Person:

Nowak, Martin

Direct reciprocity is a mechanism for the evolution of cooperation. For the iterated prisoner’s dilemma, a new class of strategies has recently been described, the so-called zero-determinant strategies. Using such a strategy, a player can unilaterally enforce a linear relationship between his own payoff and the co-player’s payoff. In particular the player may act in such a way that it becomes optimal for the co-player to cooperate unconditionally. In this way, a player can manipulate and extort his co-player, thereby ensuring that the own payoff never falls below the co-player’s payoff. However, using a compliant strategy instead, a player can also ensure that his own payoff never exceeds the co-player’s payoff. Here, we use adaptive dynamics to study when evolution leads to extortion and when it leads to compliance. We find a remarkable cyclic dynamics: in sufficiently large populations, extortioners play a transient role, helping the population to move from selfish strategies to compliance. Compliant strategies, however, can be subverted by altruists, which in turn give rise to selfish strategies. Whether cooperative strategies are favored in the long run critically depends on the size of the population; we show that cooperation is most abundant in large populations, in which case average payoffs approach the social optimum. Our results are not restricted to the case of the prisoners dilemma, but can be extended to other social dilemmas, such as the snowdrift game. Iterated social dilemmas in large populations do not lead to the evolution of strategies that aim to dominate their co-player. Instead, generosity succeeds.

We show that the times separating the birth of benign, invasive, and metastatic tumor cells can be determined by analysis of the mutations they have in common. When combined with prior clinical observations, these analyses suggest the following general conclusions about colorectal tumorigenesis: (i) It takes ≈17 years for a large benign tumor to evolve into an advanced cancer but <2 years for cells within that cancer to acquire the ability to metastasize; (ii) it requires few, if any, selective events to transform a highly invasive cancer cell into one with the capacity to metastasize; (iii) the process of cell culture ex vivo does not introduce new clonal mutations into colorectal tumor cell populations; and (iv) the rates at which point mutations develop in advanced cancers are similar to those of normal cells. These results have important implications for understanding human tumor pathogenesis, particularly those associated with metastasis.

Background: The basic idea of tag-based models for cooperation is that individuals recognize each other via arbitrary signals, so-called tags. If there are tags of different colors, then cooperators can always establish new signals of recognition. The resulting “chromodynamics” is a mechanism for the evolution of cooperation. Cooperators use a secret tag until they are discovered by defectors who then destroy cooperation based on this tag. Subsequently, a fraction of the population manages to establish cooperation based on a new tag. Methodology/Principal Findings: We derive a mathematical description of stochastic evolutionary dynamics of tag-based cooperation in populations of finite size. Benefit and cost of cooperation are given by b and c. We find that cooperators are more abundant than defectors if (b/c > 1+2u/v), where (u) is the mutation rate changing only the strategy and v is the mutation rate changing strategy and tag. We study specific assumptions for (u) and (v) in two genetic models and one cultural model. Conclusions/Significance: In a genetic model, tag-based cooperation only evolves if a gene encodes both strategy and tag. In a cultural model with equal mutation rates between all possible phenotypes (tags and behaviors), the crucial condition is (b/c > (K+1)/(K-1)), where (K) is the number of tags. A larger number of tags requires a smaller benefit-to-cost ratio. In the limit of many different tags, the condition for cooperators to have a higher average abundance than defectors becomes (b > c).

The idea of evolutionary game theory is to relate the payoff of a game to reproductive success (= fitness). An underlying assumption in most models is that fitness is a linear function of the payoff. For stochastic evolutionary dynamics in finite populations, this leads to analytical results in the limit of weak selection, where the game has a small effect on overall fitness. But this linear function makes the analysis of strong selection difficult. Here, we show that analytical results can be obtained for any intensity of selection, if fitness is defined as an exponential function of payoff. This approach also works for group selection (= multi-level selection). We discuss the difference between our approach and that of inclusive fitness theory.

Cancer results from genetic alterations that disturb the normal cooperative behavior of cells. Recent high-throughput genomic studies of cancer cells have shown that the mutational landscape of cancer is complex and that individual cancers may evolve through mutations in as many as 20 different cancer-associated genes. We use data published by Sjöblom et al. (2006) to develop a new mathematical model for the somatic evolution of colorectal cancers. We employ the Wright-Fisher process for exploring the basic parameters of this evolutionary process and derive an analytical approximation for the expected waiting time to the cancer phenotype. Our results highlight the relative importance of selection over both the size of the cell population at risk and the mutation rate. The model predicts that the observed genetic diversity of cancer genomes can arise under a normal mutation rate if the average selective advantage per mutation is on the order of 1%. Increased mutation rates due to genetic instability would allow even smaller selective advantages during tumorigenesis. The complexity of cancer progression can be understood as the result of multiple sequential mutations, each of which has a relatively small but positive effect on net cell growth.

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.

In evolutionary games the fitness of individuals is not constant but depends on the relative abundance of the various strategies in the population. Here we study general games among n strategies in populations of large but finite size. We explore stochastic evolutionary dynamics under weak selection, but for any mutation rate. We analyze the frequency dependent Moran process in well-mixed populations, but almost identical results are found for the Wright–Fisher and Pairwise Comparison processes. Surprisingly simple conditions specify whether a strategy is more abundant on average than 1/n, or than another strategy, in the mutation-selection equilibrium. We find one condition that holds for low mutation rate and another condition that holds for high mutation rate. A linear combination of these two conditions holds for any mutation rate. Our results allow a complete characterization of n×n games in the limit of weak selection.

Direct reciprocity relies on repeated encounters between the same two individuals. Here we examine the evolution of cooperation under direct reciprocity in dynamically structured populations. Individuals occupy the vertices of a graph, undergoing repeated interactions with their partners via the edges of the graph. Unlike the traditional approach to evolutionary game theory, where individuals meet at random and have no control over the frequency or duration of interactions, we consider a model in which individuals differ in the rate at which they seek new interactions. Moreover, once a link between two individuals has formed, the productivity of this link is evaluated. Links can be broken off at different rates. Whenever the active dynamics of links is sufficiently fast, population structure leads to a simple transformation of the payoff matrix, effectively changing the game under consideration, and hence paving the way for reciprocators to dominate defectors. We derive analytical conditions for evolutionary stability.

Recently, the frequency-dependent Moran process has been introduced in order to describe evolutionary game dynamics in finite populations. Here, an alternative to this process is investigated that is based on pairwise comparison between two individuals. We follow a long tradition in the physics community and introduce a temperature (of selection) to account for stochastic effects. We calculate the fixation probabilities and fixation times for any symmetric 2×2 game, for any intensity of selection and any initial number of mutants. The temperature can be used to gauge continuously from neutral drift to the extreme selection intensity known as imitation dynamics. For some payoff matrices the distribution of fixation times can become so broad that the average value is no longer very meaningful.

How fast does a population evolve from one fitness peak to another? We study the dynamics of evolving, asexually reproducing populations in which a certain number of mutations jointly confer a fitness advantage. We consider the time until a population has evolved from one fitness peak to another one with a higher fitness. The order of mutations can either be fixed or random. If the order of mutations is fixed, then the population follows a metaphorical ridge, a single path. If the order of mutations is arbitrary, then there are many ways to evolve to the higher fitness state. We address the time required for fixation in such scenarios and study how it is affected by the order of mutations, the population size, the fitness values and the mutation rate.