Open research questions in Evolutionary Game Theory and Cooperation

What the literature leaves open

• The challenge of testing a gene-culture coevolutionary model of Christian denominational differentiation. The difficulty of incorporating phylogenetically controlled regression models into the analysis. The need to address the complexity of the relationships between gene-culture coevolution, parasite stress, and Christian denominational differentiation.

  The Role of Gene–Culture Coevolution and Parasite Stress in Diversifying Christian Denominations · 2026 · DOI

• The lack of understanding of the role of gene-culture coevolution in shaping religious diversity. The need for a model that explains the diversification of Christian denominations. The limited use of phylogenetic methods in the study of religious diversity.

  The Role of Gene–Culture Coevolution and Parasite Stress in Diversifying Christian Denominations · 2026 · DOI

• The paper identifies a gap in the literature regarding the effects of altruism on socio-economic outcomes. It argues that the conventional narrative that altruism leads to beneficial outcomes is incomplete, and that a public choice perspective is needed to analyze the effects of altruism. The paper also highlights the importance of considering the incentives and rules that govern behavior, rather than relying solely on benevolence.

  Dysfunctional effects of altruism: an introduction to the symposium · 2026 · DOI

• However, there often will be ecological reasons not to intervene, insofar as the effects of predation enable the flourishing of other animals (including but not limited to the predator).

  Equality, Ecology, and the Problem of Predation · 2026 · DOI

• The nature of intelligence remains scientifically poorly understood and the subject of many disagreements both biologically and psychologically.

  Intelligence, from Origins to Human Culture: An Eco-Evolutionary Perspective · 2025 · DOI

• Its true form, the nature of its variation and its evolutionary origins remain unclear.

  Intelligence, from Origins to Human Culture: An Eco-Evolutionary Perspective · 2025 · DOI

• In reality, human activities often involve temporal interactions, where links are impermanent, and understanding the evolution of cooperation on such temporal networks is an open problem.

  Promoting collective cooperation through temporal interactions · 2025 · DOI

• However, achieving collective, cooperative behavior-in which intelligent actors in complex environments jointly improve their well-being-remains poorly understood.

  Collective cooperative intelligence · 2025 · DOI

• In hunter-gatherer societies, women's subsistence activities are crucial for food provisioning and children's social learning but are understudied relative to men's activities.

  Women’s subsistence networks scaffold cultural transmission among BaYaka foragers in the Congo Basin · 2024 · DOI

• While this switching behavior has been mainly attributed to random environmental factors, it remains unclear the extent to which internal community dynamics affect the switching capacity of microbial systems.

  Structured community transitions explain the switching capacity of microbial systems · 2024 · DOI

• PRACTICAL IMPLICATIONS: Originality/value While research on negative gossip exists, this study specifically examines its impact on frontline service employees, a crucial but under-studied group in service organizations.

  Workplace gossip erodes proactive work behavior: anxiety and neuroticism as underlying mechanisms · 2024 · DOI

• The study identifies a gap in our understanding of the impact of slow spatial migration on cooperative antimicrobial resistance in time-varying environments. Prior work has shown that drug resistance is enhanced by migration in static conditions, but this changes in time-fluctuating spatially structured environments.

  Slow spatial migration can help eradicate cooperative antimicrobial resistance in time-varying environments · 2026 · DOI

• Translation to the laboratory: chemostats, microfluidic setups, and batch cultures. Our modelling approach is mainly inspired by chemostat setups, which are commonly used in laboratory-controlled experiments to modulate the influx of nutrients and drugs in microbial communities. In such systems, the concentrations of resources and toxins can be adjusted to impose harsh conditions that generate population bottlenecks, whose eco-evolutionary impacts are the subject of intense study [25,88,93]. Here, we focus on the biologically relevant regime of intermediate environmental time variation [25,94,119], characterised by ν ≲ 1 and 0 carrying capacity, whereas the local composition (number of S and R cells in a deme) relaxes more slowly, on a timescale δ ≲ 1, in which the population size within each deme rapidly tracks the ≤ 1/s with typically s ≲ 10–1 (s = 0.1 in all figures here; see “Background” in Model & Methods and S1 Appendix Sec. ∼ 1.2.1). This regime corresponds to conditions fluctuating between mild (K = K+) and harsh (K = K–) environmental states with a frequency between once per hour and once per day, that is, approximately every 1 – 100 microbial generations (ν = 0.01 – 1; see Model & Methods). The drug influx is kept constant, and each environmental switch, theoretically treated as instantaneous, occurs rapidly in practice. While we have conveniently represented the switching of the carrying capacity as a random process at rates ν 1/ν+ + 1/ν–, would not change the qualitative results of our study (“Background” in Model & Methods, S1 Appendix Sec. 1.2.3), and could be seen as a potential laboratory implementation of this model. (Fig 1A), the case where K varies periodically between K+ and K–, with period ± Since all the above conditions can be practically implemented [25,94], we believe that our theoretical predictions can, in principle, be probed in prospective laboratory-controlled experiments. The environmental switching of PLOS Computational Biology | https://doi.org/10.1371/journal.pcbi.1013997 March 16, 2026 22 / 31 K(t) would be realised using a sequence of spatially connected, fixed-volume chemostats, each acting as a deme, see, e.g., Ref.. The rate of cell migration would be set by the rate of volume exchange between neighbouring demes-chemostats (0.001% – 10% of the volume every hour). Moreover, with microfluidic devices and single-cell techniques, it is possible to perform spatially structured experiments involving as few as 10 – 100 cells per microhabitat patch [9,107,149]. These conditions are consistent with our modelling parameters, notably those corresponding to demes of relatively small size under harsh conditions (e.g., K = K– = 80). The migration rate between microhabitats in such setups largely depends on the experimental design (e.g., number of patch-to-patch channels, channel cross-section).

  Slow spatial migration can help eradicate cooperative antimicrobial resistance in time-varying environments · 2026 · DOI

• Further application of reinforcement learning to ecological systems. Investigation of the role of reinforcement learning in the emergence of other traits, such as trust and fairness.

  A brief review of evolutionary game dynamics in the reinforcement learning paradigm · 2026 · DOI

• The gap between theoretical predictions and behavioral experiments in evolutionary game theory. The lack of a unified framework for understanding social and ecological phenomena.

  A brief review of evolutionary game dynamics in the reinforcement learning paradigm · 2026 · DOI

• The experiment was conducted with a limited number of participants. The experiment was conducted in a laboratory setting, which may not reflect real-world scenarios.

  Cooperative protection against stochastic losses: Experimental evidence on behavioral dynamics · 2026 · DOI

• The paper predicts that cooperative equilibria with exactly two cooperating players (Q=2) may only exist under DamRed and only for sufficiently risk-averse subjects, yet no analysis of actual observed equilibrium coordination on specific Q values is provided, nor experimental conditions designed to investigate which Qmin thresholds subjects actually implement.

  Cooperative protection against stochastic losses: Experimental evidence on behavioral dynamics · 2026 · DOI

• Further research could explore the implications of the Leviathan Paradox for political and social systems. The authors' alternative architectural model could be tested and refined through further research.

  Cooperation as Attractor: A Multi-Level Formal Refutation of Hobbes's Bellum Omnium Contra Omnes · 2026 · DOI

• The paper identifies a gap in the literature regarding the refutation of Hobbes's thesis. The authors argue that prior work has not provided a systematic, multi-level refutation of Hobbes's claim.

  Cooperation as Attractor: A Multi-Level Formal Refutation of Hobbes's Bellum Omnium Contra Omnes · 2026 · DOI

• Further studies can investigate the effects of other factors on cooperation in the three-strategy snowdrift game. The paper's results can be used to develop new strategies to promote cooperation in social and biological systems.

  Strategy Dynamics of Three-Strategy Snowdrift Game Induced by Reward Strategy and Payoff Delay · 2026 · DOI

• The evolutionary game dynamics of a three-strategy snowdrift game with special reward strategy and payoff delay has not been fully investigated. The effects of reward and payoff delay on cooperation are not well understood.

  Strategy Dynamics of Three-Strategy Snowdrift Game Induced by Reward Strategy and Payoff Delay · 2026 · DOI

• the approach requires numerical integration of the governing equations, - the numerical experiments presented in this work focus on small and intermediate-size systems, - extending the framework to much larger networks remains an important direction for future research

  Dissimilarity measures for generalized Lotka-Volterra systems on networks · 2026 · DOI

• applying the method to empirically derived interaction networks, - developing approximate or reduced descriptions of the dynamics capable of preserving the relevant dynamical information while remaining computationally tractable

  Dissimilarity measures for generalized Lotka-Volterra systems on networks · 2026 · DOI

• Further study is needed to understand the impact of environment heterogeneity on mutant fixation in different types of graphs. The study's findings could be extended to other types of populations, such as animal or plant populations. The development of new experimental systems to test the study's predictions is needed.

  Environment heterogeneity creates fast amplifiers of natural selection in graph-structured populations · 2026 · DOI

• Although RPS systems have provided a classic framework for understanding cyclic competition and have been extended to include HOIs, the specific effects of these interactions on biodiversity in increasingly complex systems remain uncertain.

  Nonlinear effects of higher-order interactions on pattern formation and species persistence in cyclic ecosystems · 2026 · DOI

Most-cited papers in Evolutionary Game Theory and Cooperation

• Five Rules for the Evolution of Cooperation · Science · 2006 · 4,913 citations
• Evolutionary games and spatial chaos · Nature · 1992 · 3,717 citations
• Collective Action and the Evolution of Social Norms · The Journal of Economic Perspectives · 2000 · 2,636 citations
• The ecology of the microbiome: Networks, competition, and stability · Science · 2015 · 2,541 citations
• The diversity–stability debate · Nature · 2000 · 2,513 citations
• Patterns of Aggressive Behavior in Experimentally Created “Social Climates” · The Journal of Social Psychology · 1939 · 1,779 citations
• An Evolutionary Approach to Norms · American Political Science Review · 1986 · 1,562 citations
• The evolution of cooperation in strategic alliances: Initial conditions or learning processes? · Strategic Management Journal · 2007 · 1,179 citations
• Cultural group selection, coevolutionary processes and large-scale cooperation · Journal of Economic Behavior & Organization · 2003 · 722 citations
• Ants, Rationality, and Recruitment · The Quarterly Journal of Economics · 1993 · 689 citations

Most recent work

• Public goods games on any population structure · Science Advances · 2026
• Predicting competition and substrate preferences for targeted microbiome alteration · Cell · 2026
• Pattern formation in a predator–prey model on network and non-network environments · Chaos, Solitons & Fractals · 2026
• A higher-order equivalence of Lotka-Volterra and replicator dynamics reveals tight connections between ecology and evolution · bioRxiv · 2026
• Social structure as a form of collective intelligence: a new framework · Philosophical Transactions B · 2026
• Cooperation in collective dilemmas under opinion-based risk perceptions · Applied Mathematical Modelling · 2026
• Repeated evolution of supergenes on an ancient social chromosome · Current Biology · 2026
• Community coalescence reveals strong selection and coexistence within species in complex microbial communities · bioRxiv · 2026
• Promotion of cooperation in deme-structured populations with growth-merging dynamics · bioRxiv · 2026
• Nonlinear public goods game in dynamical environments · Applied Mathematical Modelling · 2026

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