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Ancestral processes with selection: Branching and Moran models

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 Added by Ellen Baake
 Publication date 2007
  fields Biology
and research's language is English




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115 - Sandra Kluth , Ellen Baake 2013
We reconsider the Moran model in continuous time with population size $N$, two allelic types, and selection. We introduce a new particle representation, which we call the labelled Moran model, and which has the same distribution of type frequencies as the original Moran model, provided the initial values are chosen appropriately. In the new model, individuals are labelled $1,2, dots, N$; neutral resampling events may take place between arbitrary labels, whereas selective events only occur in the direction of increasing labels. With the help of elementary methods only, we not only recover fixation probabilities, but also obtain detailed insight into the number and nature of the selective events that play a role in the fixation process forward in time.
Recently, the selection-recombination equation with a single selected site and an arbitrary number of neutral sites was solved by means of the ancestral selection-recombination graph. Here, we introduce a more accessible approach, namely the ancestral initiation graph. The construction is based on a discretisation of the selection-recombination equation. We apply our method to systematically explain a long-standing observation concerning the dynamics of linkage disequilibrium between two neutral loci hitchhiking along with a selected one. In particular, this clarifies the nontrivial dependence on the position of the selected site.
We study the evolution of the population genealogy in the classic neutral Moran Model of finite size and in discrete time. The stochastic transformations that shape a Moran population can be realized directly on its genealogy and give rise to a process with a state space consisting of the finite set of Yule trees of a certain size. We derive a number of properties of this process, and show that they are in agreement with existing results on the infinite-population limit of the Moran Model. Most importantly, this process admits time reversal, which gives rise to another tree-valued Markov Chain and allows for a thorough investigation of the Most Recent Common Ancestor process.
In a (two-type) Wright-Fisher diffusion with directional selection and two-way mutation, let $x$ denote todays frequency of the beneficial type, and given $x$, let $h(x)$ be the probability that, among all individuals of todays population, the individual whose progeny will eventually take over in the population is of the beneficial type. Fearnhead [Fearnhead, P., 2002. The common ancestor at a nonneutral locus. J. Appl. Probab. 39, 38-54] and Taylor [Taylor, J. E., 2007. The common ancestor process for a Wright-Fisher diffusion. Electron. J. Probab. 12, 808-847] obtained a series representation for $h(x)$. We develop a construction that contains elements of both the ancestral selection graph and the lookdown construction and includes pruning of certain lines upon mutation. Besides being interesting in its own right, this construction allows a transparent derivation of the series coefficients of $h(x)$ and gives them a probabilistic meaning.
We define the Sampled Moran Genealogy Process, a continuous-time Markov process on the space of genealogies with the demography of the classical Moran process, sampled through time. To do so, we begin by defining the Moran Genealogy Process using a novel representation. We then extend this process to include sampling through time. We derive exact conditional and marginal probability distributions for the sampled process under a stationarity assumption, and an exact expression for the likelihood of any sequence of genealogies it generates. This leads to some interesting observations pertinent to existing phylodynamic methods in the literature. Throughout, our proofs are original and make use of strictly forward-in-time calculations and are exact for all population sizes and sampling processes.
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