Furthermore, cooperators may interact more locally than defectors. This tilt is because the individual contributes to the public good if it is a cooperator, and detracts from the collective public good if it is a defector. These finite neighborhoods create a form of assortative interaction: an individual always experiences a slightly higher frequency of its own strategy within a neighborhood than present in the entire population. Such interactions can strongly influence the eco-evolutionary dynamics between cooperators (public good producers) and defectors (free-riders).īenefits that are produced and shared are often only available within a finite neighborhood 21, 22. It benefits a cancer cell to recruit blood vessels, signal normal cells and defend against the immune system, which benefit neighbors too. As ecological engineers cancer cells can promote favorable environments 16, 17, suggesting that they evolve cooperation 18, 19, or form mutualistic relationships with other cells 20. Cancer cells, too, can engage in public goods games 13– 15. Yeast consuming and synthesizing amino acids for themselves and others were also observed to coevolve in the form of a different social dilemma, the snowdrift game 11, in which stable coexistence can exist with or without assortment 12. Yeast synthesize and spill essential nutrients, such as key amino acids into their surroundings 10. Microbes provide public goods by secreting defensive chemicals 8, 9. Via associational refuges, plants defended by spines or toxins may dissuade herbivores from feeding on any plants in a neighborhood 6, 7. Predator inspection by guppies 4, or mobbing of hawks by crows 5 provide safety to self and others regardless of participation. Prairie dogs maintain sight lines around their burrows that become available to others in the colony 3. In banded mongooses, an individual in the foraging line flushes insects for others 2. Nature offers many examples where individuals provide public goods that benefit self and others, which can be maintained by population assortment 1. Our findings advance the understanding of how neighborhood-size effects in PGG shape the dynamics of growing populations. We integrate our model with experiments of cancer cell growth and confirm that our framework describes PGG dynamics observed in cellular populations. Stochastic or strategy-dependent variations in neighborhood sizes favor coexistence by destabilizing monomorphic states. This mechanism leads to diverse evolutionarily stable strategies, including monomorphic and polymorphic populations, and neighborhood-size-driven state changes, resulting in hysteresis between equilibria. At high cooperation, increases provide diminishing returns. At low cooperation, increases in the public good provide increasing returns. We model the population dynamics of a non-linear PGG and consider density-dependence on the global level, while the game occurs within local neighborhoods. Public goods games (PGG) cover the essence of such dilemmas in which cooperators are prone to exploitation by defectors. An evolutionary game emerges when a subset of individuals incur costs to provide benefits to all individuals.
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