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Daniel B. Cooney

Publications and source records attributed to Daniel B. Cooney.

14 recordsLinked to original sources

Kernel-Dependent Pattern Formation in a Population Model with Nonlocal Facilitation and Competition

Spatial patterns, such as those in dryland vegetation models, have historically been studied in systems of reaction-diffusion systems with pattern onset via a Turing bifurcation from a spatially uniform state. More recently, spatial patterns have been considered in models that incorporate spatially extended interactions via nonlocal interaction kernels. It remains largely underexplored if and how the choice of nonlocal interaction kernel contributes to differences in pattern formation and persistence, particularly in models that contain competition and facilitation. Here, we investigate spatial patterns in a reaction-diffusion model for a single species that includes nonlocal competition and facilitation processes; Gaussian, exponential, algebraic, hat, and smooth hat kernels are considered as specific examples. Via a center manifold analysis, and using the relative spatial scale of competition to facilitation and the death rate as bifurcation parameters, we identify that the choice of kernel has impacts on the pattern forming bifurcation. Bifurcations using the Gaussian, exponential, and algebraic kernels largely follow expectations of Turing patterns, but patterns in the hat and smooth hat kernels can form even when the scale of competition is less than that of facilitation. The dynamics of patterns far from onset are investigated via numerical continuation methods. The model produces the so-called "Turing-before-Tipping" phenomenon demonstrating that the arrangement into spatial patterns is an effective resilience mechanism against harsh conditions. Again, there is a kernel-dependent dichotomy in pattern behavior. Early warning signs for population extinction are observed with the Gaussian, exponential, and algebraic kernels, but not under the hat or smooth hat cases.

math.DS

Pattern Formation in a Spatial Public Goods Dilemma due to Diffusive or Directed Motion

The costly provision of public goods serves as a model problem for the evolution of cooperative behavior, presenting a social dilemma between the collective benefits of shared resources and the individual incentive to free-ride in resource production. The spatial structure of populations can also impact cooperation over public goods, as diffusion of public goods and intentional motion of individuals towards regions with greater resources can interact with population and public goods dynamics to produce heterogeneous patterns in the spatial distribution of strategies and resources. In this paper, we build off a model introduced by Young and Belmonte for the reaction dynamics of interacting individuals and explicit public good, deriving a system of PDEs that describes the spatial profiles of strategies and the public good in the presence of both diffusive motion of individuals and resources and chemotaxis-like directed motion of individuals in response to gradients in the concentration of public goods. Through linear stability analysis, we show that spatial patterns in strategic and public goods profiles can emerge due to either Turing instability with high defector diffusivity or a directed-motion instability through strong sensitivity of cooperators towards increasing resource concentration. We further explore the emergent spatial patterns with a mix of weakly nonlinear stability analysis and numerical simulation, showing that, for a wide range of reaction parameters, diffusion-driven instability appears to increase cooperation and public goods across the spatial domain, while directed motion of cooperators towards public goods tends to decrease cooperation and environmental quality across the environment.

q-bio.PE

Pattern Formation in Agent-Based and PDE Models for Evolutionary Games with Payoff-Driven Motion

Spatial structure can play an important role in the evolution of cooperative behavior and the achievement of collective success of a population. In this paper, we explore the role of random and directed motion on spatial pattern formation and the payoff achieved by populations in both stochastic and deterministic models of spatial populations who engage in social interactions following a hawk-dove game. For the case of purely diffusive motion, both a stochastic spatial model and a partial differential equation model show that Turing patterns can emerge when hawks have a greater movement rate than doves, and in both models hawks and doves see an increase in population size and average payoff as hawk mobility increases. For the case of the payoff-driven motion, the stochastic model shows an overall decrease in population size and average payoff, but the PDE model displays more subtle behavior in this setting and will depend on the relative diffusivities of the two strategies. The PDE model also displays a biologically infeasible short-wave instability in the case of payoff-driven motion and equal diffusivities, indicating that we need to be careful about the mathematical properties of PDE models with payoff-driven directed motion and indicating potential use for nonlocal PDE models for spatial patterns in evolutionary games with directed motion.

q-bio.PE

Spatial Pattern Formation in Eco-Evolutionary Games with Environment-Driven Motion

The sustainable management of common resources often leads to a social dilemma known as the tragedy of the commons: individuals benefit from rapid extraction of resources, while communities as a whole benefit from more sustainable extraction strategies. Such a social dilemma can be further complicated by the role played by space for both resources and harvesters, where spatial diffusion of resources and directed motion of harvesters can potentially feature the emergence of clusters of environmental resource and sustainable harvesting strategies. In this paper, we explore a PDE model of evolutionary game theory with environmental feedback, describing how the spatial distribution of resource extraction strategies and environmental resources can change due to both local eco-evolutionary dynamics and environmental-driven directed motion of harvesters. Through linear stability analysis, we show that this biased motion towards higher-quality environments can lead to spatial patterns in the distribution of extraction strategies, creating local regions with improved environmental quality and increase payoff for resource extractors. However, by measuring the average payoff and environmental quality across the spatial domain, we see that this pattern-forming mechanism can actually decrease the overall success of the population relative to the equilibrium outcome in the absence of spatial motion. This suggests that environmental-driven motion can produce a spatial social dilemma, in which biased motion towards more beneficial regions can produce emergent patterns featuring a worse overall environment for the population.

q-bio.PE

Exploring the Evolution of Altruistic Punishment with a PDE Model of Cultural Multilevel Selection

Two mechanisms that have been used to study the evolution of cooperative behavior are altruistic punishment, in which cooperative individuals pay additional costs to punish defection, and multilevel selection, in which competition between groups can help to counteract individual-level incentives to cheat. Boyd, Gintis, Bowles, and Richerson have used simulation models of cultural evolution to suggest that altruistic punishment and pairwise group-level competition can work in concert to promote cooperation, even when neither mechanism can do so on its own. In this paper, we formulate a PDE model for multilevel selection motivated by the approach of Boyd and coauthors, modeling individual-level birth-death competition with a replicator equation based on individual payoffs and describing group-level competition with pairwise conflicts based on differences in the average payoffs of the competing groups. Building off of existing PDE models for multilevel selection with frequency-independent group-level competition, we use analytical and numerical techniques to understand how the forms of individual and average payoffs can impact the long-time ability to sustain altruistic punishment in group-structured populations. We find several interesting differences between the behavior of our new PDE model with pairwise group-level competition and existing multilevel PDE models, including the observation that our new model can feature a non-monotonic dependence of the long-time collective payoff on the strength of altruistic punishment. Going forward, our PDE framework can serve as a way to connect and compare disparate approaches for understanding multilevel selection across the literature in evolutionary biology and anthropology.

q-bio.PE

Steady-State and Dynamical Behavior of a PDE Model of Multilevel Selection with Pairwise Group-Level Competition

Evolutionary competition often occurs simultaneously at multiple levels of organization, in which traits or behaviors that are costly for an individual can provide collective benefits to groups to which the individual belongs. Building off of recent work that has used ideas from game theory to study evolutionary competition within and among groups, we study a PDE model for multilevel selection that considers group-level evolutionary dynamics through a pairwise conflict depending on the strategic composition of the competing groups. This model allows for incorporation of group-level frequency dependence, facilitating the exploration for how the form of probabilities for victory in a group-level conflict can impact the long-time support for cooperation via multilevel selection. We characterize well-posedness properties for measure-valued solutions of our PDE model and apply these properties to show that the population will converge to a delta-function at the all-defector equilibrium when between-group selection is sufficiently weak. We further provide necessary conditions for the existence of bounded steady state densities for the multilevel dynamics of Prisoners' Dilemma and Hawk-Dove scenarios, using a mix of analytical and numerical techniques to characterize the relative strength of between-group selection required to ensure the long-time survival of cooperation via multilevel selection. We also see that the average payoff at steady state appears to be limited by the average payoff of the all-cooperator group, even for games in which groups achieve maximal average payoff at intermediate levels of cooperation, generalizing behavior that has previously been observed in PDE models of multilevel selection with frequency-indepdent group-level competition.

q-bio.PE

Evolutionary Dynamics Within and Among Competing Groups

Biological and social systems are structured at multiple scales, and the incentives of individuals who interact in a group may diverge from the collective incentive of the group as a whole. Mechanisms to resolve this tension are responsible for profound transitions in evolutionary history, including the origin of cellular life, multi-cellular life, and even societies. Here we synthesize a growing literature that extends evolutionary game theory to describe multilevel evolutionary dynamics, using nested birth-death processes and partial differential equations to model natural selection acting on competition within and among groups of individuals. We apply this theory to analyze how mechanisms known to promote cooperation within a single group -- including assortment, reciprocity, and population structure -- alter evolutionary outcomes in the presence of competition among groups. We find that population structures most conducive to cooperation in multi-scale systems may differ from those most conducive within a single group. Likewise, for competitive interactions with a continuous range of strategies we find that among-group selection may fail to produce socially optimal outcomes, but it can nonetheless produce second-best solutions that balance individual incentives to defect with the collective incentives for cooperation. We conclude by describing the broad applicability of multi-scale evolutionary models to problems ranging from the production of diffusible metabolites in microbes to the management of common-pool resources in human societies.

q-bio.PE

Social dilemmas of sociality due to beneficial and costly contagion

Levels of sociality in nature vary widely. Some species are solitary; others live in family groups; some form complex multi-family societies. Increased levels of social interaction can allow for the spread of useful innovations and beneficial information, but can also facilitate the spread of harmful contagions, such as infectious diseases. It is natural to assume that these contagion processes shape the evolution of complex social systems, but an explicit account of the dynamics of sociality under selection pressure imposed by contagion remains elusive. We consider a model for the evolution of sociality strategies in the presence of both a beneficial and costly contagion. We study the dynamics of this model at three timescales: using a susceptible-infectious-susceptible (SIS) model to describe contagion spread for given sociality strategies, a replicator equation to study the changing fractions of two different levels of sociality, and an adaptive dynamics approach to study the long-time evolution of the population level of sociality. For a wide range of assumptions about the benefits and costs of infection, we identify a social dilemma: the evolutionarily-stable sociality strategy (ESS) is distinct from the collective optimum -- the level of sociality that would be best for all individuals. In particular, the ESS level of social interaction is greater (respectively less) than the social optimum when the good contagion spreads more (respectively less) readily than the bad contagion. Our results shed light on how contagion shapes the evolution of social interaction, but reveals that evolution may not necessarily lead populations to social structures that are good for any or all.

physics.soc-ph

How do classroom-turnover times depend on lecture-hall size?

Academic spaces in colleges and universities span classrooms for 10 students to lecture halls that hold over 600 people. During the break between consecutive classes, students from the first class must leave and the new class must find their desks, regardless of whether the room holds 10 or 600 people. Here we address the question of how the size of large lecture halls affects classroom-turnover times, focusing on non-emergency settings. By adapting the established social-force model, we treat students as individuals who interact and move through classrooms to reach their destinations. We find that social interactions and the separation time between consecutive classes strongly influence how long it takes entering students to reach their desks, and that these effects are more pronounced in larger lecture halls. While the median time that individual students must travel increases with decreased separation time, we find that shorter separation times lead to shorter classroom-turnover times overall. This suggests that the effects of scheduling gaps and lecture-hall size on classroom dynamics depends on the perspective - individual student or whole class - that one chooses to take.

physics.soc-ph

Assortment and Reciprocity Mechanisms for Promotion of Cooperation in a Model of Multilevel Selection

In the study of the evolution of cooperation, many mechanisms have been proposed to help overcome the self-interested cheating that is individually optimal in the Prisoners' Dilemma game. These mechanisms include assortative or networked social interactions, other-regarding preferences considering the payoffs of others, reciprocity rules to establish cooperation as a social norm, and multilevel selection involving simultaneous competition between individuals favoring cheaters and competition between groups favoring cooperators. In this paper, we build on recent work studying PDE replicator equations for multilevel selection to understand how within-group mechanisms of assortment, other-regarding preferences, and both direct and indirect reciprocity can help to facilitate cooperation in concert with evolutionary competition between groups. We consider a group-structured population in which interactions between individuals consist of Prisoners' Dilemma games, and study the dynamics of multilevel competition determined by the payoffs individuals receive when interacting according to these within-group mechanisms. We find that the presence of each of these mechanisms acts synergistically with multilevel selection for the promotion of cooperation, decreasing the strength of between-group competition required to sustain long-time cooperation and increasing the collective payoff achieved by the population. However, we find that only other-regarding preferences allows for the achievement of socially optimal collective payoffs for Prisoners' Dilemma games in which average payoff is maximized by an intermediate mix of cooperators and defectors. For the other three mechanisms, the multilevel dynamics remain susceptible to a shadow of lower-level selection, as the collective outcome fails to exceed the payoff of the all-cooperator group.

physics.soc-ph

Long-Time Behavior of a PDE Replicator Equation for Multilevel Selection in Group-Structured Populations

In many biological systems, natural selection acts simultaneously on multiple levels of organization. This scenario typically presents an evolutionary conflict between the incentive of individuals to cheat and the collective incentive to establish cooperation within a group. Generalizing previous work on multilevel selection in evolutionary game theory, we consider a hyperbolic PDE model of a group-structured population, in which members within a single group compete with each other for individual-level replication; while the group also competes against other groups for group-level replication. We derive a threshold level of the relative strength of between-group competition such that defectors take over the population below the threshold while cooperation persists in the long-time population above the threshold. Under stronger assumptions on the initial distribution of group compositions, we further prove that the population converges to a steady state density supporting cooperation for between-group selection strength above the threshold. We further establish long-time bounds on the time-average of the collective payoff of the population, showing that the long-run population cannot outperform the payoff of a full-cooperator group even in the limit of infinitely-strong between-group competition. When the group replication rate is maximized by an intermediate level of within-group cooperation, individual-level selection casts a long shadow on the dynamics of multilevel selection: no level of between-group competition can erase the effects of the individual incentive to defect. We further extend our model to study the case of multiple types of groups, showing how the games that groups play can coevolve with the level of cooperation.

q-bio.PE

A PDE Model for Protocell Evolution and the Origin of Chromosomes via Multilevel Selection

The evolution of complex cellular life involved two major transitions: the encapsulation of self-replicating genetic entities into cellular units and the aggregation of individual genes into a collectively replicating genome. In this paper, we formulate a minimal model of the evolution of proto-chromosomes within protocells. We model a simple protocell composed of two types of genes: a "fast gene" with an advantage for gene-level self-replication and a "slow gene" that replicates more slowly at the gene level, but which confers an advantage for protocell-level reproduction. Protocell-level replication capacity depends on cellular composition of fast and slow genes. We use a partial differential equation to describe how the composition of genes within protocells evolves over time under within-cell and between-cell competition. We find that the gene-level advantage of fast replicators casts a long shadow on the multilevel dynamics of protocell evolution: no level of between-protocell competition can produce coexistence of the fast and slow replicators when the two genes are equally needed for protocell-level reproduction. By introducing a "dimer replicator" consisting of a linked pair of the slow and fast genes, we show analytically that coexistence between the two genes can be promoted in pairwise multilevel competition between fast and dimer replicators, and provide numerical evidence for coexistence in trimorphic competition between fast, slow, and dimer replicators. Our results suggest that dimerization, or the formation of a simple chromosome-like dimer replicator, can help to overcome the shadow of lower-level selection and work in concert with deterministic multilevel selection to allow for the coexistence of two genes that are complementary at the protocell-level but compete at the level of individual gene-level replication.

q-bio.PE

Analysis of Multilevel Replicator Dynamics for General Two-Strategy Social Dilemmas

Here we consider a game theoretic model of multilevel selection in which individuals compete based on their payoff and groups also compete based on the average payoff of group members. Our focus is on multilevel social dilemmas: games in which individuals are best off cheating, while groups of individuals do best when composed of many cooperators. We analyze the dynamics of the two-level replicator dynamics, a nonlocal hyperbolic PDE describing deterministic birth-death dynamics for both individuals and groups. While past work on such multilevel dynamics has restricted attention to scenarios with exactly solvable within-group dynamics, we use comparison principles and an invariant property of the tail of the population distribution to extend our analysis to all possible two-player, two-strategy social dilemmas. We characterize the threshold level of between-group selection dividing a regime in which the population converges to a delta function at the equilibrium of the within-group dynamics from a regime in which between-group competition facilitates the existence of steady-state densities supporting greater levels of cooperation. In particular, we see that the threshold selection strength and average payoff at steady state depend on a tug-of-war between the individual-level incentive to be a defector in a many-cooperator group and the group-level incentive to have many cooperators over many defectors. We also find that lower-level selection casts a long shadow: if groups are best off with a mix of cooperators and defectors, then there will always be fewer cooperators than optimal at steady state, even in the limit of infinitely strong competition between groups.

q-bio.PE

The Replicator Dynamics for Multilevel Selection in Evolutionary Games

We consider a stochastic model for evolution of group-structured populations in which interactions between group members correspond to the Prisoner's Dilemma or the Hawk-Dove game. Selection operates at two organization levels: individuals compete with peer group members based on individual payoff, while groups also compete with other groups based on average payoff of group members. In the Prisoner's Dilemma, this creates a tension between the two levels of selection, as defectors are favored at the individual level, whereas groups with at least some cooperators outperform groups of defectors at the between-group level. In the limit of infinite group size and infinite number of groups, we derive a non-local PDE that describes the probability distribution of group compositions in the population. For special families of payoff matrices, we characterize the long-time behavior of solutions of our equation, finding a threshold level of between-group selection required to sustain density steady states and the survival of cooperation. When all-cooperator groups are most fit, the average and most abundant group composition at steady state range from featuring all-defector groups when individual-level selection dominates to featuring all-cooperator groups when group-level selection dominates. When the most fit groups have a mix of cooperators and defectors, then the average and most abundant group compositions always feature a smaller fraction of cooperators than required for the optimal mix, even in the limit where group-level selection is infinitely stronger than individual-level selection. In such cases, the conflict between the two levels of selection cannot be decoupled, and cooperation cannot be sustained at all in the case when between-group competition favors an even mix of cooperators and defectors.

q-bio.PE