Consider two ancestral lineages sampled from a system of two-dimensional branching random walks with logistic regulation in the stationary regime. We study the asymptotics of their coalescence time for large initial separation and find that it agrees with well known results for a suitably scaled two-dimensional stepping stone model and also with Malécot's continuous-space approximation for the probability of identity by descent as a function of sampling distance.
This can be viewed as a justification for the replacement of locally fluctuating population sizes by fixed effective sizes. Our main tool is a joint regeneration construction for the spatial embeddings of the two ancestral lineages.[-]

We analyse patterns of genetic variability of populations in the presence of a large seed bank with the help of a new coalescent structure called seed bank coalescent. This ancestral process appears naturally as scaling limit of the genealogy of large populations that sustain seed banks, if the seed bank size and individual dormancy times are of the same order as the active population. Mutations appear as Poisson process on the active lineages, and potentially at reduced rate also on the dormant lineages. The presence of ‘dormant' lineages leads to qualitatively altered times to the most recent common ancestor and non-classical patterns of genetic diversity. To illustrate this we provide a Wright-Fisher model with seed bank component and mutation, motivated from recent models of microbial dormancy, whose genealogy can be described by the seed bank coalescent. Based on our coalescent model, we derive recursions for the expectation and variance of the time to most recent common ancestor, number of segregating sites, pairwise differences, and singletons. Commonly employed distance statistics, in the presence and absence of a seed bank, are compared. The effect of a seed bank on the expected site-frequency spectrum is also investigated. Our results indicate that the presence of a large seed bank considerably alters the distribution of some distance statistics, as well as the site-frequency spectrum. Thus, one should be able to detect the presence of a large seed bank in genetic data. Joint work with Bjarki Eldon, Adrián González Casanova, Noemi Kurt, Maite Wilke-Berenguer[-]

Pervasive natural selection can strongly influence observed patterns of genetic variation, but these effects remain poorly understood when multiple selected variants segregate in nearby regions of the genome. Classical population genetics fails to account for interference between linked mutations, which grows increasingly severe as the density of selected polymorphisms increases. I will describe a simple limit that emerges when interference is common, in which the fitness effects of individual mutations play a relatively minor role. Instead, similar to models of quantitative genetics, molecular evolution is determined by the variance in fitness within the population, defined over an effectively asexual segment of the genome (a "linkage block"). I will describe how we can exploit this insensitivity in a new "coarse-grained" coalescent framework, which approximates the effects of many weakly selected mutations with a smaller number of strongly selected mutations that create the same variance in fitness. This approximation generates accurate and efficient predictions for silent site variability when interference is common. However, these results suggest that there is reduced power to resolve individual selection pressures when interference is sufficiently widespread, since a broad range of parameters possess nearly identical patterns of silent site variability.[-]

Maladapted individuals can only colonise a new habitat if they can evolve a positive growth rate fast enough to avoid extinction - evolutionary rescue. We use the infinitesimal model to follow the evolution of the growth rate, and find that the probability that a single migrant can establish depends on just two parameters: the mean and genetic variance of fitness. With continued migration, establishment is inevitable. However, above a threshold migration rate, the population may be trapped in a sink state, in which adaptation is held back by gene flow. By assuming a constant genetic variance, we develop a diffusion approximation for the joint distribution of population size and trait mean.[-]

The infinitesimal model is based on the assumption that, conditional on the pedigree, the joint distribution of trait values is multivariate normal, then, selecting parents does not alter the variance amongst offspring. We explain how the infinitesimal model extends to include dominance as well as epistasis. Then, the evolution of a population depends on just a few quantities, which define the components of genetic variance and the inbreeding depression. In practice, the main difficulty in applying the infinitesimal model in the presence of dominance is that one must calculate the probabilities of identity by descent amongst up to four genes, which means that very many identity coefficients must be traced. We show how these coefficients can be calculated and approximated, allowing the infinitesimal model to be applied to help understand the evolutionary consequences of inbreeding depression.[-]

In the infinitesimal model, one or several quantitative traits are described as the sum of a genetic and a non-genetic component, the first being distributed within families as a normal random variable centred at the average of the parental genetic components, and with a variance independent of the parental traits. The idea behind the normal distribution of the genetic component is that the genetic part of the trait of interest is the sum of the ‘infinitesimal' contributions of the allelic states at a very large number of loci. This model has been widely used in quantitative genetics, but less so in evolutionary biology and the precise conditionsunder which it holds has remained rather vague. In this talk, we shall provide a mathematical justification of the model as the limit as the number M of loci tends to infinity of a model with Mendelian inheritance, which includes different evolutionary processes (genetic drift, recombination, selection, mutation, population structure, ...). Generalisations of the simple version of the infinitesimal model presented here, as well as some applications, will be presented in the following talks by Nick Barton and Alison Etheridge.[-]