arXiv subjects

Arne Traulsen

Publications and source records attributed to Arne Traulsen.

At least 19 recordsLinked to original sources

arXiv2026

Changes in behaviour when adherers to an intervention experience a different epidemic than non-adherers

Non-pharmaceutical interventions (NPIs), including mask-wearing, physical distancing, and hygiene measures, provide the primary means of reducing transmission in the early stages of an epidemic. Individuals adopt one of two strategies-adherence (A) or non-adherence (N) to NPIs. These strategies influence the transmission rate and thus the number of infections, but they also come with inherent costs and benefits. We propose a model coupling behavior and disease dynamics in adherers and non-adherers based on the SIR framework. This gives rise to six behavioral-epidemiological compartments. Using numerical simulations and analytical considerations, we first examine the case where strategies are fixed. Stronger NPIs and more initial adherers lead to fewer infections, and adherers consistently experience lower infection risk than non-adherers. We then introduce behavioral switching based on the benefits and costs of the two strategies. When NPIs are effective, higher transmission rates promote adherence, resulting in fewer infections. Strikingly, in high-severity outbreaks, even modestly effective NPIs can significantly reduce infections. These findings highlight the critical role of the coupling between behavior and disease dynamics, and underscore how individual choices can compromise or compensate public health interventions.

q-bio.PE↗

arXiv2026

Universal principles of cell population growth follow from local contact inhibition

Cancer cell populations often exhibit remarkably similar growth laws despite their heterogeneity. Explanations of universal cell population growth remain partly unresolved to this day. Here, we present a growth-law unification by investigating the connection between microscopic assumptions and the expected contact inhibition, which leads to five classical tumor growth laws: exponential, radial growth, fractal growth, generalized logistic, and Gompertzian growth. All five can be seen as manifestations of a single microscopic model. Agent-based simulations substantiate our theory, and we can explain differences in growth curves in experimental data from em in vitro cancer cell population growth. Thus, our framework offers a possible explanation for many mean-field laws used to empirically capture seemingly unrelated cancer or microbial growth dynamics. Our results highlight that the interplay between contact inhibition and other assumptions (e.g., well-mixed) can influence our quantitative understanding of how cancer cells grow and, in turn, how they may interact.

q-bio.PE↗

arXiv2025

Fluctuating growth rates link turnover and unevenness in species-rich communities

The maintenance of diversity, the `commonness of rarity', and compositional turnover are ubiquitous features of species-rich communities. Through a minimal model, we consider how these features reflect the interplay between environmental stochasticity, intra- and interspecific competition, and dispersal. We show that, even if species have the same time-average fitness, fluctuations tend to drive the community towards ever-growing unevenness and species extinctions, but self-limitation and/or dispersal allow species-rich states to be sustained. Species abundance--distributions vary systematically in a Buffering--Stabilization parameter plane that describes the relative strength of the underlying ecological processes, and cover different empirically relevant power-law and unimodal shapes. A model describing the effective dynamics of a focal species relates static abundance distributions with turnover dynamics, also when species have different mean fitness. The model suggests how community statistics and time series of individual species can inform on the relative importance of the ecological processes that structure diversity.

q-bio.PE↗

arXiv2025

Chaos and noise in evolutionary game dynamics

Evolutionary game theory has traditionally employed deterministic models to describe population dynamics. These models, due to their inherent nonlinearities, can exhibit deterministic chaos, where population fluctuations follow complex, aperiodic patterns. Recently, the focus has shifted towards stochastic models, quantifying fixation probabilities and analysing systems with constants of motion. Yet, the role of stochastic effects in systems with chaotic dynamics remains largely unexplored within evolutionary game theory. This study addresses how demographic noise -- arising from probabilistic birth and death events -- impacts chaotic dynamics in finite populations. We show that despite stochasticity, large populations retain a signature of chaotic dynamics, as evidenced by comparing a chaotic deterministic system with its stochastic counterpart. More concretely, the strange attractor observed in the deterministic model is qualitatively recovered in the stochastic model, where the term deterministic chaos loses its meaning. We employ tools from nonlinear dynamics to quantify how the population size influences the dynamics. We observe that for small populations, stochasticity dominates, overshadowing deterministic selection effects. However, as population size increases, the dynamics increasingly reflect the underlying chaotic structure. This resilience to demographic noise can be essential for maintaining diversity in populations, even in non-equilibrium dynamics. Overall, our results broaden our understanding of population dynamics, and revisit the boundaries between chaos and noise, showing how they maintain structure when considering finite populations in systems that are chaotic in the deterministic limit.

q-bio.PE↗

arXiv2024

Chaotic turnover of rare and abundant species in a strongly interacting model community

The composition of ecological communities varies not only between different locations but also in time. Understanding the fundamental processes that drive species towards rarity or abundance is crucial to assessing ecosystem resilience and adaptation to changing environmental conditions. In plankton communities in particular, large temporal fluctuations in species abundances have been associated with chaotic dynamics. On the other hand, microbial diversity is overwhelmingly sustained by a `rare biosphere' of species with very low abundances. We consider here the possibility that interactions within a species-rich community can relate both phenomena. We use a Lotka-Volterra model with weak immigration and strong, disordered, and mostly competitive interactions between hundreds of species to bridge single-species temporal fluctuations and abundance distribution patterns. We highlight a generic chaotic regime where a few species at a time achieve dominance, but are continuously overturned by the invasion of formerly rare species. We derive a focal-species model that captures the intermittent boom-and-bust dynamics that every species undergoes. Although species cannot be treated as effectively uncorrelated in their abundances, the community's effect on a focal species can nonetheless be described by a time-correlated noise characterized by a few effective parameters that can be estimated from time series. The model predicts a non-unitary exponent of the power-law abundance decay, which varies weakly with ecological parameters, consistent with observation in marine protist communities. The chaotic turnover regime is thus poised to capture relevant ecological features of species-rich microbial communities.

q-bio.PE↗

arXiv2024

Seasonal social dilemmas

Social dilemmas where the good of a group is at odds with individual interests are usually considered as static -- the dilemma does not change over time. In the COVID-19 pandemic, social dilemmas occurred in the mitigation of epidemic spread: Should I reduce my contacts or wear a mask to protect others? In the context of respiratory diseases, which are predominantly spreading during the winter months, some of these situations re-occur seasonally. We couple a game theoretical model, where individuals can adjust their behavior, to an epidemiological model with seasonal forcing. We find that social dilemmas can occur annually and that behavioral reactions to them can either decrease or increase the peaks of infections in a population. Our work has not only implications for seasonal infectious diseases, but also more generally for oscillatory social dilemmas: A complex interdependence between behavior and external dynamics emerges. To be effective and to exploit behavioral dynamics, intervention measures to mitigate re-occuring social dilemmas have to be timed carefully.

physics.soc-ph↗

arXiv2022

Diversity enables the jump towards cooperation for the Traveler's Dilemma

Social dilemmas are situations in which collective welfare is at odds with individual gain. One widely studied example, due to the conflict it poses between human behaviour and game theoretic reasoning, is the Traveler's Dilemma. The dilemma relies on the players' incentive to undercut their opponent at the expense of losing a collective high payoff. Such individual incentive leads players to a systematic mutual undercutting until the lowest possible payoff is reached, which is the game's unique Nash equilibrium. However, if players were satisfied with a high payoff -- that is not necessarily higher than their opponent's -- they would both be better off individually and collectively. Here, we explain how it is possible to converge to this cooperative high payoff equilibrium. Our analysis focuses on decomposing the dilemma into a local and a global game. We show that players need to escape the local maximisation and jump to the global game, in order to reach the cooperative equilibrium. Using a dynamic approach, based on evolutionary game theory and learning theory models, we find that diversity, understood as the presence of suboptimal strategies, is the general mechanism that enables the jump towards cooperation.

cs.GT↗

arXiv2022

Sharp thresholds limit the benefit of defector avoidance in cooperation on networks

Consider a cooperation game on a spatial network of habitat patches, where players can relocate between patches if they judge the local conditions to be unfavorable. In time, the relocation events may lead to a homogeneous state where all patches harbor the same relative densities of cooperators and defectors or they may lead to self-organized patterns, where some patches become safe havens that maintain an elevated cooperator density. Here we analyze the transition between these states mathematically. We show that safe havens form once a certain threshold in connectivity is crossed. This threshold can be analytically linked to the structure of the patch network and specifically to certain network motifs. Surprisingly, a forgiving defector avoidance strategy may be most favorable for cooperators. Our results demonstrate that the analysis of cooperation games in ecological metacommunity models is mathematically tractable and has the potential to link topics such as macroecological patterns, behavioral evolution, and network topology.

q-bio.PE↗

arXiv2020

Understanding evolutionary and ecological dynamics using a continuum limit

This manuscript contains nothing new, but synthesizes known results: For the theoretical population geneticist with a probabilistic background, we provide a summary of some key results on stochastic differential equations. For the evolutionary game theorist, we give a new perspective on the derivations of results obtained when using discrete birth-death processes. For the theoretical biologist familiar with deterministic modeling, we outline how to derive and work with stochastic versions of classical ecological and evolutionary processes.

q-bio.PE↗

arXiv2018

Exploring and mapping the universe of evolutionary graphs

Population structure can be modelled by evolutionary graphs, which can have a substantial, but very subtle influence on the fate of the arising mutants. Individuals are located on the nodes of these graphs, competing with each other to eventually take over the graph via the links. Many applications for this framework can be envisioned, from the ecology of river systems and cancer initiation in colonic crypts to biotechnological search for optimal mutations. In all these applications, it is not only important where and when novel variants arise and how likely it is that they ultimately take over, but also how long this process takes. More concretely, how is the probability to take over the population related to the associated time? We study this problem for all possible undirected and unweighted graphs up to a certain size. To move beyond the graph size where an exhaustive search is possible, we devise a genetic algorithm to find graphs with either high or low fixation probability and either short or long fixation time and study their structure in detail searching for common themes. Our work unravels structural properties that maximize or minimize fixation probability and time, which allows us to contribute to a first map of the universe of evolutionary graphs.

q-bio.PE↗

arXiv2017

Extinction dynamics from meta-stable coexistences in an evolutionary game

Deterministic evolutionary game dynamics can lead to stable coexistences of different types. Stochasticity, however, drives the loss of such coexistences. This extinction is usually accompanied by population size fluctuations. We investigate the most probable extinction trajectory under such fluctuations by mapping a stochastic evolutionary model to a problem of classical mechanics using the Wentzel-Kramers-Brillouin (WKB) approximation. Our results show that more abundant types in a coexistence can be more likely to go extinct first well agreed with previous results, and also the distance between the coexistence and extinction point is not a good predictor of extinction. Instead, the WKB method correctly predicts the type going extinct first.

physics.bio-ph↗

arXiv2017

Chaotic provinces in the kingdom of the Red Queen

The interplay between parasites and their hosts is found in all kinds of species and plays an important role in understanding the principles of evolution and coevolution. Usually, the different genotypes of hosts and parasites oscillate in their abundances. The well-established theory of oscillatory Red Queen dynamics proposes an ongoing change in frequencies of the different types within each species. So far, it is unclear in which way Red Queen dynamics persists with more than two types of hosts and parasites. In our analysis, an arbitrary number of types within two species are examined in a deterministic framework with constant or changing population size. This general framework allows for analytical solutions for internal fixed points and their stability. For more than two species, apparently chaotic dynamics has been reported. Here we show that even for two species, once more than two types are considered per species, irregular dynamics in their frequencies can be observed in the long run. The nature of the dynamics depends strongly on the initial configuration of the system; the usual regular Red Queen oscillations are only observed in some parts of the parameter region.

q-bio.PE↗

arXiv2017

Fixation probabilities in populations under demographic fluctuations

We study the fixation probability of a mutant type when introduced into a resident population. As opposed to the usual assumption of constant pop- ulation size, we allow for stochastically varying population sizes. This is implemented by a stochastic competitive Lotka-Volterra model. The compe- tition coefficients are interpreted in terms of inverse payoffs emerging from an evolutionary game. Since our study focuses on the impact of the competition values, we assume the same birth and death rates for both types. In this gen- eral framework, we derive an approximate formula for the fixation probability ϕ of the mutant type under weak selection. The qualitative behavior of ϕ when compared to the neutral scenario is governed by the invasion dynamics of an initially rare type. Higher payoffs when competing with the resident type yield higher values of ϕ. Additionally, we investigate the influence of the remaining parameters and find an explicit dependence of ϕ on the mixed equilibrium value of the corresponding deterministic system (given that the parameter values allow for its existence).

q-bio.PE↗

arXiv2017

Evolutionary games on cycles with strong selection

Evolutionary games on graphs describe how strategic interactions and population structure determine evolutionary success, quantified by the probability that a single mutant takes over a population. Graph structures, compared to the well-mixed case, can act as amplifiers or suppressors of selection by increasing or decreasing the fixation probability of a beneficial mutant. Properties of the associated mean fixation times can be more intricate, especially when selection is strong. The intuition is that fixation of a beneficial mutant happens fast (in a dominance game), that fixation takes very long (in a coexistence game), and that strong selection eliminates demographic noise. Here we show that these intuitions can be misleading in structured populations. We analyze mean fixation times on the cycle graph under strong frequency-dependent selection for two different microscopic evolutionary update rules (death-birth and birth-death). We establish exact analytical results for fixation times under strong selection, and show that there are coexistence games in which fixation occurs in time polynomial in population size. Depending on the underlying game, we observe inherence of demographic noise even under strong selection, if the process is driven by random death before selection for birth of an offspring (death-birth update). In contrast, if selection for an offspring occurs before random removal (birth-death update), strong selection can remove demographic noise almost entirely.

q-bio.PE↗

arXiv2016

Exact numerical calculation of fixation probability and time on graphs

The Moran process on graphs is a popular model to study the dynamics of evolution in a spatially structured population. Exact analytical solutions for the fixation probability and time of a new mutant have been found for only a few classes of graphs so far. Simulations are time-expensive and many realizations are necessary, as the variance of the fixation times is high. We present an algorithm that numerically computes these quantities for arbitrary small graphs by an approach based on the transition matrix. The advantage over simulations is that the calculation has to be executed only once. Building the transition matrix is automated by our algorithm. This enables a fast and interactive study of different graph structures and their effect on fixation probability and time. We provide a fast implementation in C with this note https://github.com/hindersin/efficientFixation. Our code is very flexible, as it can handle two different update mechanisms (Birth-death or death-Birth), as well as arbitrary directed or undirected graphs.

q-bio.PE↗

arXiv2016

Most undirected random graphs are amplifiers of selection for Birth-death dynamics, but suppressors of selection for death-Birth dynamics

We analyze evolutionary dynamics on graphs, where the nodes represent individuals of a population. The links of a node describe which other individuals can be displaced by the offspring of the individual on that node. Amplifiers of selection are graphs for which the fixation probability is increased for advantageous mutants and decreased for disadvantageous mutants. A few examples of such amplifiers have been developed, but so far it is unclear how many such structures exist and how to construct them. Here, we show that almost any undirected random graph is an amplifier of selection for Birth-death updating, where an individual is selected to reproduce with probability proportional to its fitness and one of its neighbors is replaced by that offspring at random. If we instead focus on death-Birth updating, in which a random individual is removed and its neighbors compete for the empty spot, then the same ensemble of graphs consists of almost only suppressors of selection for which the fixation probability is decreased for advantageous mutants and increased for disadvantageous mutants. Thus, the impact of population structure on evolutionary dynamics is a subtle issue that will depend on seemingly minor details of the underlying evolutionary process.

q-bio.PE↗

arXiv2015

When the mean is not enough: Calculating fixation time distributions in birth-death processes

Studies of fixation dynamics in Markov processes predominantly focus on the mean time to absorption. This may be inadequate if the distribution is broad and skewed. We compute the distribution of fixation times in one-step birth-death processes with two absorbing states. These are expressed in terms of the spectrum of the process, and we provide different representations as forward-only processes in eigenspace. These allow efficient sampling of fixation time distributions. As an application we study evolutionary game dynamics, where invading mutants can reach fixation or go extinct. We also highlight the median fixation time as a possible analog of mixing times in systems with small mutation rates and no absorbing states, whereas the mean fixation time has no such interpretation.

cond-mat.stat-mech↗

arXiv2015

Stochastic evolutionary games in dynamic populations

Frequency dependent selection and demographic fluctuations play important roles in evolutionary and ecological processes. Under frequency dependent selection, the average fitness of the population may increase or decrease based on interactions between individuals within the population. This should be reflected in fluctuations of the population size even in constant environments. Here, we propose a stochastic model, which naturally combines these two evolutionary ingredients by assuming frequency dependent competition between different types in an individual-based model. In contrast to previous game theoretic models, the carrying capacity of the population and thus the population size is determined by pairwise competition of individuals mediated by evolutionary games and demographic stochasticity. In the limit of infinite population size, the averaged stochastic dynamics is captured by the deterministic competitive Lotka-Volterra equations. In small populations, demographic stochasticity may instead lead to the extinction of the entire population. As the population size is driven by the fitness in evolutionary games, a population of cooperators is less prone to go extinct than a population of defectors, whereas in the usual systems of fixed size, the population would thrive regardless of its average payoff.

q-bio.PE↗