The embedding problem of Markov transition matrices into continuous-time Markov semigroups is a classic problem that regained a lot of impetus and activities in recent years. We consider it here for the following generalisation of the well-known coupon collection process: from a finite set of distinct objects, a subset is drawn repeatedly according to some probability distribution, independently and with replacement, and each time united with the set of objects sampled so far. We derive and interpret properties of and explicit conditions for the resulting discrete-time Markov chain to be representable within a semigroup or a flow of a continuous-time process of the same type.
We establish a genealogical framework for an existing analytical moment duality between a Wright–Fisher type SDE and a counting process with interaction. To achieve this, we construct a finite-population Moran model featuring interactive neutral reproduction as a novel mechanism. In the corresponding events, an individual, regardless of its own type, can only reproduce if a randomly encountered partner is of the “fit” type. This Moran model has a relatively simple counting process as its factorial moment dual, whose genealogical meaning appears to be cryptic: after all, the line-counting process of the natural genealogical process of the model, namely the ancestral influence graph (AIG), exhibits a complex hierarchical structure not reflected in the factorial moment dual. Since moment duality is a property in expectation, we are allowed to systematically remove information from the AIG and merge different realizations of the ancestry. We call the result the Frankenstein process. Based on this, we establish the factorial moment duality from a genealogical perspective. The moment duality in the diffusion limit follows in a natural way.
The classic recombination equation, both in discrete and in continuous time, can be solved in a way that derives from the Markov chain of a partitioning process. Here, we revisit this structure from the point of view of the Markov embedding problem. In particular, we analyse when a discrete-time Markov matrix of recombination type can occur in a time-homogeneous Markov semigroup that is generated by a recombination rate matrix. En route, we also show that such rate matrices (or Markov generators) generally do not form a matrix algebra, but span a real Lie algebra.
We consider a Moran-type model of cultural evolution, which describes how traits emerge, are transmitted, and get lost in populations. Our analysis focuses on the underlying cultural genealogies; they were first described by Aguilar and Ghirlanda (2015) and are closely related to the ancestral selection graph of population genetics, wherefore we call them ancestral learning graphs. We investigate their dynamical behaviour, that is, we are concerned with evolving genealogies. In particular, we consider the total length of the genealogy of the entire population as a function of the (forward) time where we start looking back. This quantity shows a sawtooth-like dynamics with linear increase interrupted by collapses to near-zero at random times. We relate this to the metastable behaviour of the stochastic logistic model, which describes the evolution of the number of ancestors as well as the number of descendants of a given sample. We superpose types to the model by assuming that new inventions appear independently in every individual, and all traits of the cultural parent are transmitted to the learner in any given learning event. The set of traits of an individual then agrees with the set of innovations along its genealogy. The properties of the genealogy thus translate into the properties of the trait set of a sample. In particular, the moments of the number of traits are obtained from the moments of the total length of the genealogy.
The embeddability of reversible Markov matrices into time-homogeneous Markov semigroups is revisited, with some focus on simplifications and extensions. In particular, we do not demand irreducibility and consider weakly reversible matrices as well as reversible matrices with negative eigenvalues.
We consider the long-term behaviour of critical multitype branching processes conditioned on non-extinction, both with respect to the forward and the ancestral processes. Forward in time, we prove a functional limit theorem in the space of trajectories of the linearly-scaled h-transformed process; the change of measure allows us to work on the same probability space for all times. Backward in time, we trace the lines of descent of individuals sampled from a (stationary) population and analyse the ancestral type distribution, that is, the type distribution of the ancestors in the distant past, as well as the type process along the ancestral line.
We consider the Moran model of population genetics with two types, mutation, and selection, and investigate the line of descent of a randomly-sampled individual from a contemporary population. We trace this ancestral line back into the distant past, far beyond the most recent common ancestor of the population (thus connecting population genetics to phylogeny), and analyse the mutation process along this line.To this end, we use the pruned lookdown ancestral selection graph (Lenz et al., 2015), which consists of a set of potential ancestors of the sampled individual at any given time. Relative to the neutral case (that is, without selection), we obtain a general bias towards the beneficial type, an increase in the beneficial mutation rate, and a decrease in the deleterious mutation rate. This sheds new light on previous analytical results. We discuss our findings in the light of a well-known observation at the interface of phylogeny and population genetics, namely, the difference in the mutation rates (or, more precisely, mutation fluxes) estimated via phylogenetic methods relative to those observed in pedigree studies.
We establish connections between the absorption probabilities of a class of birth-death processes with killing, and the stationary tail of a related class of birth-death processes with catastrophes. The major ingredients of the proofs are a decomposition of the dynamics of these processes, a Feynman–Kac type relationship for Markov chains with reset and rebirth, and the concept of Siegmund duality, which allows us to invert the relationship between the processes. We apply our results to a pair of ancestral processes in population genetics, namely the killed ancestral selection graph and the pruned lookdown ancestral selection graph, in a finite population setting and its diffusion limit.
The computation of matrix functions is a well-studied problem. Of special importance are the exponential and the logarithm of a matrix, where the latter also raises existence and uniqueness questions. This is particularly relevant in the context of matrix semigroups and their generators. Here, we look at matrix functions of triangular matrices, where a recursive approach is possible when the matrix has simple spectrum. The special feature is that no knowledge of eigenvectors is required, and that the same recursion applies to the computation of multiple functions or semigroups simultaneously.
We study ancestral structures for the two-type Moran model with mutation and frequency-dependent selection under the nonlinear dominance or fittest-type-wins scheme. Under appropriate conditions, both lead, in distribution, to the same type-frequency process. Reasoning through the mutations on the ancestral selection graph (ASG), we develop the corresponding killed and pruned lookdown ASGs and use them to determine the present and ancestral type distributions. To this end, we establish factorial moment dualities to the Moran model and a relative. We extend the results to the diffusion limit and present applications for finite populations, as well as for moderate and weak selection limits.
We consider the mutation-selection differential equation with pairwise interaction (or, equivalently, the diploid mutation-selection equation) and establish the corresponding ancestral process, which is a random tree and a variant of the ancestral selection graph. The formal relation to the forward model is given via duality. To make the tree tractable, we prune branches upon mutations, thus reducing it to its informative parts. The hierarchies inherent in the tree are encoded systematically via tripod trees with weighted leaves; this leads to the stratified ancestral selection graph. The latter also satisfies a duality relation with the mutation-selection equation. Each of the dualities provides a stochastic representation of the solution of the differential equation. This allows us to connect the equilibria and their bifurcations to the long-term behaviour of the ancestral process. Furthermore, with the help of the stratified ancestral selection graph, we obtain explicit results about the ancestral type distribution in the case of unidirectional mutation.
The deterministic selection-recombination equation describes the evolution of the genetic type composition of a population under selection and recombination in a law of large numbers regime. So far, an explicit solution has seemed out of reach; only in the special case of three sites with selection acting on one of them has an approximate solution been found, but without an obvious path to generalisation. We use both an analytical and a probabilistic, genealogical approach for the case of an arbitrary number of neutral sites linked to one selected site. This leads to a recursive integral representation of the solution. Starting from a variant of the ancestral selection-recombination graph, we develop an efficient genealogical structure, which may, equivalently, be represented as a weighted partitioning process, a family of Yule processes with initiation and resetting, and a family of initiation processes. We prove them to be dual to the solution of the differential equation forward in time and thus obtain a stochastic representation of the deterministic solution, along with the Markov semigroup in closed form.
We consider the discrete-time migration–recombination equation, a deterministic, nonlinear dynamical system that describes the evolution of the genetic type distribution of a population evolving under migration and recombination in a law of large numbers setting. We relate this dynamics (forward in time) to a Markov chain, namely a labelled partitioning process, backward in time. This way, we obtain a stochastic representation of the solution of the migration–recombination equation. As a consequence, one obtains an explicit solution of the nonlinear dynamics, simply in terms of powers of the transition matrix of the Markov chain. The limiting and quasi-limiting behaviour of the Markov chain are investigated, which gives immediate access to the asymptotic behaviour of the dynamical system. We finally sketch the analogous situation in continuous time.
Recently, the selection–recombination equation with a single selected site and an arbitrary number of neutral sites was solved by Alberti and Baake (2021) 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.
This chapter gives a synopsis of recent approaches to model and analyse the evolution of microbial populations under selection. The first part reviews two population genetic models of Lenski's long-term evolution experiment with Escherichia coli, where models aim at explaining the observed curve of the evolution of the mean fitness. The second part describes a model of a host-pathogen system where the population of pathogenes experiences balancing selection, migration, and mutation, as motivated by observations of the genetic diversity of HCMV (the human cytomegalovirus) across hosts.
Solving the recombination equation has been a long-standing challenge of \emph{deterministic} population genetics. We review recent progress obtained by introducing ancestral processes, as traditionally used in the context of \emph{stochastic} models of population genetics, into the deterministic setting. With the help of an ancestral partitioning process, which is obtained by letting population size tend to infinity (without rescaling parameters or time) in an ancestral recombination graph, we obtain the solution to the recombination equation in a transparent form.
We revisit the model by Wiser et al. (2013), which describes how the mean fitness increases over time due to beneficial mutations in Lenski’s long-term evolution experiment. We develop the model further both conceptually and mathematically. Conceptually, we describe the experiment with the help of a Cannings model with mutation and selection, where the latter includes diminishing returns epistasis. The analysis sheds light on the growth dynamics within every single day and reveals a runtime effect, that is, the shortening of the daily growth period with increasing fitness; and it allows to clarify the contribution of epistasis to the mean fitness curve. Mathematically, we explain rigorous results in terms of a law of large numbers (in the limit of infinite population size and for a certain asymptotic parameter regime), and present approximations based on heuristics and supported by simulations for finite populations.
Reaction systems and population dynamics constitute two highly developed areas of research that build on well-defined model classes, both in terms of dynamical systems and stochastic processes. Despite a significant core of common structures, the two fields have largely led separate lives. The workshop brought the communities together and emphasised concepts, methods and results that have, so far, appeared in one area but are potentially useful in the other as well.
We consider a discrete-time Markov chain, called fragmentation process, that describes a specific way of successively removing objects from a linear arrangement. The process arises in population genetics and describes the ancestry of the genetic material of individuals in a population experiencing recombination. We aim at the law of the process over time. To this end, we investigate sets of realisations of this process that agree with respect to a specific order of events and represent each such set by a rooted (binary) tree. The probability of each tree is, in turn, obtained by M\obius inversion on a suitable poset of all rooted forests that can be obtained from the tree by edge deletion; we call this poset the \textit{pruning poset}. Dependencies within the fragments make it difficult to obtain explicit expressions for the probabilities of the trees. We therefore construct an auxiliary process for every given tree, which is i.i.d. over time, and which allows to give a pathwise construction of realisations that match the tree.
Thomas Wiehe合作论文数University of Cologne
Institute for Genetics2