In this paper, we investigate the consequences of dormancy in the `rare mutation' and `large population' regime of stochastic adaptive dynamics. Starting from an individual-based micro-model, we first derive the polymorphic evolution sequence of the population, based on previous work by Baar and Bovier (2018). After passing to a second `small mutations' limit, we arrive at the canonical equation of adaptive dynamics, and state a corresponding criterion for evolutionary branching, extending a previous result of Champagnat and Méléard (2011).
The criterion allows a quantitative and qualitative analysis of the effects of dormancy in the well-known model of Dieckmann and Doebeli (1999) for sympatric speciation. In fact, a quite intuitive picture merges: Dormancy enlarges the parameter range for evolutionary branching, increases the carrying capacity and niche width of the post-branching sub-populations, and, depending on the model parameters, can either increase or decrease the `speed of adaptation' of populations. Finally, dormancy increases diversity by increasing the genetic distance between subpopulations.
In the present article, we investigate the effects of dormancy on an abstract population genetic level. We first provide a short review of seed bank models in population genetics, and the role of dormancy for the interplay of evolutionary forces in general, before we discuss two recent paradigmatic models, referring to spontaneous resp. simultaneous switching of individuals between the active and the dormant state. We show that both mechanisms give rise to non-trivial mathematical objects, namely the (continuous) seed bank diffusion and the seed bank diffusion with jumps, as well as their dual…
The interplay of selection and dormancy in a Moran model can lead to coexistence of types
Goethe-Universität Frankfurt（法兰克福歌德大学）；Budapest University of Technology and Economics（布达佩斯技术与经济大学）；HUN-REN Alfréd Rényi Institute of Mathematics（匈牙利研究与教育网络阿尔弗雷德·雷尼数学研究所）
We study the interplay of population growth and evolutionary dynamics using a stochastic model based on birth and death events. In contrast to the common assumption of an independent population size, evolution can be strongly affected by population dynamics in general. Especially for fast reproducing microbes which are subject to selection, both types of dynamics are often closely intertwined. We illustrate this by considering different growth scenarios. Depending on whether microbes die or stop to reproduce (dormancy), qualitatively different behaviors emerge. For cooperating bacteria, a perm…
Evolutionary branching points are a paradigmatic feature of adaptive dynamics, because they are potential starting points for adaptive diversification. The antithesis to evolutionary branching points are Continuously stable strategies (CSS's), which are convergent stable and evolutionarily stable equilibrium points of the adaptive dynamics and hence are thought to represent endpoints of adaptive processes. However, this assessment is based on situations in which the invasion fitness function determining the adaptive dynamics have non-zero second derivatives at a CSS. Here we show that the scop…
Game theory research on the snowdrift game has showed that gradual evolution of the continuously varying level of cooperation in joint enterprises can demonstrate evolutionary merging as well as evolutionary branching. However, little is known about the consequences of changes in diversity at the cooperation level. In the present study I consider effects of costly rewards on the continuous snowdrift game. I show that not evolutionary merging but evolutionary branching can promote the emergence of pool reward, which can then enhance social welfare.
We analyze evolutionary dynamics in a confluent, branching cellular population, such as in a growing duct, vasculature, or in a branching microbial colony. We focus on the coarse-grained features of the evolution and build a statistical model that captures the essential features of the dynamics. Using simulations and analytic approaches, we show that the survival probability of strains within the growing population is sensitive to the branching geometry: Branch bifurcations enhance survival probability due to an overall population growth (i.e., "inflation"), while branch termination and the sm…
Studies of cooperation have traditionally focused on discrete games such as the well-known prisoner's dilemma, in which players choose between two pure strategies: cooperation and defection. Increasingly, however, cooperation is being studied in continuous games that feature a continuum of strategies determining the level of cooperative investment. For the continuous snowdrift game, it has been shown that a gradually evolving monomorphic population may undergo evolutionary branching, resulting in the emergence of a defector strategy that coexists with a cooperator strategy. This phenomenon has…
Investigating the emergence of a particular cell type is a recurring theme in models of growing cellular populations. The evolution of resistance to therapy is a classic example. Common questions are: when does the cell type first occur, and via which sequence of steps is it most likely to emerge? For growing populations, these questions can be formulated in a general framework of branching processes spreading through a graph from a root to a target vertex. Cells have a particular fitness value on each vertex and can transition along edges at specific rates. Vertices represents cell states, sa…
The central goal of a dynamical theory of evolution is to abstract the mean evolutionary trajectory in the trait space by considering ecological processes at the level of the individual. In this work, we develop such a theory for a new class of deterministic individual based models describing individual births and deaths, which captures the essential features of standard stochastic individual-based models and become identical with the latter under maximal competition. The key motivation is to derive the canonical equation of adaptive dynamics from this microscopic ecological model, which can b…
We propose a faster algorithm for individual based simulations for adaptive dynamics based on a simple modification to the standard Gillespie Algorithm for simulating stochastic birth-death processes. We provide an analytical explanation that shows that simulations based on the modified algorithm, in the deterministic limit, lead to the same equations of adaptive dynamics as well as same conditions for evolutionary branching as those obtained from the standard Gillespie algorithm. Based on this algorithm, we provide an intuitive and simple interpretation of the canonical equation of adaptive d…
The interplay between mutation and selection plays a fundamental role in the behaviour of evolutionary algorithms (EAs). However, this interplay is still not completely understood. This paper presents a rigorous runtime analysis of a non-elitist population-based EA that uses the linear ranking selection mechanism. The analysis focuses on how the balance between parameter $η$, controlling the selection pressure in linear ranking, and parameter $χ$ controlling the bit-wise mutation rate, impacts the runtime of the algorithm. The results point out situations where a correct balance between select…
Understanding if and how mutants reach fixation in populations is an important question in evolutionary biology. We study the impact of population growth has on the success of mutants. To systematically understand the effects of growth we decouple competition from reproduction; competition follows a birth--death process and is governed by an evolutionary game, while growth is determined by an externally controlled branching rate. In stochastic simulations we find non-monotonic behaviour of the fixation probability of mutants as the speed of growth is varied; the right amount of growth can lead…
Although mutations drive the evolutionary process, the rates at which the mutations occur are themselves subject to evolutionary forces. Our purpose here is to understand the role of selection and random genetic drift in the evolution of mutation rates, and we address this question in asexual populations at mutation-selection equilibrium neglecting selective sweeps. Using a multitype branching process, we calculate the fixation probability of a rare nonmutator in a large asexual population of mutators, and find that a nonmutator is more likely to fix when the deleterious mutation rate of the m…
In this work we present a flexible tool for tumor progression, which simulates the evolutionary dynamics of cancer. Tumor progression implements a multi-type branching process where the key parameters are the fitness landscape, the mutation rate, and the average time of cell division. The fitness of a cancer cell depends on the mutations it has accumulated. The input to our tool could be any fitness landscape, mutation rate, and cell division time, and the tool produces the growth dynamics and all relevant statistics.
We derive an expression for the variation between parallel trajectories in phenotypic evolution, extending the well known result that predicts the mean evolutionary path in adaptive dynamics or quantitative genetics. We show how this expression gives rise to the notion of fluctuation domains - parts of the fitness landscape where the rate of evolution is very predictable (due to fluctuation dissipation) and parts where it is highly variable (due to fluctuation enhancement). These fluctuation domains are determined by the curvature of the fitness landscape. Regions of the fitness landscape with…
Most theories of evolutionary diversification are based on equilibrium assumptions: they are either based on optimality arguments involving static fitness landscapes, or they assume that populations first evolve to an equilibrium state before diversification occurs, as exemplified by the concept of evolutionary branching points in adaptive dynamics theory. Recent results indicate that adaptive dynamics may often not converge to equilibrium points and instead generate complicated trajectories if evolution takes place in high-dimensional phenotype spaces. Even though some analytical results on d…
University of Toronto（多伦多大学）；Ontario Institute for Cancer Research（安大略癌症研究所）
High-throughput sequencing allows the detection and quantification of frequencies of somatic single nucleotide variants (SNV) in heterogeneous tumor cell populations. In some cases, the evolutionary history and population frequency of the subclonal lineages of tumor cells present in the sample can be reconstructed from these SNV frequency measurements. However, automated methods to do this reconstruction are not available and the conditions under which reconstruction is possible have not been described. We describe the conditions under which the evolutionary history can be uniquely reconstruct…
We are interested in modeling the Darwinian evolution resulting from the interplay of phenotypic variation and natural selection through ecological interactions, in the specific scales of the biological framework of adaptive dynamics. Adaptive dynamics so far has been put on a rigorous footing only for direct competition models (Lotka-Volterra models) involving a competition kernel which describes the competition pressure from one individual to another one. We extend this to a multi-resources chemostat model, where the competition between individuals results from the sharing of several resourc…
The aim of this paper is to discuss the role of robustness and diversity in population dynamics in particular to some properties of the multi-step from healthy tissue to fully malignant tumours. Recent evidence shows that diversity within the cell population of a neoplasm, a pre-tumoural lession that can develop into a fully malignant tumour, is the best predictor for its evolving into a tumour. By studying the dynamics of a population described by a multi-type, population-size limited branching process in terms of the evolutionary formalism, we show some general principles regarding the proba…
The evolution of leadership in migratory populations depends not only on costs and benefits of leadership investments but also on the opportunities for individuals to rely on cues from others through social interactions. We derive an analytically tractable adaptive dynamic network model of collective migration with fast timescale migration dynamics and slow timescale adaptive dynamics of individual leadership investment and social interaction. For large populations, our analysis of bifurcations with respect to investment cost explains the observed hysteretic effect associated with recovery of …
Branching processes are widely used to model evolutionary and population dynamics as well as the spread of infectious diseases. To characterize the dynamics of their growth or spread, the basic reproduction number $R_0$ has received considerable attention. In the context of infectious diseases, it is usually defined as the expected number of secondary cases produced by an infectious case in a completely susceptible population. Typically $R_0>1$ indicates that an outbreak is expected to continue and to grow exponentially, while $R_0<1$ usually indicates that an outbreak is expected to terminate…
