Emergence of order in selection-mutation dynamics

We characterize the time evolution of a d-dimensional probability distribution by the value of its final entropy. If it is near the maximally-possible value we call the evolution mixing, if it is near zero we say it is purifying. The evolution is determined by the simplest non-linear equation and contains a d times d matrix as input. Since we are not interested in a particular evolution but in the general features of evolutions of this type, we take the matrix elements as uniformly-distributed random numbers between zero and some specified upper bound. Computer simulations show how the final entropies are distributed over this field of random numbers. The result is that the distribution crowds at the maximum entropy, if the upper bound is unity. If we restrict the dynamical matrices to certain regions in matrix space, for instance to diagonal or triangular matrices, then the entropy distribution is maximal near zero, and the dynamics typically becomes purifying.Comment: 8 pages, 8 figure

Spatial Invasion of Cooperation

The evolutionary puzzle of cooperation describes situations where cooperators provide a fitness benefit to other individuals at some cost to themselves. Under Darwinian selection, the evolution of cooperation is a conundrum, whereas non-cooperation (or defection) is not. In the absence of supporting mechanisms, cooperators perform poorly and decrease in abundance. Evolutionary game theory provides a powerful mathematical framework to address the problem of cooperation using the prisoner's dilemma. One well-studied possibility to maintain cooperation is to consider structured populations, where each individual interacts only with a limited subset of the population. This enables cooperators to form clusters such that they are more likely to interact with other cooperators instead of being exploited by defectors. Here we present a detailed analysis of how a few cooperators invade and expand in a world of defectors. If the invasion succeeds, the expansion process takes place in two stages: first, cooperators and defectors quickly establish a local equilibrium and then they uniformly expand in space. The second stage provides good estimates for the global equilibrium frequencies of cooperators and defectors. Under hospitable conditions, cooperators typically form a single, ever growing cluster interspersed with specks of defectors, whereas under more hostile conditions, cooperators form isolated, compact clusters that minimize exploitation by defectors. We provide the first quantitative assessment of the way cooperators arrange in space during invasion and find that the macroscopic properties and the emerging spatial patterns reveal information about the characteristics of the underlying microscopic interactions.MathematicsOrganismic and Evolutionary Biolog

Synergy and Discounting of Cooperation in Social Dilemmas

The emergence and maintenance of cooperation by natural selection is an enduring conundrum in evolutionary biology, which has been studied using a variety of game theoretical models inspired by different biological situations. The most widely studied games are the Prisoner's Dilemma, the Snowdrift game and by-product mutualism for pairwise interactions, as well as Public Goods games in larger groups of interacting individuals. Here, we present a general framework for cooperation in social dilemmas in which all the traditional scenarios can be recovered as special cases. In social dilemmas, cooperators provide a benefit to the group at some cost, while defectors exploit the group by reaping the benefits without bearing the costs of cooperation. Using the concepts of discounting and synergy for describing how benefits accumulate when more than one cooperator is present in a group of interacting individuals, we recover the four basic scenarios of evolutionary dynamics given by (i) dominating defection, (ii) coexistence of defectors and cooperators, (iii) dominating cooperation and (iv) bi-stability, in which cooperators and defectors cannot invade each other. Generically, for groups of three or more interacting individuals further, more complex, dynamics can occur. Our framework provides the first unifying approach to model cooperation in different kinds of social dilemmas.Mathematic

Evolutionary Dynamics on Graphs: Efficient Method for Weak Selection

Investigating the evolutionary dynamics of game theoretical interactions in populations where individuals are arranged on a graph can be challenging in terms of computation time. Here, we propose an efficient method to study any type of game on arbitrary graph structures for weak selection. In this limit, evolutionary game dynamics represents a first-order correction to neutral evolution. Spatial correlations can be empirically determined under neutral evolution and provide the basis for formulating the game dynamics as a discrete Markov process by incorporating a detailed description of the microscopic dynamics based on the neutral correlations. This framework is then applied to one of the most intriguing questions in evolutionary biology: the evolution of cooperation. We demonstrate that the degree heterogeneity of a graph impedes cooperation and that the success of tit for tat depends not only on the number of rounds but also on the degree of the graph. Moreover, considering the mutation-selection equilibrium shows that the symmetry of the stationary distribution of states under weak selection is skewed in favor of defectors for larger selection strengths. In particular, degree heterogeneity—a prominent feature of scale-free networks—generally results in a more pronounced increase in the critical benefit-to-cost ratio required for evolution to favor cooperation as compared to regular graphs. This conclusion is corroborated by an analysis of the effects of population structures on the fixation probabilities of strategies in general 2×2 games for different types of graphs. Computer simulations confirm the predictive power of our method and illustrate the improved accuracy as compared to previous studies.MathematicsOrganismic and Evolutionary Biolog

Overlay Accuracy Limitations of Soft Stamp UV Nanoimprint Lithography and Circumvention Strategies for Device Applications

In this work multilevel pattering capabilities of Substrate Conformal Imprint Lithography (SCIL) have been explored. A mix & match approach combining the high throughput of nanoimprint lithography with the excellent overlay accuracy of electron beam lithography (EBL) has been exploited to fabricate nanoscale devices. An EBL system has also been utilized as a benchmarking tool to measure both stamp distortions and alignment precision of this mix & match approach. By aligning the EBL system to 20 mm x 20 mm and 8 mm x 8 mm cells to compensate pattern distortions of order of $3 \mu m$ over 6 inch wafer area, overlay accuracy better than $1.2 \mu m$ has been demonstrated. This result can partially be attributed to the flexible SCIL stamp which compensates deformations caused by the presence of particles which would otherwise significantly reduce the alignment precision

Evolutionary prisoner's dilemma games with optional participation

Competition among cooperators, defectors, and loners is studied in an evolutionary prisoner's dilemma game with optional participation. Loners are risk averse i.e. unwilling to participate and rather rely on small but fixed earnings. This results in a rock-scissors-paper type cyclic dominance of the three strategies. The players are located either on square lattices or random regular graphs with the same connectivity. Occasionally, every player reassesses its strategy by sampling the payoffs in its neighborhood. The loner strategy efficiently prevents successful spreading of selfish, defective behavior and avoids deadlocks in states of mutual defection. On square lattices, Monte Carlo simulations reveal self-organizing patterns driven by the cyclic dominance, whereas on random regular graphs different types of oscillatory behavior are observed: the temptation to defect determines whether damped, periodic or increasing oscillations occur. These results are compared to predictions by pair approximation. Although pair approximation is incapable of distinguishing the two scenarios because of the equal connectivity, the average frequencies as well as the oscillations on random regular graphs are well reproduced.Comment: 6 pages, 7 figure

Public Goods with Punishment and Abstaining in Finite and Infinite Populations

The evolution and maintenance of cooperation in human and animal societies challenges various disciplines ranging from evolutionary biology, to anthropology, social sciences and economics. In social interactions, cooperators increase the welfare of the group at some cost to themselves whereas defectors attempt to free-ride and neither provide benefits nor incur costs. The problem of cooperation becomes even more pronounced when increasing the number of interacting individuals. Punishment and voluntary participation have been identified as possible factors to support cooperation and prevent cheating. Typically, punishment behavior is unable to gain a foothold in a population, while volunteering alone can efficiently prevent deadlocks in states of mutual defection but is unable to stabilize cooperation. The combined effects of the two mechanisms have surprisingly different consequences in finite and infinite populations. Here we provide a detailed comparison of the two scenarios and demonstrate that driven by the inherent stochasticity of finite populations, the possibility to abstain from social interactions plays a pivotal role, which paves the way for the establishment of cooperation and punishment.Organismic and Evolutionary Biolog

Via Freedom to Coercion: The Emergence of Costly Punishment

In human societies, cooperative behavior in joint enterprises is often enforced through institutions that impose sanctions on defectors. Many experiments on so-called public goods games have shown that in the absence of such institutions, individuals are willing to punish defectors, even at a cost to themselves. Theoretical models confirm that social norms prescribing the punishment of uncooperative behavior are stable—once established, they prevent dissident minorities from spreading. But how can such costly punishing behavior gain a foothold in the population? A surprisingly simple model shows that if individuals have the option to stand aside and abstain from the joint endeavor, this paves the way for the emergence and establishment of cooperative behavior based on the punishment of defectors. Paradoxically, the freedom to withdraw from the common enterprise leads to enforcement of social norms. Joint enterprises that are compulsory rather than voluntary are less likely to lead to cooperation.MathematicsOrganismic and Evolutionary BiologyMathematic