{"title":"Evolutionarily stable strategies in population games: An invader’s perspective","authors":"Felipe A. Murgel , Max O. Souza","doi":"10.1016/j.jtbi.2025.112108","DOIUrl":"10.1016/j.jtbi.2025.112108","url":null,"abstract":"<div><div>We revisit the various definitions of an Evolutionarily Stable Strategy (<span><math><mi>ESS</mi></math></span>) in nonlinear population games from the standpoint of barrier functions. We show the equivalence between an <span><math><mi>ESS</mi></math></span> being uniformly uninvadable and the corresponding barrier function being lower semi-continuous (LSC). Moreover, it is sufficient to check this for strategies that are near an alternative best reply lying on an opposite face. We also provide some counterexamples that show that uniform stability cannot be taken for granted in nonlinear population games; we denote such <span><math><mi>ESS</mi></math></span>s as singular <span><math><mi>ESS</mi></math></span>s. Furthermore, we obtain conditions that are equivalent to the barrier function being LSC and are typically easier to verify. As a by-product, we identify a number of instances where being an <span><math><mi>ESS</mi></math></span> is equivalent to being uniformly uninvadable: 3-player games, payoffs inducing convex incentives, or differentiable payoffs with negative definite first derivative, when considered on alternative best replies.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"607 ","pages":"Article 112108"},"PeriodicalIF":1.9,"publicationDate":"2025-04-16","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143839761","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Ancestral process for infectious disease outbreaks with superspreading","authors":"Xavier Didelot , David Helekal , Ian Roberts","doi":"10.1016/j.jtbi.2025.112109","DOIUrl":"10.1016/j.jtbi.2025.112109","url":null,"abstract":"<div><div>When an infectious disease outbreak is of a relatively small size, describing the ancestry of a sample of infected individuals is difficult because most ancestral models assume large population sizes. Given a set of infected individuals, we show that it is possible to express exactly the probability that they have the same infector, either inclusively (so that other individuals may have the same infector too) or exclusively (so that they may not). To compute these probabilities requires knowledge of the offspring distribution, which determines how many infections each infected individual causes. We consider transmission both without and with superspreading, in the form of a Poisson and a Negative-Binomial offspring distribution, respectively. We show how our results can be incorporated into a new Lambda-coalescent model which allows multiple lineages to coalesce together. We call this new model the Omega-coalescent, we compare it with previously proposed alternatives, and advocate its use in future studies of infectious disease outbreaks.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"607 ","pages":"Article 112109"},"PeriodicalIF":1.9,"publicationDate":"2025-04-14","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143830184","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"What does the tree of life look like as it grows? Evolution and the multifractality of time","authors":"Kevin Hudnall , Raissa M. D’Souza","doi":"10.1016/j.jtbi.2025.112121","DOIUrl":"10.1016/j.jtbi.2025.112121","url":null,"abstract":"<div><div>By unifying three foundational principles of modern biology, we develop a mathematical framework to analyze the growing tree of life. Contrary to the static case, where the analogy between phylogenetic trees and the tree that grows in soil is drawn, our framework shows that the living tree of life is analogous to a Cantor dust where each branch is a distinct fractal curve. The system as a whole is therefore multifractal in the sense that it consists of many unique fractals. The three foundational principles for the mathematical framework are that phylogeny is nested, phylogeny is dualistic (i.e., transitive between singularities and populations), and phylogeny is stochastic. Integrating these three principles, we model the dynamic (i.e., <em>living</em>) tree of life as a random iterated function system that generates unique convexly related sequences of branching random variables (visualized in <span><span><strong>Animation 1</strong></span></span>). The multifractal nature of this dynamic tree of life implies that, for any two living entities, the time interval from their last common ancestor to the present moment is a distinct fractal curve for each. Thus, the length of a time interval along each distinct branch is unique, so that time is also multifractal and not an ultrametric on the tree of life.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"607 ","pages":"Article 112121"},"PeriodicalIF":1.9,"publicationDate":"2025-04-11","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143870413","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Vivi Rottschäfer , Willem G.N. Kuppers , Jiao Chen , Ed van Bavel
{"title":"Arterial arcades and collaterals regress under hemodynamics-based diameter adaptation: A computational and mathematical analysis","authors":"Vivi Rottschäfer , Willem G.N. Kuppers , Jiao Chen , Ed van Bavel","doi":"10.1016/j.jtbi.2025.112111","DOIUrl":"10.1016/j.jtbi.2025.112111","url":null,"abstract":"<div><div>Arterial networks exhibit a wide range of segment radii, largely thought to result from adaptation to wall shear stress (WSS). Segments remodel outward or inward if WSS is higher or lower than a reference value. While this mechanism seems straightforward for arterial trees, real networks contain arcades, collaterals, and loops. We investigated the stability of these looping structures under WSS control using simulation models of small networks and published coronary and cerebral artery data.</div><div>Adaptation was modeled as changes in segment radius proportional to deviations from reference WSS. A generalized model included other hemodynamic stimuli like flow and velocity. Simulations consistently predicted loop regression due to the loss of one or more segments, both for the WSS model and the generalized model, regardless of initial conditions or model parameters. This loop loss was also observed in networks with heterogeneous adaptation rates or under dynamic conditions.</div><div>A mathematical analysis confirmed that loop instability is a direct consequence of Kirchhoff’s circuit law, leading to unstable equilibria. Thus, loss of loops is an inherent outcome of arterial networks adapting to local hemodynamics. Additional mechanisms, such as communication between connected segments, may be needed to explain the presence of loops in real networks.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"607 ","pages":"Article 112111"},"PeriodicalIF":1.9,"publicationDate":"2025-04-06","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143812910","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Marco Colnaghi , Fernando P. Santos , Paul A.M. Van Lange , Daniel Balliet
{"title":"Power asymmetry destabilizes reciprocal cooperation in social dilemmas","authors":"Marco Colnaghi , Fernando P. Santos , Paul A.M. Van Lange , Daniel Balliet","doi":"10.1016/j.jtbi.2025.112106","DOIUrl":"10.1016/j.jtbi.2025.112106","url":null,"abstract":"<div><div>Direct reciprocity has been long identified as a mechanism to support the evolution of cooperation in social dilemmas. While most research on reciprocal cooperation has focused on symmetrical interactions, real world interactions often involve differences in power. Verbal theories have either claimed that power differences enhance or destabilize cooperation, indicating the need for a comprehensive theoretical model of how power asymmetries affect direct reciprocity. Here, we investigate the relationship between power and cooperation in two frequently studied social dilemmas, the prisoner’s dilemma (PD) and the snowdrift game (SD). Combining evolutionary game theory and agent-based models, we demonstrate that power asymmetries are detrimental to the evolution of cooperation. Strategies that are contingent on power within an interaction provide a selective advantage in the iterated SD, but not in the iterated PD. In both games, the rate of cooperation declines as power asymmetry increases, indicating that a more egalitarian distribution of the benefits of cooperation is the prerequisite for direct reciprocity to evolve and be maintained.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"606 ","pages":"Article 112106"},"PeriodicalIF":1.9,"publicationDate":"2025-04-06","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143812913","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"A single-locus quantitative genetic model incorporating DNA methylation","authors":"L. Ayres, H. Bovenhuis, M.P.L. Calus","doi":"10.1016/j.jtbi.2025.112110","DOIUrl":"10.1016/j.jtbi.2025.112110","url":null,"abstract":"<div><div>We describe a single-locus quantitative genetic model that incorporates effects due to DNA methylation. Extending Fisher’s decomposition of the genotypic value, we distinguish two quantities to predict an individual’s phenotypic or genetic values: the “basic genetic value” and the “expressed genetic value”. We show how these quantities relate to the concept of breeding value and derive their corresponding formulas, along with those for phenotypic variance and covariance between relatives. The resulting parameters are influenced by several factors, including the population distribution of DNA methylation levels, the functional relationship between methylation and phenotype, the magnitudes of genetic and methylation effects, and allele frequencies. We show that under the conditions modeled, the presence of DNA methylation does not bias estimated breeding values.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"607 ","pages":"Article 112110"},"PeriodicalIF":1.9,"publicationDate":"2025-04-04","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143796982","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Kathryn H. Bowers , Daniela De Angelis , Paul J. Birrell
{"title":"Modelling with SPEED: a Stochastic Predictor of Early Epidemic Detection","authors":"Kathryn H. Bowers , Daniela De Angelis , Paul J. Birrell","doi":"10.1016/j.jtbi.2025.112120","DOIUrl":"10.1016/j.jtbi.2025.112120","url":null,"abstract":"<div><div>The frequency of emerging infectious disease outbreaks continues to rise, necessitating predictive frameworks for public health decision-making. This study introduces the Stochastic Predictor of Early Epidemic Detection (SPEED) model, an adaptation of the classic Susceptible-Infected-Recovered model, employing a Gillespie-like algorithm to simulate early-stage stochastic disease transmission. SPEED incorporates individual-level detection probabilities based on the infection time and the lag from GP consultation to lab confirmation. The model dynamically adjusts to public health responses by enhancing testing and reducing detection times once a single case has been identified. SPEED serves two key functionalities. First, as a statistical inference tool refining reproduction number estimates following the detection of a small number of cases. SPEED inference uses specified prior distributions for the reproduction number to provide reliable posterior estimates. Second, to simulate epidemic scenarios under specified values of the reproduction number in order to construct a distribution of the time to subsequent detections. The model is used to evaluate how second case timings can rule out higher values of the reproduction number. Comparisons with simulations under heightened surveillance scenarios demonstrate the model’s utility in assessing response efficacy on the initial outbreak spread. Our results demonstrate SPEED applied to a single case of influenza A(H1N2)v, detected through routine flu surveillance on the 23rd November 2023.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"607 ","pages":"Article 112120"},"PeriodicalIF":1.9,"publicationDate":"2025-04-04","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143797051","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Nadiah P. Kristensen , Ryan A. Chisholm , Hisashi Ohtsuki
{"title":"Many-strategy games in groups with relatives and the evolution of coordinated cooperation","authors":"Nadiah P. Kristensen , Ryan A. Chisholm , Hisashi Ohtsuki","doi":"10.1016/j.jtbi.2025.112089","DOIUrl":"10.1016/j.jtbi.2025.112089","url":null,"abstract":"<div><div>Humans often cooperate in groups with friends and family members with varying degrees of genetic relatedness. Past kin selection can also be relevant to interactions between strangers, explaining how the cooperation first arose in the ancestral population. However, modelling the effects of relatedness is difficult when the benefits of cooperation scale nonlinearly with the number of cooperators (e.g., economies of scale). Here, we present a direct fitness method for rigorously accounting for kin selection in <span><math><mi>n</mi></math></span>-player interactions with <span><math><mi>m</mi></math></span> discrete strategies, where a genetically homophilic group-formation model is used to calculate the necessary higher-order relatedness coefficients. Our approach allows us to properly account for non-additive fitness effects between relatives (synergy). Analytical expressions for dynamics are obtained, and they can be solved numerically for modestly sized groups and numbers of strategies. We illustrate with an example where group members can verbally agree (cheap talk) to contribute to a public good with a sigmoidal benefit function, and we find that such coordinated cooperation is favoured by kin selection. As interactions switched from family to strangers, in order for coordinated cooperation to persist and for the population to resist invasion by liars, either some level of homophily must be maintained or following through on the agreement must be in the self-interests of contributors. Our approach is useful for scenarios where fitness effects are non-additive and the strategies are best modelled in a discrete way, such as behaviours that require a cognitive ‘leap’ of insight into the situation (e.g., shared intentionality, punishment).</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"605 ","pages":"Article 112089"},"PeriodicalIF":1.9,"publicationDate":"2025-03-28","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143725288","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Alistair D. Falconer , Zhiyong Li , Dietmar B. Oelz
{"title":"Two-lane track geometry facilitates coordination of collective cell migration","authors":"Alistair D. Falconer , Zhiyong Li , Dietmar B. Oelz","doi":"10.1016/j.jtbi.2025.112105","DOIUrl":"10.1016/j.jtbi.2025.112105","url":null,"abstract":"<div><div>Collective cell migration is an essential biological process in the formation and maintenance of tissues, yet its regulation is still not well understood. We formulate a 2D particle model for the collective cell migration in a confluent monolayer of epithelial cells within constrained annular geometries. Key elements of the model are directional persistence modelled by a mutual reinforcement between cell polarity and velocity, as well as a process to determine neighbourhoods of cells through adjacency in the associated Voronoi tessellation. We test this model using recently published experimental results concerning the onset of large-scale global migratory behaviour in annular geometries. Our simulations predict that tracks which are narrow enough to prevent the presence of neighbouring cells lateral to the direction of motion lead to less efficient organisation of globally coordinated migration.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"606 ","pages":"Article 112105"},"PeriodicalIF":1.9,"publicationDate":"2025-03-25","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143733230","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Plant–pollinator interaction model with separate pollen and nectar dynamics","authors":"Tomás A. Revilla","doi":"10.1016/j.jtbi.2025.112096","DOIUrl":"10.1016/j.jtbi.2025.112096","url":null,"abstract":"<div><div>The mutualism between plants and pollinators involves the exchange between plant resources and pollen dispersal services among con-specific plants. Since many pollinators are generalist foragers, the quality of pollination is compromised by inter-specific pollen transfer (IPT). This article proposes a mechanistic approach to model plant–pollinator interactions that considers the dynamics of pollen pick-up and its delivery to con-specific and hetero-specific targets, in parallel but separated from the consumption dynamics of plant rewards by the pollinators. This mechanism can model the interference effect caused by IPT on plant fitness, and predicts saturating effects on the quantity and efficiency of pollination. By coupling the mechanism with population dynamics, the resulting model demonstrates how plant–pollinator associations can shift between net mutualism and parasitism, depending on the ecological context and species traits.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"606 ","pages":"Article 112096"},"PeriodicalIF":1.9,"publicationDate":"2025-03-24","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143733229","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}