arXiv subjects

Arne Traulsen

Publications and source records attributed to Arne Traulsen.

At least 37 records · Page 2Linked to original sources

arXiv2015

Counterintuitive properties of the fixation time in network-structured populations

Evolutionary dynamics on graphs can lead to many interesting and counterintuitive findings. We study the Moran process, a discrete time birth-death process, that describes the invasion of a mutant type into a population of wild-type individuals. Remarkably, the fixation probability of a single mutant is the same on all regular networks. But non-regular networks can increase or decrease the fixation probability. While the time until fixation formally depends on the same transition probabilities as the fixation probabilities, there is no obvious relation between them. For example, an amplifier of selection, which increases the fixation probability and thus decreases the number of mutations needed until one of them is successful, can at the same time slow down the process of fixation. Based on small networks, we show analytically that (i) the time to fixation can decrease when links are removed from the network and (ii) the node providing the best starting conditions in terms of the shortest fixation time depends on the fitness of the mutant. Our results are obtained analytically on small networks, but numerical simulations show that they are qualitatively valid even in much larger populations.

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arXiv2015

Cancer initiation with epistatic interactions between driver and passenger mutations

We investigate the dynamics of cancer initiation in a mathematical model with one driver mutation and several passenger mutations. Our analysis is based on a multi type branching process: We model individual cells which can either divide or undergo apoptosis. In case of a cell division, the two daughter cells can mutate, which potentially confers a change in fitness to the cell. In contrast to previous models, the change in fitness induced by the driver mutation depends on the genetic context of the cell, in our case on the number of passenger mutations. The passenger mutations themselves have no or only a very small impact on the cell's fitness. While our model is not designed as a specific model for a particular cancer, the underlying idea is motivated by clinical and experimental observations in Burkitt Lymphoma. In this tumor, the hallmark mutation leads to deregulation of the MYC oncogene which increases the rate of apoptosis, but also the proliferation rate of cells. This increase in the rate of apoptosis hence needs to be overcome by mutations affecting apoptotic pathways, naturally leading to an epistatic fitness landscape. This model shows a very interesting dynamical behavior which is distinct from the dynamics of cancer initiation in the absence of epistasis. Since the driver mutation is deleterious to a cell with only a few passenger mutations, there is a period of stasis in the number of cells until a clone of cells with enough passenger mutations emerges. Only when the driver mutation occurs in one of those cells, the cell population starts to grow rapidly.

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arXiv2014

When do microscopic assumptions determine the outcome in evolutionary game dynamics?

The modelling of evolutionary game dynamics in finite populations requires microscopic processes that determine how strategies spread. The exact details of these processes are often chosen without much further consideration. Different types of microscopic models, including in particular fitness-based selection rules and imitation-based dynamics, are often used as if they were interchangeable. We challenge this view and investigate how robust these choices on the micro-level really are. Focusing on a key macroscopic observable, the probability for a single mutant to take over a population of wild-type individuals, we show that there is a unique pair of a fitness-based process and an imitation process leading to identical outcomes for arbitrary games and for all intensities of selection. This highlights the perils of making arbitrary choices at the micro-level without regard of the consequences at the macro-level.

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arXiv2014

Zero-determinant alliances in multiplayer social dilemmas

Direct reciprocity and conditional cooperation are important mechanisms to prevent free riding in social dilemmas. But in large groups these mechanisms may become ineffective, because they require single individuals to have a substantial influence on their peers. However, the recent discovery of the powerful class of zero-determinant strategies in the iterated prisoner's dilemma suggests that we may have underestimated the degree of control that a single player can exert. Here, we develop a theory for zero-determinant strategies for multiplayer social dilemmas, with any number of involved players. We distinguish several particularly interesting subclasses of strategies: fair strategies ensure that the own payoff matches the average payoff of the group; extortionate strategies allow a player to perform above average; and generous strategies let a player perform below average. We use this theory to explore how individuals can enhance their strategic options by forming alliances. The effects of an alliance depend on the size of the alliance, the type of the social dilemma, and on the strategy of the allies: fair alliances reduce the inequality within their group; extortionate alliances outperform the remaining group members; but generous alliances increase welfare. Our results highlight the critical interplay of individual control and alliance formation to succeed in large groups.

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arXiv2014

Evolutionary Multiplayer Games

Evolutionary game theory has become one of the most diverse and far reaching theories in biology. Applications of this theory range from cell dynamics to social evolution. However, many applications make it clear that inherent non-linearities of natural systems need to be taken into account. One way of introducing such non-linearities into evolutionary games is by the inclusion of multiple players. An example is of social dilemmas, where group benefits could e.g.\ increase less than linear with the number of cooperators. Such multiplayer games can be introduced in all the fields where evolutionary game theory is already well established. However, the inclusion of non-linearities can help to advance the analysis of systems which are known to be complex, e.g. in the case of non-Mendelian inheritance. We review the diachronic theory and applications of multiplayer evolutionary games and present the current state of the field. Our aim is a summary of the theoretical results from well-mixed populations in infinite as well as finite populations. We also discuss examples from three fields where the theory has been successfully applied, ecology, social sciences and population genetics. In closing, we probe certain future directions which can be explored using the complexity of multiplayer games while preserving the promise of simplicity of evolutionary games.

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arXiv2013

Mutualism and evolutionary multiplayer games: revisiting the Red King

Coevolution of two species is typically thought to favour the evolution of faster evolutionary rates helping a species keep ahead in the Red Queen race, where `it takes all the running you can do to stay where you are'. In contrast, if species are in a mutualistic relationship, it was proposed that the Red King effect may act, where it can be beneficial to evolve slower than the mutualistic species. The Red King hypothesis proposes that the species which evolves slower can gain a larger share of the benefits. However, the interactions between the two species may involve multiple individuals. To analyse such a situation, we resort to evolutionary multiplayer games. Even in situations where evolving slower is beneficial in a two-player setting, faster evolution may be favoured in a multiplayer setting. The underlying features of multiplayer games can be crucial for the distribution of benefits. They also suggest a link between the evolution of the rate of evolution and group size.

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arXiv2012

The mechanics of stochastic slowdown in evolutionary games

We study the stochastic dynamics of evolutionary games, and focus on the so-called `stochastic slowdown' effect, previously observed in (Altrock et. al, 2010) for simple evolutionary dynamics. Slowdown here refers to the fact that a beneficial mutation may take longer to fixate than a neutral one. More precisely, the fixation time conditioned on the mutant taking over can show a maximum at intermediate selection strength. We show that this phenomenon is present in the prisoner's dilemma, and also discuss counterintuitive slowdown and speedup in coexistence games. In order to establish the microscopic origins of these phenomena, we calculate the average sojourn times. This allows us to identify the transient states which contribute most to the slowdown effect, and enables us to provide an understanding of slowdown in the takeover of a small group of cooperators by defectors: Defection spreads quickly initially, but the final steps to takeover can be delayed substantially. The analysis of coexistence games reveals even more intricate behavior. In small populations, the conditional average fixation time can show multiple extrema as a function of the selection strength, e.g., slowdown, speedup, and slowdown again. We classify two-player games with respect to the possibility to observe non-monotonic behavior of the conditional average fixation time as a function of selection strength.

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arXiv2012

Mixing times in evolutionary game dynamics

Without mutation and migration, evolutionary dynamics ultimately leads to the extinction of all but one species. Such fixation processes are well understood and can be characterized analytically with methods from statistical physics. However, many biological arguments focus on stationary distributions in a mutation-selection equilibrium. Here, we address the equilibration time required to reach stationarity in the presence of mutation, this is known as the mixing time in the theory of Markov processes. We show that mixing times in evolutionary games have the opposite behaviour from fixation times when the intensity of selection increases: In coordination games with bistabilities, the fixation time decreases, but the mixing time increases. In coexistence games with metastable states, the fixation time increases, but the mixing time decreases. Our results are based on simulations and the WKB approximation of the master equation.

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arXiv2012

Stochastic differential equations for evolutionary dynamics with demographic noise and mutations

We present a general framework to describe the evolutionary dynamics of an arbitrary number of types in finite populations based on stochastic differential equations (SDE). For large, but finite populations this allows to include demographic noise without requiring explicit simulations. Instead, the population size only rescales the amplitude of the noise. Moreover, this framework admits the inclusion of mutations between different types, provided that mutation rates, $μ$, are not too small compared to the inverse population size 1/N. This ensures that all types are almost always represented in the population and that the occasional extinction of one type does not result in an extended absence of that type. For $μN\ll1$ this limits the use of SDE's, but in this case there are well established alternative approximations based on time scale separation. We illustrate our approach by a Rock-Scissors-Paper game with mutations, where we demonstrate excellent agreement with simulation based results for sufficiently large populations. In the absence of mutations the excellent agreement extends to small population sizes.

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arXiv2011

Strategy abundance in evolutionary many-player games with multiple strategies

Evolutionary game theory is an abstract and simple, but very powerful way to model evolutionary dynamics. Even complex biological phenomena can sometimes be abstracted to simple two-player games. But often, the interaction between several parties determines evolutionary success. Rather than pair-wise interactions, in this case we must take into account the interactions between many players, which are inherently more complicated than the usual two-player games, but can still yield simple results. In this manuscript we derive the composition of a many-player multiple strategy system in the mutation-selection equilibrium. This results in a simple expression which can be obtained by recursions using coalescence theory. This approach can be modified to suit a variety of contexts, e.g. to find the equilibrium frequencies of a finite number of alleles in a polymorphism or that of different strategies in a social dilemma in a cultural context.

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arXiv2010

Universality of weak selection

Weak selection, which means a phenotype is slightly advantageous over another, is an important limiting case in evolutionary biology. Recently it has been introduced into evolutionary game theory. In evolutionary game dynamics, the probability to be imitated or to reproduce depends on the performance in a game. The influence of the game on the stochastic dynamics in finite populations is governed by the intensity of selection. In many models of both unstructured and structured populations, a key assumption allowing analytical calculations is weak selection, which means that all individuals perform approximately equally well. In the weak selection limit many different microscopic evolutionary models have the same or similar properties. How universal is weak selection for those microscopic evolutionary processes? We answer this question by investigating the fixation probability and the average fixation time not only up to linear, but also up to higher orders in selection intensity. We find universal higher order expansions, which allow a rescaling of the selection intensity. With this, we can identify specific models which violate (linear) weak selection results, such as the one--third rule of coordination games in finite but large populations.

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arXiv2010

A homoclinic route to asymptotic full cooperation in adaptive networks and its failure

We consider the evolutionary dynamics of a cooperative game on an adaptive network, where the strategies of agents (cooperation or defection) feed back on their local interaction topology. While mutual cooperation is the social optimum, unilateral defection yields a higher payoff and undermines the evolution of cooperation. Although no a priori advantage is given to cooperators, an intrinsic dynamical mechanism can lead asymptotically to a state of full cooperation. In finite systems, this state is characterized by long periods of strong cooperation interrupted by sudden episodes of predominant defection, suggesting a possible mechanism for the systemic failure of cooperation in real-world systems.

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arXiv2010

Stochastic slowdown in evolutionary processes

We examine birth--death processes with state dependent transition probabilities and at least one absorbing boundary. In evolution, this describes selection acting on two different types in a finite population where reproductive events occur successively. If the two types have equal fitness the system performs a random walk. If one type has a fitness advantage it is favored by selection, which introduces a bias (asymmetry) in the transition probabilities. How long does it take until advantageous mutants have invaded and taken over? Surprisingly, we find that the average time of such a process can increase, even if the mutant type always has a fitness advantage. We discuss this finding for the Moran process and develop a simplified model which allows a more intuitive understanding. We show that this effect can occur for weak but non--vanishing bias (selection) in the state dependent transition rates and infer the scaling with system size. We also address the Wright-Fisher model commonly used in population genetics, which shows that this stochastic slowdown is not restricted to birth-death processes.

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arXiv2010

Deterministic evolutionary game dynamics in finite populations

Evolutionary game dynamics describes the spreading of successful strategies in a population of reproducing individuals. Typically, the microscopic definition of strategy spreading is stochastic, such that the dynamics becomes deterministic only in infinitely large populations. Here, we introduce a new microscopic birth--death process that has a fully deterministic strong selection limit in well--mixed populations of any size. Additionally, under weak selection, from this new process the frequency dependent Moran process is recovered. This makes it a natural extension of the usual evolutionary dynamics under weak selection. We find simple expressions for the fixation probabilities and average fixation times of the new process in evolutionary games with two players and two strategies. For cyclic games with two players and three strategies, we show that the resulting deterministic dynamics crucially depends on the initial condition in a non--trivial way.

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arXiv2010

The pace of evolution across fitness valleys

How fast does a population evolve from one fitness peak to another? We study the dynamics of evolving, asexually reproducing populations in which a certain number of mutations jointly confer a fitness advantage. We consider the time until a population has evolved from one fitness peak to another one with a higher fitness. The order of mutations can either be fixed or random. If the order of mutations is fixed, then the population follows a metaphorical ridge, a single path. If the order of mutations is arbitrary, then there are many ways to evolve to the higher fitness state. We address the time required for fixation in such scenarios and study how it is affected by the order of mutations, the population size, the fitness values and the mutation rate.

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arXiv2010

Evolutionary games in the multiverse

Evolutionary game dynamics of two players with two strategies has been studied in great detail. These games have been used to model many biologically relevant scenarios, ranging from social dilemmas in mammals to microbial diversity. Some of these games may in fact take place between a number of individuals and not just between two. Here, we address one-shot games with multiple players. As long as we have only two strategies, many results from two player games can be generalized to multiple players. For games with multiple players and more than two strategies, we show that statements derived for pairwise interactions do no longer hold. For two player games with any number of strategies there can be at most one isolated internal equilibrium. For any number of players $\boldsymbol{d}$ with any number of strategies n, there can be at most (d-1)^(n-1) isolated internal equilibria. Multiplayer games show a great dynamical complexity that cannot be captured based on pairwise interactions. Our results hold for any game and can easily be applied for specific cases, e.g. public goods games or multiplayer stag hunts.

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arXiv2010

Human strategy updating in evolutionary games

Evolutionary game dynamics describes not only frequency dependent genetical evolution, but also cultural evolution in humans. In this context, successful strategies spread by imitation. It has been shown that the details of strategy update rules can have a crucial impact on evolutionary dynamics in theoretical models and e.g. significantly alter the level of cooperation in social dilemmas. But what kind of strategy update rules can describe imitation dynamics in humans? Here, we present a way to measure such strategy update rules in a behavioral experiment. We use a setting in which individuals are virtually arranged on a spatial lattice. This produces a large number of different strategic situations from which we can assess strategy updating. Most importantly, spontaneous strategy changes corresponding to mutations or exploration behavior are more frequent than assumed in many models. Our experimental approach to measure properties of the update mechanisms used in theoretical models will be useful for mathematical models of cultural evolution.

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arXiv2009

Mutation-selection equilibrium in games with multiple strategies

In evolutionary games the fitness of individuals is not constant but depends on the relative abundance of the various strategies in the population. Here we study general games among n strategies in populations of large but finite size. We explore stochastic evolutionary dynamics under weak selection, but for any mutation rate. We analyze the frequency dependent Moran process in well-mixed populations, but almost identical results are found for the Wright-Fisher and Pairwise Comparison processes. Surprisingly simple conditions specify whether a strategy is more abundant on average than 1/n, or than another strategy, in the mutation-selection equilibrium. We find one condition that holds for low mutation rate and another condition that holds for high mutation rate. A linear combination of these two conditions holds for any mutation rate. Our results allow a complete characterization of n*n games in the limit of weak selection.

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