Interaction intensity in strategic fitness: A quantifying yardstick of selection optimization for evolutionary game.

The notion of the fitness of a strategy has been assimilated as the reproductive success in the evolutionary game. Initially, this fitness was tied to the game's pay-off and the strategy's relative frequency. However, density dependence becomes exigent in order to make ecologically reliable fitness. However, the contributions of each different type of interaction to the species's overall growth process were surprisingly under-explored. This oversight has occasionally led to either more or less prediction of strategy selection compared to the actual possibility. Moreover, density regulation of the population has always been analysed in a general way compared to strategy selection. In this context, our study introduces the concept of mean relative death payoff, which helps in assessing interaction intensity coefficients and integrates them into strategic fitness. Based on this fitness function, we develop the frequency-density replicator dynamics, which eventually provides distinguishing criteria for directional and balancing selection. Our optimized, evolutionarily stable strategy emerges as a superior alternative to the conventional trade-off between selection forces and ecological processes. More significantly, mean relative death pay-off has both conditional and quantitative roles in getting a stable population size. As a case study, we have extensively analysed the evolution of aggression using the Hawk-Dove game. We have shown that pure Dove selection is always beneficial for species growth rather than pure Hawk selection, and the condition of selection is dependent on external mortality pressure. However, the condition of coexistence is independent of external mortality pressure, representing a strong evolutionary selection that optimizes population density governed by interaction intensity.

Evolution of Cooperation in Spatio-Temporal Evolutionary Games with Public Goods Feedback.

In biology, evolutionary game-theoretical models often arise in which players' strategies impact the state of the environment, driving feedback between strategy and the surroundings. In this case, cooperative interactions can be applied to studying ecological systems, animal or microorganism populations, and cells producing or actively extracting a growth resource from their environment. We consider the framework of eco-evolutionary game theory with replicator dynamics and growth-limiting public goods extracted by population members from some external source. It is known that the two sub-populations of cooperators and defectors can develop spatio-temporal patterns that enable long-term coexistence in the shared environment. To investigate this phenomenon and unveil the mechanisms that sustain cooperation, we analyze two eco-evolutionary models: a well-mixed environment and a heterogeneous model with spatial diffusion. In the latter, we integrate spatial diffusion into replicator dynamics. Our findings reveal rich strategy dynamics, including bistability and bifurcations, in the temporal system and spatial stability, as well as Turing instability, Turing-Hopf bifurcations, and chaos in the diffusion system. The results indicate that effective mechanisms to promote cooperation include increasing the player density, decreasing the relative timescale, controlling the density of initial cooperators, improving the diffusion rate of the public goods, lowering the diffusion rate of the cooperators, and enhancing the payoffs to the cooperators. We provide the conditions for the existence, stability, and occurrence of bifurcations in both systems. Our analysis can be applied to dynamic phenomena in fields as diverse as human decision-making, microorganism growth factors secretion, and group hunting.

Revealing the hierarchical structure of microbial communities.

Measuring the dynamics of microbial communities results in high-dimensional measurements of taxa abundances over time and space, which is difficult to analyze due to complex changes in taxonomic compositions. This paper presents a new method to investigate and visualize the intrinsic hierarchical community structure implied by the measurements. The basic idea is to identify significant intersection sets, which can be seen as sub-communities making up the measured communities. Using the subset relationship, the intersection sets together with the measurements form a hierarchical structure visualized as a Hasse diagram. Chemical organization theory (COT) is used to relate the hierarchy of the sets of taxa to potential taxa interactions and to their potential dynamical persistence. The approach is demonstrated on a data set of community data obtained from bacterial 16S rRNA gene sequencing for samples collected monthly from four groundwater wells over a nearly 3-year period (n = 114) along a hillslope area. The significance of the hierarchies derived from the data is evaluated by showing that they significantly deviate from a random model. Furthermore, it is demonstrated how the hierarchy is related to temporal and spatial factors; and how the idea of a core microbiome can be extended to a set of interrelated core microbiomes. Together the results suggest that the approach can support developing models of taxa interactions in the future.

Managing ecological thresholds of a risky commons.

Common resources are often overexploited and appear subject to critical transitions from one stable state to another antagonistic state. Many times resulting in tragedy of the commons (TOC)-exploitation of shared resources for personal gain/payoffs, leading to worse outcomes or extinction. An adequate response would be strategic interaction, such as inspection and punishment by institutions to avoid TOC. This strategic interaction is often coupled with dynamically changing common resources. However, effect of strategic interaction in complex, coupled socio-ecological systems is less studied. Here, we develop replicator equations using evolving games in which strategy and common resources co-evolve. We consider the shared commons as fish dynamics governed by the intrinsic growth rate, predation and harvesting. The joint dynamics exhibit an oscillatory TOC, revealing that institutions need to pay special attention to intrinsic growth rate and nonlinear interaction. Our research shows that the co-evolving system exhibits a broader range of dynamics when predation is present compared to the disengaged fishery system. We conclude that the usefulness, chances and challenges of modelling co-evolutionary games to create sustainable systems merit further research.

Multi-objective control optimization of isolated bridge using replicator controller and NSGA-II.

Earthquakes can cause significant damage to constructed structures, leading engineers to design systems that effectively reduce damage and improve real-time vibration control. While base isolation is a commonly used passive method for seismic protection in highway structures, it has limitations such as a lack of immediate adaptation, constrained power dissipation capacity, and poor performance during earthquakes. To address the limitations of passive base isolation bearings, a hybrid control system that includes semi-active MR dampers is being introduced into isolated highway bridge structures. The aim is to enhance vibration reduction and improve overall performance. One of the major challenges in implementing this technology is developing appropriate control algorithms to handle the nonlinear behavior of semi-active devices. This paper proposes an adaptive data-driven control algorithm, informed by evolutionary game theory and a multi-objective optimization process, to optimize the distribution of voltage to semi-active MR dampers based on measurements of the damper's response to input signals. The algorithm is designed to provide optimal seismic protection. The performance of the replicator dynamics in the control system depends on three critical parameters: total population, which represents the total available resources or the sum of actuator forces; growth rate, which is the rate at which resources are distributed among control devices; and the fictitious fitness function, which regulates power consumption. Previous studies used sensitivity analysis to ascertain the best values for population size and growth rate, a time-consuming and unreliable process. This study aims to improve the performance of the system by solving a multi-objective problem. The proposed approach integrates a control algorithm with a multi-objective optimization algorithm, namely NSGA-II, to find Pareto optimal values for all parameters of the replicator dynamics. These parameters include total population, growth rate, and the fictitious function, with the aim of ensuring sustainability. By considering multiple objectives simultaneously, the proposed approach can provide a more comprehensive and effective solution for the bridge control problem. The effectiveness of this proposed approach is demonstrated through sample results Utilizing a case study centered around the Southern California Interstate 91/5 Overcrossing Highway Bridge, which is exposed to seismic activities.

Policy choices and compliance behavior in pandemic times.

In this paper, we model an evolutionary noncooperative game between politicians and citizens that, given the level of infection, describes the observed variety of mitigation policies and citizens' compliance during the COVID-19 pandemic period. Our results show that different stable equilibria exist and that different ways/paths exist to reach these equilibria may be present, depending on the choice of parameters. When the parameters are chosen opportunistically, in the short run, our model generates transitions between hard and soft policy measures to deal with the pandemic. In the long-run, convergence is achieved toward one of the possible stable steady states (obey or not obey lockdown rules) as functions of politicians' and citizens' incentives.

Weak selection and stochastic evolutionary stability in a stochastic replicator dynamics.

It is a very challenging problem whether natural selection is able to effectively resist the continuous disturbance of environmental noise such that the direction or outcome of evolution determined by the deterministic selection pressure will not be changed. By analyzing the impact of weak selection on the evolutionary stability of a stochastic replicator dynamics with n possible pure strategies, we found that the weak selection is able to enhance the evolutionary stability, that is, under weak selection, the stochastic evolutionary stability of the system is determined by the mean payoff matrix. This finding strongly implies that the weak selection should be regarded as an important mechanism to ensure evolutionary stability in stochastic environments.

Evolutionary analysis of replicator dynamics about anti-cancer combination therapy.

The emergence and growth of drug-resistant cancer cell subpopulations during anti-cancer treatment is a major challenge for cancer therapies. Combination therapies are usually applied for overcoming drug resistance. In the present paper, we explored the evolution outcome of tumor cell populations under different combination schedules of chemotherapy and p53 vaccine, by construction of replicator dynamical model for sensitive cells, chemotherapy-resistant cells and p53 vaccine-resistant cells. The local asymptotic stability analysis of the evolutionary stable points revealed that cancer population could evolve to the population with single subpopulation, or coexistence of sensitive cells and p53 vaccine-resistant cells, or coexistence of chemotherapy-resistant cells and p53 vaccine-resistant cells under different monotherapy or combination schedules. The design of adaptive therapy schedules that maintain the subpopulations under control is also demonstrated by sequential and periodic application of combination treatment strategies based on the evolutionary velocity and evolutionary absorbing regions. Applying a new replicator dynamical model, we further explored the supportive effects of sensitive cancer cells on targeted therapy-resistant cells revealed in mice experiments. It was shown that the supportive effects of sensitive cells could drive the evolution of cell population from sensitive cells to coexistence of sensitive cells and one type of targeted therapy-resistant cells.

Evolution of cooperation in multigame with environmental space and delay.

Replicator dynamics is widely used in evolutionary game theory, however, most previous studies on replicator dynamics focus on single games and ignore multiple social dilemmas encountered by individuals in a population. This paper uses replicator dynamics to construct a multigame system with environmental space and delay based on three social dilemmas. For the non-delayed and delayed multigame systems, rich dynamics for stability, bistability, transcritical bifurcation, Hopf bifurcation, and the direction, stability and periodic variation of periodic solutions are comprehensively investigated. Accordingly, we use numerical simulations to assist in exploring the effects of multigame, environmental space, and time delay on strategic dynamics. The results show that large proportions of snowdrift game and stag hunt game are conducive to the prosperity of cooperators, and defectors are easy to survive when the proportion of prisoner's dilemma is large. The cooperator gains the advantage of benefit distribution from environmental space, or the defector gets less benefit distribution as punishment, which will make pure cooperation the dominant strategy. Furthermore, environmental space can allow cooperation and defection to coexist oscillatingly. Interestingly, large delays reverse the coexistence of cooperation and defection to a situation dominated by the purely cooperative strategy.

Evolution of trustfulness in the case where resources for cooperation are sometimes absent.

It is worth investigating the existence of cooperation, which is costly for the actor but beneficial to the recipient (precisely because it is costly for the former). If players, when they approach defectors, stop their relationship with them, cooperation can pay off and favorably emerge in the course of evolutionary dynamics. The present study examines the situation in which animals, even when they want to cooperate, sometimes lack the necessary resources, and are thereby prevented from cooperating with others. In addition, it is also considered that the underlying information about the presence or absence of these resources can be conveyed to the opponent player. Here, the opponent who defects-has no resources for cooperation-may be a cooperator or a defector. Therefore, it is not clear which behavior is more likely to evolve, if it is keeping the interaction with such an opponent (i.e., being trustful) or stopping the interaction with such an opponent (i.e., being not trustful). By using evolutionary game theory, it is revealed that those who want to keep the interaction with those without the resources to cooperate are favored by natural selection. This study sheds new light on the role of keeping and stopping interaction in the evolution of cooperation under variable availability of resources.

Bayes and Darwin: How replicator populations implement Bayesian computations.

Bayesian learning theory and evolutionary theory both formalize adaptive competition dynamics in possibly high-dimensional, varying, and noisy environments. What do they have in common and how do they differ? In this paper, we discuss structural and dynamical analogies and their limits, both at a computational and an algorithmic-mechanical level. We point out mathematical equivalences between their basic dynamical equations, generalizing the isomorphism between Bayesian update and replicator dynamics. We discuss how these mechanisms provide analogous answers to the challenge of adapting to stochastically changing environments at multiple timescales. We elucidate an algorithmic equivalence between a sampling approximation, particle filters, and the Wright-Fisher model of population genetics. These equivalences suggest that the frequency distribution of types in replicator populations optimally encodes regularities of a stochastic environment to predict future environments, without invoking the known mechanisms of multilevel selection and evolvability. A unified view of the theories of learning and evolution comes in sight.

COVID-19 and Stigma: Evolution of Self-restraint Behavior.

Social stigma can effectively prevent people from going out and possibly spreading COVID-19. Using the framework of replicator dynamics, we analyze the interaction between self-restraint behavior, infection with viruses such as COVID-19, and stigma against going out. Our model is analytically solvable with respect to an interior steady state in contrast to the previous model of COVID-19 with stigma. We show that a non-legally binding policy reduces the number of people going out in a steady state.

Weak selection helps cheap but harms expensive cooperation in spatial threshold dilemmas.

Public Goods Games (PGGs) are n-person games with dependence of individual fitness benefits on the collective investment by the players. We have studied a simple PGG scenario played out by cooperating (C) and defecting (D) agents, applying the highly nonlinear threshold benefit function in an individual-based lattice model. A semi-analytical approximation of the lattice model has been developed and shown to describe the dynamics fairly well in the vicinity of the steady state. Besides the expected outcomes (i.e., the negative effect on cooperator persistence of higher cooperation costs and/or more intensive mixing of the population) we have found a surprising, counter-intuitive effect of the strength of selection on the steady state of the model. The effect is different at low and high cooperation costs, and it shows up only in the lattice model, suggesting that stochastic effects and higher order spatial correlations due to the emergent spatial clustering of cooperators (not taken into account in the semi-analytical approximation) must be responsible for the unexpected results for which we propose an intuitive explanation, present a tentative demonstration, and shortly discuss their biological relevance.

Comparison between best-response dynamics and replicator dynamics in a social-ecological model of lake eutrophication.

Social-ecological models are often used to investigate the mutual interactions between an ecological system and human behaviour at a collective level. The social system is widely represented either by the replicator dynamics or by the best-response dynamics. We investigate the consequences of choosing one or the other with the example of a social-ecological model for eutrophication in shallow lakes, where the anthropogenic discharge of pollutants into the water is determined by a behavioural model using the replicator or a best-response dynamics. We discuss a fundamental difference between the replicator dynamics and the logit formulation of the best-response dynamics. This fundamental difference results in a different number of equilibria. We show that the replicator equation is a limit case of the best-response model, when agents are assumed to behave with infinite rationality. If agents act less rationally in the model using the best-response dynamics, the correspondence with the model using the replicator dynamics decreases. Finally, we show that sustained oscillations observed in both cases may differ substantially. The replicator dynamics makes the amplitude of the limit cycle become larger and makes the system come closer to full cooperation or full defection. Thus, the dynamics along the limit cycle imply a different risk for the system to be pushed by a perturbation into a desirable or an undesirable outcome depending on the socioeconomic dynamics assumed in the model. When analyzing social-ecological models, the choice of a socioeconomic dynamics is often little justified but our results show that it may have dramatic impacts on the coupled human-environment system.

Replicator dynamics for the game theoretic selection models based on state.

The paper presents an attempt to integrate the classical evolutionary game theory based on replicator dynamics and the state-based approach of Houston and McNamara. In the new approach, individuals have different heritable strategies; however, individuals carrying the same strategy can differ in terms of state, role or the situation in which they act. Thus, the classical replicator dynamics is completed by the additional subsystem of differential equations describing the dynamics of transitions between different states. In effect, the interactions described by game structure, in addition to the demographic payoffs (constituted by births and deaths), can lead to the change in state of the competing individuals. Special cases of reversible and irreversible incremental stage-structured models, where the state changes can describeenergy accumulation, developmental steps or aging, are derived for discrete and continuous versions. The new approach is illustrated using the example of the Owner-Intruder game with explicit dynamics of the role changes. The new model presents a generalization of the demographic version of the Hawk-Dove game,with the difference being that the opponents in the game are drawn from two separate subpopulations consisting of Owners and Intruders. Here, the Intruders check random nest sites and play the Hawk-Dove game with the Owner if they are occupied. Meanwhile, the Owners produce newborns that become Intruders, since they must find a free nest site to reproduce. An interesting feedback mechanism is produced via the fluxes of individuals between the different subpopulations. In addition, the population growth suppression mechanism resulting from the fixation Bourgeois strategy is analyzed.

Cancer immunoediting: A game theoretical approach.

The role of the immune system in tumor development increasingly includes the idea of cancer immunoediting. It comprises three phases: elimination, equilibrium, and escape. In the first phase, elimination, transformed cells are recognized and destroyed by immune system. The rare tumor cells that are not destroyed in this phase may then enter the equilibrium phase, where their growth is prevented by immunity mechanisms. The escape phase represents the final phase of this process, where cancer cells begin to grow unconstrained by the immune system. In this study, we describe and analyze an evolutionary game theoretical model of proliferating, quiescent, and immune cells interactions for the first time. The proposed model is evaluated with constant and dynamic approaches. Population dynamics and interactions between the immune system and cancer cells are investigated. Stability of equilibria or critical points are analyzed by applying algebraic analysis. This model allows us to understand the process of cancer development and might help us design better treatment strategies to account for immunoediting.

The ESS for evolutionary matrix games under time constraints and its relationship with the asymptotically stable rest point of the replicator dynamics.

Recently we interpreted the notion of ESS for matrix games under time constraints and investigated the corresponding state in the polymorphic situation. Now we give two further static (monomorphic) characterizations which are the appropriate analogues of those known for classical evolutionary matrix games. Namely, it is verified that an ESS can be described as a neighbourhood invader strategy independently of the dimension of the strategy space in our non-linear situation too, that is, a strategy is an ESS if and only if it is able to invade and completely replace any monomorphic population which totally consists of individuals following a strategy close to the ESS. With the neighbourhood invader property at hand, we establish a dynamic characterization under the replicator dynamics in two dimensions which corresponds to the strong stability concept for classical evolutionary matrix games. Besides, in some special cases, we also prove the stability of the corresponding rest point in higher dimensions.

The stabilization of equilibria in evolutionary game dynamics through mutation: mutation limits in evolutionary games.

The multi-population replicator dynamics is a dynamic approach to coevolving populations and multi-player games and is related to Cross learning. In general, not every equilibrium is a Nash equilibrium of the underlying game, and the convergence is not guaranteed. In particular, no interior equilibrium can be asymptotically stable in the multi-population replicator dynamics, e.g. resulting in cyclic orbits around a single interior Nash equilibrium. We introduce a new notion of equilibria of replicator dynamics, called mutation limits, based on a naturally arising, simple form of mutation, which is invariant under the specific choice of mutation parameters. We prove the existence of mutation limits for a large class of games, and consider a particularly interesting subclass called attracting mutation limits. Attracting mutation limits are approximated in every (mutation-)perturbed replicator dynamics, hence they offer an approximate dynamic solution to the underlying game even if the original dynamic is not convergent. Thus, mutation stabilizes the system in certain cases and makes attracting mutation limits near attainable. Hence, attracting mutation limits are relevant as a dynamic solution concept of games. We observe that they have some similarity to Q-learning in multi-agent reinforcement learning. Attracting mutation limits do not exist in all games, however, raising the question of their characterization.

Phenotypic variability promotes diversity and stability in competitive communities.

Intraspecific variation is at the core of evolutionary theory, and yet, from an ecological perspective, we have few robust expectations for how this variation should affect the dynamics of large communities. Here, by adapting an approach from evolutionary game theory, we show that the incorporation of phenotypic variability into competitive networks dramatically alters the dynamics across ecological timescales, stabilising the systems and buffering the communities against demographic perturbations. The beneficial effects of phenotypic variability are strongest when there are substantial differences among phenotypes and when phenotypes are inherited with moderately high fidelity; yet even low levels of variation lead to significant increases in diversity, stability, and robustness. By identifying a simple and ubiquitous stabilising force in competitive communities, this work contributes to our core understanding of how biological diversity is maintained in natural systems.

The public goods game with shared punishment cost in well-mixed and structured populations.

Both experimental and theoretical studies have shown that punishment plays an important role in promoting cooperation. Various forms of punishment are proposed to explain why costly punishment could be maintained in the population and stabilize cooperation. Here we consider an altruistic behavior that cooperators perform cooperation and punishment simultaneously and share the punishment cost. We investigate the role of punishment cost shared among cooperators in the evolution of cooperation in public goods game. We show that the punishment can promote and stabilize cooperation when the penalty imposed on defectors is large enough compared to the punishment cost incurred by cooperators in well-mixed populations. In structured populations, cooperation could emerge under lower fine threshold and coexist with defection. However, as the penalty increases, cooperation will have a larger basin of attraction in the well-mixed population than that in the structured population. Our analytical and simulated results indicate that punishment indeed can effectively promote the evolution of cooperation. We also find that population structure can promote the coexistence of cooperation and defection but not always be beneficial to cooperation.