depending on λ < µ or λ > µ . Similarly if M2(t) = ∞ 1 n2 Pn(t) one can show M2(t) = 2(λ − µ)M2(t) + (λ + µ)M(t) and when λ > µ, the variance is n0e2(λ−µ)t 1 − e(µ−λ)t λ + µ λ − µ 247