scholarly journals Wald’s martingale and the conditional distributions of absorption time in the Moran process

Author(s):  
Travis Monk ◽  
André van Schaik

Many models of evolution are stochastic processes, where some quantity of interest fluctuates randomly in time. One classic example is the Moranbirth–death process, where that quantity is the number of mutants in a population. In such processes, we are often interested in their absorption (i.e. fixation) probabilities and the conditional distributions of absorption time. Those conditional time distributions can be very difficult to calculate, even for relatively simple processes like the Moran birth–death model. Instead of considering the time to absorption, we consider a closely related quantity: the number of mutant population size changes before absorption. We use Wald’s martingale to obtain the conditional characteristic functions of that quantity in the Moran process. Our expressions are novel, analytical and exact, and their parameter dependence is explicit. We use our results to approximate the conditional characteristic functions of absorption time. We state the conditions under which that approximation is particularly accurate. Martingales are an elegant framework to solve principal problems of evolutionary stochastic processes. They do not require us to evaluate recursion relations, so when they are applicable, we can quickly and tractably obtain absorption probabilities and times of evolutionary models.

2020 ◽  
Author(s):  
Travis Monk ◽  
André van Schaik

AbstractMany models of evolution are stochastic processes, where some quantity of interest fluctuates randomly in time. One classic example is the Moran birth-death process, where that quantity is the number of mutants in a population. In such processes we are often interested in their absorption (i.e. fixation) probabilities, and the conditional distributions of absorption time. Those conditional time distributions can be very difficult to calculate, even for relatively simple processes like the Moran birth-death model. Instead of considering the time to absorption, we consider a closely-related quantity: the number of mutant population size changes before absorption. We use Wald’s martingale to obtain the conditional characteristic functions of that quantity in the Moran process. Our expressions are novel, analytical, and exact. The parameter dependence of the characteristic functions is explicit, so it is easy to explore their properties in parameter space. We also use them to approximate the conditional characteristic functions of absorption time. We state the conditions under which that approximation is particularly accurate. Martingales are an elegant framework to solve principal problems of evolutionary stochastic processes. They do not require us to evaluate recursion relations, so we can quickly and tractably obtain absorption probabilities and times of evolutionary stochastic processes.Author summaryThe Moran process is a probabilistic birth-death model of evolution. A mutant is introduced to an indigenous population, and we randomly choose organisms to live or die on subsequent time steps. Our goals are to calculate the probabilities that the mutant eventually dominates the population or goes extinct, and the distribution of time it requires to do so. The conditional distributions of time are difficult to obtain for the Moran process, so we consider a slightly different but related problem. We instead calculate the conditional distributions of the number of times that the mutant population size changes before it dominates the population or goes extinct. We use a martingale identified by Abraham Wald to obtain elegant and exact expressions for those distributions. We then use them to approximate conditional time distributions, and we show when that approximation is accurate. Our analysis outlines the basic concepts martingales and demonstrates why they are a formidable tool for studying probabilistic evolutionary models such as the Moran process.


2021 ◽  
Vol 8 (10) ◽  
Author(s):  
Travis Monk ◽  
André van Schaik

Evolutionary graph theory investigates how spatial constraints affect processes that model evolutionary selection, e.g. the Moran process. Its principal goals are to find the fixation probability and the conditional distributions of fixation time, and show how they are affected by different graphs that impose spatial constraints. Fixation probabilities have generated significant attention, but much less is known about the conditional time distributions, even for simple graphs. Those conditional time distributions are difficult to calculate, so we consider a close proxy to it: the number of times the mutant population size changes before absorption. We employ martingales to obtain the conditional characteristic functions (CCFs) of that proxy for the Moran process on the complete bipartite graph. We consider the Moran process on the complete bipartite graph as an absorbing random walk in two dimensions. We then extend Wald’s martingale approach to sequential analysis from one dimension to two. Our expressions for the CCFs are novel, compact, exact, and their parameter dependence is explicit. We show that our CCFs closely approximate those of absorption time. Martingales provide an elegant framework to solve principal problems of evolutionary graph theory. It should be possible to extend our analysis to more complex graphs than we show here.


1980 ◽  
Vol 17 (02) ◽  
pp. 363-372 ◽  
Author(s):  
C. Park ◽  
F. J. Schuurmann

Let {W(t), 0≦t<∞} be the standard Wiener process. The computation schemes developed in the past are not computationally efficient for the absorption probabilities of the type P{sup0≦t≦T W(t) − f(t) ≧ 0} when either T is large or f(0) > 0 is small. This paper gives an efficient and accurate algorithm to compute such probabilities, and some applications to other Gaussian stochastic processes are discussed.


2006 ◽  
Vol 43 (4) ◽  
pp. 1107-1118 ◽  
Author(s):  
Alexander D. Kolesnik

We consider the random motion of a particle that moves with constant finite speed in the space ℝ4 and, at Poisson-distributed times, changes its direction with uniform law on the unit four-sphere. For the particle's position, X(t) = (X1(t), X2(t), X3(t), X4(t)), t > 0, we obtain the explicit forms of the conditional characteristic functions and conditional distributions when the number of changes of directions is fixed. From this we derive the explicit probability law, f(x, t), x ∈ ℝ4, t ≥ 0, of X(t). We also show that, under the Kac condition on the speed of the motion and the intensity of the switching Poisson process, the density, p(x,t), of the absolutely continuous component of f(x,t) tends to the transition density of the four-dimensional Brownian motion with zero drift and infinitesimal variance σ2 = ½.


1983 ◽  
Vol 20 (2) ◽  
pp. 305-321 ◽  
Author(s):  
J. Grasman ◽  
D. Ludwig

Asymptotic approximations and numerical computations are used to estimate the accuracy of the diffusion approximation for the expected time to extinction for some stochastic processes. The results differ for processes with a continuant transition matrix (e.g. a birth and death process), and those with a noncontinuant transition matrix (e.g. a non-linear branching process). In the latter case, the diffusion equation does not hold near the point of exit. Consequently, high-order corrections do not result in substantial improvement over the diffusion approximation.


2005 ◽  
Vol 42 (01) ◽  
pp. 185-198 ◽  
Author(s):  
Erik A. Van Doorn ◽  
Alexander I. Zeifman

We study birth-death processes on the nonnegative integers, where {1, 2,…} is an irreducible class and 0 an absorbing state, with the additional feature that a transition to state 0 may occur from any state. We give a condition for absorption (extinction) to be certain and obtain the eventual absorption probabilities when absorption is not certain. We also study the rate of convergence, as t → ∞, of the probability of absorption at time t, and relate it to the common rate of convergence of the transition probabilities that do not involve state 0. Finally, we derive upper and lower bounds for the probability of absorption at time t by applying a technique that involves the logarithmic norm of an appropriately defined operator.


2003 ◽  
Vol 40 (3) ◽  
pp. 821-825 ◽  
Author(s):  
Damian Clancy ◽  
Philip K. Pollett

For Markov processes on the positive integers with the origin as an absorbing state, Ferrari, Kesten, Martínez and Picco studied the existence of quasi-stationary and limiting conditional distributions by characterizing quasi-stationary distributions as fixed points of a transformation Φ on the space of probability distributions on {1, 2, …}. In the case of a birth–death process, the components of Φ(ν) can be written down explicitly for any given distribution ν. Using this explicit representation, we will show that Φ preserves likelihood ratio ordering between distributions. A conjecture of Kryscio and Lefèvre concerning the quasi-stationary distribution of the SIS logistic epidemic follows as a corollary.


Author(s):  
Burton Voorhees ◽  
Alex Murray

The problem of finding birth–death fixation probabilities for configurations of normal and mutants on an N -vertex graph is formulated in terms of a Markov process on the 2 N -dimensional state space of possible configurations. Upper and lower bounds on the fixation probability after any given number of iterations of the birth–death process are derived in terms of the transition matrix of this process. Consideration is then specialized to a family of graphs called circular flows, and we present a summation formula for the complete bipartite graph, giving the fixation probability for an arbitrary configuration of mutants in terms of a weighted sum of the single-vertex fixation probabilities. This also yields a closed-form solution for the fixation probability of bipartite graphs. Three entropy measures are introduced, providing information about graph structure. Finally, a number of examples are presented, illustrating cases of graphs that enhance or suppress fixation probability for fitness r >1 as well as graphs that enhance fixation probability for only a limited range of fitness. Results are compared with recent results reported in the literature, where a positive correlation is observed between vertex degree variance and fixation probability for undirected graphs. We show a similar correlation for directed graphs, with correlation not directly to fixation probability but to the difference between fixation probability for a given graph and a complete graph.


2018 ◽  
Author(s):  
David Hathcock ◽  
Steven H. Strogatz

Evolutionary graph theory models the effects of natural selection and random drift on structured populations of mutant and non-mutant individuals. Recent studies have shown that fixation times, which determine the rate of evolution, often have right-skewed distributions. Little is known, however, about how these distributions and their skew depend on mutant fitness. Here we calculate the fitness dependence of the fixation-time distribution for the Moran Birth-death process in populations modeled by two extreme networks: the complete graph and the one-dimensional ring lattice, each of which admits an exact solution in the limit of large network size. We find that with non-neutral fitness, the Moran process on the ring has normally distributed fixation times, independent of the relative fitness of mutants and non-mutants. In contrast, on the complete graph, the fixation-time distribution is a weighted convolution of two Gumbel distributions, with a weight depending on the relative fitness. When fitness is neutral, however, the Moran process has a highly skewed fixation-time distribution on both the complete graph and the ring. In this sense, the case of neutral fitness is singular. Even on these simple network structures, the fixation-time distribution exhibits rich fitness dependence, with discontinuities and regions of universality. Applications of our methods to a multi-fitness Moran model, times to partial fixation, and evolution on random networks are discussed.


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