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Direct reciprocity is a wide-spread mechanism for evolution of cooperation. In repeated interactions, players can condition their behavior on previous outcomes. A well known approach is given by reactive strategies, which respond to the co-player's previous move. Here we extend reactive strategies to longer memories. A reactive-$n$ strategy takes into account the sequence of the last $n$ moves of the co-player. A reactive-$n$ counting strategy records how often the co-player has cooperated during the last $n$ rounds. We derive an algorithm to identify all partner strategies among reactive-$n$ strategies. We give explicit conditions for all partner strategies among reactive-2, reactive-3 strategies, and reactive-$n$ counting strategies. Partner strategies are those that ensure mutual cooperation without exploitation. We perform evolutionary simulations and find that longer memory increases the average cooperation rate for reactive-$n$ strategies but not for reactive counting strategies. Paying attention to the sequence of moves is necessary for reaping the advantages of longer memory.

Computing the rate of evolution in spatially structured populations is difficult. A key quantity is the fixation time of a single mutant with relative reproduction rate $r$ which invades a population of residents. We say that the fixation time is "fast" if it is at most a polynomial function in terms of the population size $N$. Here we study fixation times of advantageous mutants ($r>1$) and neutral mutants ($r=1$) on directed graphs, which are those graphs that have at least some one-way connections. We obtain three main results. First, we prove that for any directed graph the fixation time is fast, provided that $r$ is sufficiently large. Second, we construct an efficient algorithm that gives an upper bound for the fixation time for any graph and any $r\ge 1$. Third, we identify a broad class of directed graphs with fast fixation times for any $r\ge 1$. This class includes previously studied amplifiers of selection, such as Superstars and Metafunnels. We also show that on some graphs the fixation time is not a monotonically declining function of $r$; in particular, neutral fixation can occur faster than fixation for small selective advantages., Comment: 22 pages, 5 figures

Direct reciprocity is a powerful mechanism for evolution of cooperation based on repeated interactions between the same individuals. But high levels of cooperation evolve only if the benefit-to-cost ratio exceeds a certain threshold that depends on memory length. For the best-explored case of one-round memory, that threshold is two. Here we report that intermediate mutation rates lead to high levels of cooperation, even if the benefit-to-cost ratio is only marginally above one, and even if individuals only use a minimum of past information. This surprising observation is caused by two effects. First, mutation generates diversity which undermines the evolutionary stability of defectors. Second, mutation leads to diverse communities of cooperators that are more resilient than homogeneous ones. This finding is relevant because many real world opportunities for cooperation have small benefit-to-cost ratios, which are between one and two, and we describe how direct reciprocity can attain cooperation in such settings. Our result can be interpreted as showing that diversity, rather than uniformity, promotes evolution of cooperation., Comment: 37 pages, 14 figures

In social systems subject to indirect reciprocity, a positive reputation is key for increasing one's likelihood of future positive interactions. The flow of gossip can amplify the impact of a person's actions on their reputation depending on how widely it spreads across the social network, which leads to a percolation problem. To quantify this notion, we calculate the expected number of individuals, the "audience", who find out about a particular interaction. For a potential donor, a larger audience constitutes higher reputational stakes, and thus a higher incentive, to perform "good" actions in line with current social norms. For a receiver, a larger audience therefore increases the trust that the partner will be cooperative. This idea can be used for an algorithm that generates social networks, which we call trust based attachment (TBA). TBA produces graphs that share crucial quantitative properties with real-world networks, such as high clustering, small-world behavior, and power law degree distributions. We also show that TBA can be approximated by simple friend-of-friend routines based on triadic closure, which are known to be highly effective at generating realistic social network structures. Therefore, our work provides a new justification for triadic closure in social contexts based on notions of trust, gossip, and social information spread. These factors are thus identified as potential significant influences on how humans form social ties.

Evolution occurs in populations of reproducing individuals. It is well known that population structure can affect evolutionary dynamics. Traditionally, natural selection is studied between mutants that differ in reproductive rate, but are subject to the same population structure. Here we study how natural selection acts on mutants that have the same reproductive rate, but experience different population structures. In our framework, mutation alters population structure, which is given by a graph that specifies the dispersal of offspring. Reproduction can be either genetic or cultural. Competing mutants disperse their offspring on different graphs. A more connected graph implies higher motility. We show that enhanced motility tends to increase an invader's fixation probability, but there are interesting exceptions. For island models, we show that the magnitude of the effect depends crucially on the exact layout of the additional links. Finally, we show that for low-dimensional lattices, the effect of altered motility is comparable to that of altered fitness: in the limit of large population size, the invader's fixation probability is either constant or exponentially small, depending on whether it is more or less motile than the resident., Comment: 20 pages, 11 figures

Motivated by COVID-19, we develop and analyze a simple stochastic model for a disease spread in human population. We track how the number of infected and critically ill people develops over time in order to estimate the demand that is imposed on the hospital system. To keep this demand under control, we consider a class of simple policies for slowing down and reopening the society and we compare their efficiency in mitigating the spread of the virus from several different points of view. We find that in order to avoid overwhelming of the hospital system, a policy must impose a harsh lockdown or it must react swiftly (or both). While reacting swiftly is universally beneficial, being harsh pays off only when the country is patient about reopening and when the neighboring countries coordinate their mitigation efforts. Our work highlights the importance of acting decisively when closing down and the importance of patience and coordination between neighboring countries when reopening.

Resources are rarely distributed uniformly within a population. Heterogeneity in the concentration of a drug, the quality of breeding sites, or wealth can all affect evolutionary dynamics. In this study, we represent a collection of properties affecting the fitness at a given location using a color. A green node is rich in resources while a red node is poorer. More colors can represent a broader spectrum of resource qualities. For a population evolving according to the birth-death Moran model, the first question we address is which structures, identified by graph connectivity and graph coloring, are evolutionarily equivalent. We prove that all properly two-colored, undirected, regular graphs are evolutionarily equivalent (where "properly colored" means that no two neighbors have the same color). We then compare the effects of background heterogeneity on properly two-colored graphs to those with alternative schemes in which the colors are permuted. Finally, we discuss dynamic coloring as a model for spatiotemporal resource fluctuations, and we illustrate that random dynamic colorings often diminish the effects of background heterogeneity relative to a proper two-coloring., Comment: 19 pages

Evolutionary Game Theory (EGT) simulations are used to model populations undergoing biological and cultural evolution in a range of fields, from biology to economics to linguistics. In this paper we present DyPy, an open source Python package that can perform evolutionary simulations for any matrix form game for three common evolutionary dynamics: Moran, Wright-Fisher and Replicator. We discuss the basic components of this package and illustrate how it can be used to run a variety of simulations. Our package allows a user to run such simulations fairly easily without much prior Python knowledge. We hope that this will be a great asset to researchers in a number of different fields.

Prosocial behaviors are encountered in the donation game, the prisoner's dilemma, relaxed social dilemmas, and public goods games. Many studies assume that the population structure is homogeneous, meaning all individuals have the same number of interaction partners, or that the social good is of one particular type. Here, we explore general evolutionary dynamics for arbitrary spatial structures and social goods. We find that heterogeneous networks, wherein some individuals have many more interaction partners than others, can enhance the evolution of prosocial behaviors. However, they often accumulate most of the benefits in the hands of a few highly-connected individuals, while many others receive low or negative payoff. Surprisingly, selection can favor producers of social goods even if the total costs exceed the total benefits. In summary, heterogeneous structures have the ability to strongly promote the emergence of prosocial behaviors, but they also create the possibility of generating large inequality., Comment: 72 pages; final version

A great number of biological organisms live in aqueous environments. Major evolutionary transitions, including the emergence of life itself, likely occurred in such environments. While the chemical aspects of the role of water in biology are well-studied, the effects of water's physical characteristics on evolutionary events, such as the control of population structure via its rich transport properties, are less clear. Evolutionary transitions such as the emergence of the first cells and of multicellularity, require cooperation among groups of individuals. However, evolution of cooperation faces challenges in unstructured "well-mixed" populations, as parasites quickly overwhelm cooperators. Models that assume population structure to promote cooperation envision such structure to arise from spatial "lattice" models (e.g. surface bound individuals) or compartmentalization models, often realized as protocells. Here we study the effect of turbulent motions in spatial models, and propose that coherent structures, i.e. flow patterns which trap fluid and arise naturally in turbulent flows, may serve many of the properties associated with compartments--collocalization, division, and merging--and thought to play a key role in the origins of life and other evolutionary transitions. These results suggest that group selection models may be applicable with fewer physical and chemical constraints than previously thought, and apply much more widely in aqueous environments.

The fixation probability of a single mutant invading a population of residents is among the most widely-studied quantities in evolutionary dynamics. Amplifiers of natural selection are population structures that increase the fixation probability of advantageous mutants, compared to well-mixed populations. Extensive studies have shown that many amplifiers exist for the Birth-death Moran process, some of them substantially increasing the fixation probability or even guaranteeing fixation in the limit of large population size. On the other hand, no amplifiers are known for the death-Birth Moran process, and computer-assisted exhaustive searches have failed to discover amplification. In this work we resolve this disparity, by showing that any amplification under death-Birth updating is necessarily \emph{bounded} and \emph{transient}. Our boundedness result states that even if a population structure does amplify selection, the resulting fixation probability is close to that of the well-mixed population. Our transience result states that for any population structure there exists a threshold $r^*$ such that the population structure ceases to amplify selection if the mutant fitness advantage $r$ is larger than $r^\star$. Finally, we also extend the above results to $\delta$-death-Birth updating, which is a combination of Birth-death and death-Birth updating. On the positive side, we identify population structures that maintain amplification for a wide range of values $r$ and $\delta$. These results demonstrate that amplification of natural selection depends on the specific mechanisms of the evolutionary process.

The environment has a strong influence on a population's evolutionary dynamics. Driven by both intrinsic and external factors, the environment is subject to continual change in nature. To capture an ever-changing environment, we consider a model of evolutionary dynamics with game transitions, where individuals' behaviors together with the games they play in one time step influence the games to be played next time step. Within this model, we study the evolution of cooperation in structured populations and find a simple rule: weak selection favors cooperation over defection if the ratio of the benefit provided by an altruistic behavior, $b$, to the corresponding cost, $c$, exceeds $k-k'$, where $k$ is the average number of neighbors of an individual and $k'$ captures the effects of the game transitions. Even if cooperation cannot be favored in each individual game, allowing for a transition to a relatively valuable game after mutual cooperation and to a less valuable game after defection can result in a favorable outcome for cooperation. In particular, small variations in different games being played can promote cooperation markedly. Our results suggest that simple game transitions can serve as a mechanism for supporting prosocial behaviors in highly-connected populations., Comment: 44 pages, 10 figures; final version

Population structure affects the outcome of natural selection. Static population structures can be described by graphs, where individuals occupy the nodes, and interactions occur along the edges. General conditions for evolutionary success on any weighted graph were recently derived, for weak selection, in terms of coalescence times of random walks. Here we show that for a special class of graphs, the conditions for success take a particularly simple form, in which all effects of graph structure are described by the graph's "effective degree"---a measure of the effective number of neighbors per individual. This result holds for all weighted graphs that are isothermal, meaning that the sum of edge weights is the same at each node. Isothermal graphs encompass a wide variety of underlying topologies, and arise naturally from supposing that each individual devotes the same amount of time to interaction. Cooperative behavior is favored on a large isothermal graph if the benefit-to-cost ratio exceeds the effective degree. We relate the effective degree of a graph to its spectral gap, thereby providing a link between evolutionary dynamics and the theory of expander graphs. As a surprising example, we report graphs of infinite average degree that are nonetheless highly conducive for promoting cooperation., Comment: 40 pages, 6 figures

In an iterated game between two players, there is much interest in characterizing the set of feasible payoffs for both players when one player uses a fixed strategy and the other player is free to switch. Such characterizations have led to extortionists, equalizers, partners, and rivals. Most of those studies use memory-one strategies, which specify the probabilities to take actions depending on the outcome of the previous round. Here, we consider "reactive learning strategies," which gradually modify their propensity to take certain actions based on past actions of the opponent. Every linear reactive learning strategy, $\mathbf{p}^{\ast}$, corresponds to a memory one-strategy, $\mathbf{p}$, and vice versa. We prove that for evaluating the region of feasible payoffs against a memory-one strategy, $\mathcal{C}\left(\mathbf{p}\right)$, we need to check its performance against at most $11$ other strategies. Thus, $\mathcal{C}\left(\mathbf{p}\right)$ is the convex hull in $\mathbb{R}^{2}$ of at most $11$ points. Furthermore, if $\mathbf{p}$ is a memory-one strategy, with feasible payoff region $\mathcal{C}\left(\mathbf{p}\right)$, and $\mathbf{p}^{\ast}$ is the corresponding reactive learning strategy, with feasible payoff region $\mathcal{C}\left(\mathbf{p}^{\ast}\right)$, then $\mathcal{C}\left(\mathbf{p}^{\ast}\right)$ is a subset of $\mathcal{C}\left(\mathbf{p}\right)$. Reactive learning strategies are therefore powerful tools in restricting the outcomes of iterated games., Comment: 18 pages

The rate of biological evolution depends on the fixation probability and on the fixation time of new mutants. Intensive research has focused on identifying population structures that augment the fixation probability of advantageous mutants. But these `amplifiers of natural selection' typically increase fixation time. Here we study population structures that achieve a trade-off between high fixation probability and short fixation time. First, we show that no amplifiers can have asymptotically lower absorption time than the well-mixed population. Then we design population structures that substantially augment the fixation probability with just a minor increase in fixation time. Finally, we show that those structures enable higher effective rate of evolution than the well-mixed population provided that the rate of generating advantageous mutants is relatively low. Our work sheds light on how population structure affects the rate of evolution. Moreover, our structures could be useful for lab-based, medical or industrial applications of evolutionary optimization.

We study a general setting of neutral evolution in which the population is of finite, constant size and can have spatial structure. Mutation leads to different genetic types ("traits"), which can be discrete or continuous. Under minimal assumptions, we show that the marginal trait distributions of the evolutionary process, which specify the probability that any given individual has a certain trait, all converge to the stationary distribution of the mutation process. In particular, the stationary frequencies of traits in the population are independent of its size, spatial structure, and evolutionary update rule, and these frequencies can be calculated by evaluating a simple stochastic process describing a population of size one (i.e. the mutation process itself). We conclude by analyzing mixing times, which characterize rates of convergence of the mutation process along the lineages, in terms of demographic variables of the evolutionary process., Comment: 18 pages

Cooperation is a major factor in the evolution of human societies. The structure of human social networks, which affects the dynamics of cooperation and other interpersonal phenomena, have common structural signatures. One of these signatures is the tendency to organize as groups. Among the generative models that network theorists use to emulate this feature is the Stochastic Block Model (SBM). In this paper, we study evolutionary game dynamics on SBM networks. Using a recently-discovered duality between evolutionary games and coalescing random walks, we obtain analytical conditions such that natural selection favors cooperation over defection. We calculate the transition point for each community to favor cooperation. We find that a critical inter-community link creation probability exists for given group density, such that the overall network supports cooperation even if individual communities inhibit it. As a byproduct, we present mean-field solutions for the critical benefit-to-cost ratio which perform with remarkable accuracy for diverse generative network models, including those with community structure and heavy-tailed degree distributions. We also demonstrate the generalizability of the results to arbitrary two-player games.

Social structure affects the emergence and maintenance of cooperation. Here we study the evolutionary dynamics of cooperation in fragmented societies, and show that conjoining segregated cooperation-inhibiting groups, if done properly, rescues the fate of collective cooperation. We highlight the essential role of inter-group ties, that sew the patches of the social network together and facilitate cooperation. We point out several examples of this phenomenon in actual settings. We explore random and non-random graphs, as well as empirical networks. In many cases we find a marked reduction of the critical benefit-to-cost ratio needed for sustaining cooperation. Our finding gives hope that the increasing worldwide connectivity, if managed properly, can promote global cooperation.

The evolution of multicellularity was a major transition in the history of life on earth. Conditions under which multicellularity is favored have been studied theoretically and experimentally. But since the construction of a multicellular organism requires multiple rounds of cell division, a natural question is whether these cell divisions should be synchronous or not. We study a simple population model in which there compete simple multicellular organisms that grow either by synchronous or asynchronous cell divisions. We demonstrate that natural selection can act differently on synchronous and asynchronous cell division, and we offer intuition for why these phenotypes are generally not neutral variants of each other.

We consider the modified Moran process on graphs to study the spread of genetic and cultural mutations on structured populations. An initial mutant arises either spontaneously (aka \emph{uniform initialization}), or during reproduction (aka \emph{temperature initialization}) in a population of $n$ individuals, and has a fixed fitness advantage $r>1$ over the residents of the population. The fixation probability is the probability that the mutant takes over the entire population. Graphs that ensure fixation probability of~1 in the limit of infinite populations are called \emph{strong amplifiers}. Previously, only a few examples of strong amplifiers were known for uniform initialization, whereas no strong amplifiers were known for temperature initialization. In this work, we study necessary and sufficient conditions for strong amplification, and prove negative and positive results. We show that for temperature initialization, graphs that are unweighted and/or self-loop-free have fixation probability upper-bounded by $1-1/f(r)$, where $f(r)$ is a function linear in $r$. Similarly, we show that for uniform initialization, bounded-degree graphs that are unweighted and/or self-loop-free have fixation probability upper-bounded by $1-1/g(r,c)$, where $c$ is the degree bound and $g(r,c)$ a function linear in $r$. Our main positive result complements these negative results, and is as follows: every family of undirected graphs with (i)~self loops and (ii)~diameter bounded by $n^{1-\epsilon}$, for some fixed $\epsilon>0$, can be assigned weights that makes it a strong amplifier, both for uniform and temperature initialization.

Proteins are responsible for the most diverse set of functions in biology. The ability to extract information from protein sequences and to predict the effects of mutations is extremely valuable in many domains of biology and medicine. However the mapping between protein sequence and function is complex and poorly understood. Here we present an embedding of natural protein sequences using a Variational Auto-Encoder and use it to predict how mutations affect protein function. We use this unsupervised approach to cluster natural variants and learn interactions between sets of positions within a protein. This approach generally performs better than baseline methods that consider no interactions within sequences, and in some cases better than the state-of-the-art approaches that use the inverse-Potts model. This generative model can be used to computationally guide exploration of protein sequence space and to better inform rational and automatic protein design., Comment: Abstract for oral presentation at NIPS 2017 Workshop on Machine Learning in Computational Biology

Many mathematical frameworks of evolutionary game dynamics assume that the total population size is constant and that selection affects only the relative frequency of strategies. Here, we consider evolutionary game dynamics in an extended Wright-Fisher process with variable population size. In such a scenario, it is possible that the entire population becomes extinct. Survival of the population may depend on which strategy prevails in the game dynamics. Studying cooperative dilemmas, it is a natural feature of such a model that cooperators enable survival, while defectors drive extinction. Although defectors are favored for any mixed population, random drift could lead to their elimination and the resulting pure-cooperator population could survive. On the other hand, if the defectors remain, then the population will quickly go extinct because the frequency of cooperators steadily declines and defectors alone cannot survive. In a mutation-selection model, we find that (i) a steady supply of cooperators can enable long-term population survival, provided selection is sufficiently strong, and (ii) selection can increase the abundance of cooperators but reduce their relative frequency. Thus, evolutionary game dynamics in populations with variable size generate a multifaceted notion of what constitutes a trait's long-term success., Comment: 21 pages; final version

Many mathematical models of evolution assume that all individuals experience the same environment. Here, we study the Moran process in heterogeneous environments. The population is of finite size with two competing types, which are exposed to a fixed number of environmental conditions. Reproductive rate is determined by both the type and the environment. We first calculate the condition for selection to favor the mutant relative to the resident wild type. In large populations, the mutant is favored if and only if the mutant's spatial average reproductive rate exceeds that of the resident. But environmental heterogeneity elucidates an interesting asymmetry between the mutant and the resident. Specifically, mutant heterogeneity suppresses its fixation probability; if this heterogeneity is strong enough, it can even completely offset the effects of selection (including in large populations). In contrast, resident heterogeneity has no effect on a mutant's fixation probability in large populations and can amplify it in small populations., Comment: 23 pages, 9 figures; final version

In evolutionary processes, population structure has a substantial effect on natural selection. Here, we analyze how motion of individuals affects constant selection in structured populations. Motion is relevant because it leads to changes in the distribution of types as mutations march toward fixation or extinction. We describe motion as the swapping of individuals on graphs, and more generally as the shuffling of individuals between reproductive updates. Beginning with a one-dimensional graph, the cycle, we prove that motion suppresses natural selection for death-birth updating or for any process that combines birth-death and death-birth updating. If the rule is purely birth-death updating, no change in fixation probability appears in the presence of motion. We further investigate how motion affects evolution on the square lattice and weighted graphs. In the case of weighted graphs we find that motion can be either an amplifier or a suppressor of natural selection. In some cases, whether it is one or the other can be a function of the relative reproductive rate, indicating that motion is a subtle and complex attribute of evolving populations. As a first step towards understanding less restricted types of motion in evolutionary graph theory, we consider a similar rule on dynamic graphs induced by a spatial flow and find qualitatively similar results indicating that continuous motion also suppresses natural selection., Comment: 25 pages; final version

Evolutionary graph theory studies the evolutionary dynamics in a population structure given as a connected graph. Each node of the graph represents an individual of the population, and edges determine how offspring are placed. We consider the classical birth-death Moran process where there are two types of individuals, namely, the residents with fitness 1 and mutants with fitness r. The fitness indicates the reproductive strength. The evolutionary dynamics happens as follows: in the initial step, in a population of all resident individuals a mutant is introduced, and then at each step, an individual is chosen proportional to the fitness of its type to reproduce, and the offspring replaces a neighbor uniformly at random. The process stops when all individuals are either residents or mutants. The probability that all individuals in the end are mutants is called the fixation probability. We present faster polynomial-time Monte-Carlo algorithms for finidng the fixation probability on undirected graphs. Our algorithms are always at least a factor O(n^2/log n) faster as compared to the previous algorithms, where n is the number of nodes, and is polynomial even if r is given in binary. We also present lower bounds showing that the upper bound on the expected number of effective steps we present is asymptotically tight for undirected graphs.

A trademark of eusocial insect species is reproductive division of labor, in which workers forego their own reproduction while the queen produces almost all offspring. The presence of the queen is key for maintaining social harmony, but the specific role of the queen in the evolution of eusociality remains unclear. A long-discussed scenario is that a queen either behaviorally or chemically sterilizes her workers. However, the demographic and ecological conditions that enable such manipulation are unknown. Accordingly, we propose a simple model of evolutionary dynamics that is based on haplodiploid genetics. We consider a mutation that acts in a queen, causing her to control the reproductive behavior of her workers. Our mathematical analysis yields precise conditions for the evolutionary emergence and stability of queen-induced worker sterility. These conditions do not depend on the queen's mating frequency. Moreover, we find that queen control is always established if it increases colony reproductive efficiency and can evolve even if it decreases colony efficiency. We further outline the conditions under which queen control is evolutionarily stable against invasion by mutant, reproductive workers.

Evolutionary games on graphs describe how strategic interactions and population structure determine evolutionary success, quantified by the probability that a single mutant takes over a population. Graph structures, compared to the well-mixed case, can act as amplifiers or suppressors of selection by increasing or decreasing the fixation probability of a beneficial mutant. Properties of the associated mean fixation times can be more intricate, especially when selection is strong. The intuition is that fixation of a beneficial mutant happens fast (in a dominance game), that fixation takes very long (in a coexistence game), and that strong selection eliminates demographic noise. Here we show that these intuitions can be misleading in structured populations. We analyze mean fixation times on the cycle graph under strong frequency-dependent selection for two different microscopic evolutionary update rules (death-birth and birth-death). We establish exact analytical results for fixation times under strong selection, and show that there are coexistence games in which fixation occurs in time polynomial in population size. Depending on the underlying game, we observe inherence of demographic noise even under strong selection, if the process is driven by random death before selection for birth of an offspring (death-birth update). In contrast, if selection for an offspring occurs before random removal (birth-death update), strong selection can remove demographic noise almost entirely., Comment: 5 Figures. Accepted for publication (Physical Review E)

Compartments are ubiquitous throughout biology, yet their importance stretches back to the origin of cells. In the context of origin of life, we assume that a protocell, a compartment enclosing functional components, requires $N$ components to be evolvable. We take interest in the timescale in which a minimal evolvable protocell is produced. We show that when protocells fuse and share information, the time to produce an evolvable protocell scales algebraically in $N$, in contrast to an exponential scaling in the absence of fusion. We discuss the implications of this result for origins of life, as well as other biological processes., Comment: Second version with minor reference updates

Evolutionary game dynamics are often studied in the context of different population structures. Here we propose a new population structure that is inspired by simple multicellular life forms. In our model, cells reproduce but can stay together after reproduction. They reach complexes of a certain size, n, before producing single cells again. The cells within a complex derive payoff from an evolutionary game by interacting with each other. The reproductive rate of cells is proportional to their payoff. We consider all two-strategy games. We study deterministic evolutionary dynamics with mutations, and derive exact conditions for selection to favor one strategy over another. Our main result has the same symmetry as the well-known sigma condition, which has been proven for stochastic game dynamics and weak selection. For a maximum complex size of n=2 our result holds for any intensity of selection. For n > 2 it holds for weak selection. As specific examples we study the prisoner's dilemma and hawk-dove games. Our model advances theoretical work on multicellularity by allowing for frequency-dependent interactions within groups., Comment: 44 pages, 11 figues

Evolution occurs in populations of reproducing individuals. The structure of a biological population affects which traits evolve. Understanding evolutionary game dynamics in structured populations is difficult. Precise results have been absent for a long time, but have recently emerged for special structures where all individuals have the same number of neighbors. But the problem of determining which trait is favored by selection in the natural case where the number of neighbors can vary, has remained open. For arbitrary selection intensity, the problem is in a computational complexity class which suggests there is no efficient algorithm. Whether there exists a simple solution for weak selection was unanswered. Here we provide, surprisingly, a general formula for weak selection that applies to any graph or social network. Our method uses coalescent theory and relies on calculating the meeting times of random walks. We can now evaluate large numbers of diverse and heterogeneous population structures for their propensity to favor cooperation. We can also study how small changes in population structure---graph surgery---affect evolutionary outcomes. We find that cooperation flourishes most in societies that are based on strong pairwise ties., Comment: 68 pages, 10 figures

Population structure and spatial heterogeneity are integral components of evolutionary dynamics, in general, and of evolution of cooperation, in particular. Structure can promote the emergence of cooperation in some populations and suppress it in others. Here, we provide results for weak selection to favor cooperation on regular graphs for any configuration, meaning any arrangement of cooperators and defectors. Our results extend previous work on fixation probabilities of single, randomly placed mutants. We find that for any configuration cooperation is never favored for birth-death (BD) updating. In contrast, for death-birth (DB) updating, we derive a simple, computationally tractable formula for weak selection to favor cooperation when starting from any configuration containing any number of cooperators and defectors. This formula elucidates two important features: (i) the takeover of cooperation can be enhanced by the strategic placement of cooperators and (ii) adding more cooperators to a configuration can sometimes suppress the evolution of cooperation. These findings give a formal account for how selection acts on all transient states that appear in evolutionary trajectories. They also inform the strategic design of initial states in social networks to maximally promote cooperation. We also derive general results that characterize the interaction of any two strategies, not only cooperation and defection., Comment: 28 pages; final version

Workers in insect societies are sometimes observed to kill male eggs of other workers, a phenomenon known as worker policing. We perform a mathematical analysis of the evolutionary dynamics of policing. We investigate the selective forces behind policing for both dominant and recessive mutations for different numbers of matings of the queen. The traditional, relatedness-based argument suggests that policing evolves if the queen mates with more than two males, but does not evolve if the queen mates with a single male. We derive precise conditions for the invasion and stability of policing alleles. We find that the relatedness-based argument is not robust with respect to small changes in colony efficiency caused by policing. We also calculate evolutionarily singular strategies and determine when they are evolutionarily stable. We use a population genetics approach that applies to dominant or recessive mutations of any effect size.

Direct reciprocity is a mechanism for the evolution of cooperation based on repeated interactions. When individuals meet repeatedly, they can use conditional strategies to enforce cooperative outcomes that would not be feasible in one-shot social dilemmas. Direct reciprocity requires that individuals keep track of their past interactions and find the right response. However, there are natural bounds on strategic complexity: Humans find it difficult to remember past interactions accurately, especially over long timespans. Given these limitations, it is natural to ask how complex strategies need to be for cooperation to evolve. Here, we study stochastic evolutionary game dynamics in finite populations to systematically compare the evolutionary performance of reactive strategies, which only respond to the co-player's previous move, and memory-one strategies, which take into account the own and the co-player's previous move. In both cases, we compare deterministic strategy and stochastic strategy spaces. For reactive strategies and small costs, we find that stochasticity benefits cooperation, because it allows for generous-tit-for-tat. For memory one strategies and small costs, we find that stochasticity does not increase the propensity for cooperation, because the deterministic rule of win-stay, lose-shift works best. For memory one strategies and large costs, however, stochasticity can augment cooperation., Comment: 18 pages, 7 figures

Most cancers in humans are large, measuring centimeters in diameter, composed of many billions of cells. An equivalent mass of normal cells would be highly heterogeneous as a result of the mutations that occur during each cell division. What is remarkable about cancers is their homogeneity - virtually every neoplastic cell within a large cancer contains the same core set of genetic alterations, with heterogeneity confined to mutations that have emerged after the last clonal expansions. How such clones expand within the spatially-constrained three dimensional architecture of a tumor, and come to dominate a large, pre-existing lesion, has never been explained. We here describe a model for tumor evolution that shows how short-range migration and cell turnover can account for rapid cell mixing inside the tumor. With it, we show that even a small selective advantage of a single cell within a large tumor allows the descendants of that cell to replace the precursor mass in a clinically relevant time frame. We also demonstrate that the same mechanisms can be responsible for the rapid onset of resistance to chemotherapy. Our model not only provides novel insights into spatial and temporal aspects of tumor growth but also suggests that targeting short range cellular migratory activity could have dramatic effects on tumor growth rates., Comment: 37 pages, 14 figures