arXiv subjects

Martin A. Nowak

Publications and source records attributed to Martin A. Nowak.

At least 37 records · Page 2Linked to original sources

arXiv2019

Fixation probability and fixation time in structured populations

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.

q-bio.PE↗

arXiv2018

Environmental fitness heterogeneity in the Moran process

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.

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arXiv2018

Stationary frequencies and mixing times for neutral drift processes with spatial structure

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.

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arXiv2018

Evolution of Cooperation on Stochastic Block Models

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.

cs.SI↗

arXiv2018

Conjoining uncooperative societies facilitates evolution of cooperation

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.

cs.SI↗

arXiv2018

Strong Amplifiers of Natural Selection: Proofs

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 $n^{1-ε}$, for some fixed $ε>0$, can be assigned weights that makes it a strong amplifier, both for uniform and temperature initialization.

cs.DM↗

arXiv2018

Selection for synchronized cell division in simple multicellular organisms

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.

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arXiv2018

Public goods games in populations with fluctuating size

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.

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arXiv2018

Variational auto-encoding of protein sequences

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.

q-bio.QM↗

arXiv2017

Primordial Sex Facilitates the Emergence of Evolution

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.

q-bio.PE↗

arXiv2017

Effects of motion in structured populations

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.

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arXiv2017

Faster Monte-Carlo Algorithms for Fixation Probability of the Moran Process on Undirected Graphs

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.

cs.NE↗

arXiv2017

The evolution of queen control over worker reproduction in the social Hymenoptera

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.

q-bio.PE↗

arXiv2017

Evolutionary games on cycles with strong selection

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.

q-bio.PE↗

arXiv2016

Evolutionary dynamics on any population structure

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.

q-bio.PE↗

arXiv2016

Fixation probabilities for any configuration of two strategies on regular graphs

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.

q-bio.PE↗

arXiv2016

Games of multicellularity

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.

q-bio.PE↗

arXiv2016

Comparing reactive and memory-one strategies of direct reciprocity

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.

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