Recent work has revealed a new class of “zero-determinant” (ZD) strategies for iterated, two-player games. ZD strategies allow a player to unilaterally enforce a linear relationship between her score and her opponent’s score, and thus to achieve an unusual degree of control over both players’ long-term payoffs. Although originally conceived in the context of classical two-player game theory, ZD strategies also have consequences in evolving populations of players. Here, we explore the evolutionary prospects for ZD strategies in the Iterated Prisoner’s Dilemma (IPD). Several recent studies have focused on the evolution of “extortion strategies,” a subset of ZD strategies, and have found them to be unsuccessful in populations. Nevertheless, we identify a different subset of ZD strategies, called “generous ZD strategies,” that forgive defecting opponents but nonetheless dominate in evolving populations. For all but the smallest population sizes, generous ZD strategies are not only robust to being replaced by other strategies but can selectively replace any noncooperative ZD strategy. Generous strategies can be generalized beyond the space of ZD strategies, and they remain robust to invasion. When evolution occurs on the full set of all IPD strategies, selection disproportionately favors these generous strategies. In some regimes, generous strategies outperform even the most successful of the well-known IPD strategies, including win-stay-lose-shift. Significance Cooperative behavior seems at odds with the Darwinian principle of survival of the fittest, yet cooperation is abundant in nature. Scientists have used the Prisoner Dilemma game, in which players must choose to cooperate or defect, to study the emergence and stability of cooperation. Recent work has uncovered a remarkable class of extortion strategies that provide one player a disproportionate payoff when facing an unwitting opponent. Extortion strategies perform very well in head-to-head competitions, but they fare poorly in large, evolving populations. Rather we identify a closely related set of generous strategies, which cooperate with others and forgive defection, that replace extortionists and dominate in large populations. Our results help to explain the evolution of cooperation.

Significance: This study offers a new perspective on an age-old question: When does cooperation emerge in populations? Two-player games used to study this question produce an array of counterintuitive results. And yet a consensus has emerged that, in an evolving population, cooperation tends to triumph over cheating––through reciprocity and generosity. But, what happens when players can influence not only their tendencies to cooperate, but also the rewards they reap for cooperation? We analyze coevolution of strategies and payoffs and find that, as individuals maximize the benefits of cooperation, they often pave the way for its collapse. Our analysis provides a framework for studying the coevolution of games and strategies, and suggests that maintaining cooperation may be more difficult than previously thought.

Gene expression is controlled primarily by transcription factors, whose DNA binding sites are typically 10 nt long. We develop a population-genetic model to understand how the length and information content of such binding sites evolve. Our analysis is based on an inherent trade-off between specificity, which is greater in long binding sites, and robustness to mutation, which is greater in short binding sites. The evolutionary stable distribution of binding site lengths predicted by the model agrees with the empirical distribution (5–31 nt, with mean 9.9 nt for eukaryotes), and it is remarkably robust to variation in the underlying parameters of population size, mutation rate, number of transcription factor targets, and strength of selection for proper binding and selection against improper binding. In a systematic data set of eukaryotic and prokaryotic transcription factors we also uncover strong relationships between the length of a binding site and its information content per nucleotide, as well as between the number of targets a transcription factor regulates and the information content in its binding sites. Our analysis explains these features as well as the remarkable conservation of binding site characteristics across diverse taxa.

Polarization can spread and become entrenched when inequality creates subpopulations that cannot afford risks.