Biology subjects

Erol Akcay

Publications and source records attributed to Erol Akcay.

7 recordsLinked to original sources

bioRxiv2016

Learning to cooperate: The evolution of social rewards in repeated interactions

Understanding the behavioral and psychological mechanisms underlying social behaviors is one of the major goals of social evolutionary theory. In particular, a persistent question about animal cooperation is to what extent it is supported by other-regarding preferences. In many situations, animals adjust their behaviors through learning by responding to the rewards they experience as a consequence of their actions. Therefore, we may ask whether learning in social situations can be driven by evolved prosocial rewards. Here we develop a mathematical model in order to ask whether the mere act of cooperating with a social partner will evolve to be inherently rewarding. Individuals interact repeatedly in pairs and adjust their behaviors through reinforcement learning. We assume that individuals associate to each game outcome an internal reward value. These perceived rewards are genetically evolving traits. We find that conditionally cooperative rewards that value mutual cooperation positively but the suckers outcome negatively tend to be evolutionarily stable. Purely other-regarding rewards can evolve only under special parameter combinations. On the other hand, selfish rewards that always learn pure defection are also evolutionary successful. These findings are consistent with empirical observations showing that humans tend to show conditionally cooperative behavior, and also exhibit diversity of preferences. Our model also demonstrates the need to further integrate multiple levels of biological causation of behavior.

Evolutionary Biology↗

bioRxiv2016

Population structure reduces the benefits from partner choice in mutualism

Mutualisms are key drivers of evolutionary and ecological processes. Understanding how different species can evolve to interact in mutually beneficial ways is an important goal of evolutionary theory, especially when the benefits require costly investments by the partners. For such costly investments to evolve, some sort of fitness feedback mechanism must exist that more than recoups the direct costs. Several such feedback mechanisms have been explored both theoretically and empirically, yet we know relatively little how they might act together, as they probably do in nature. In this paper, I model the joint action of three of the main mechanisms that can maintain symbiotic cooperation: partner choice by hosts, population structure amongst symbionts, and undirected rewards from hosts to symbionts. My results show that population structure reduces the benefit from partner choice to hosts. It may help or hinder beneficial symbionts and create positive or negative frequency dependence depending on the nature of host rewards to the symbiont. Strong population structure also makes it less likely that host choosiness and symbiont cooperation will be jointly maintained in a population. The intuition behind my results is that all else being equal, population structure reduces local variation available to the host to choose from. Thus, population structure is not always beneficial for the evolution of cooperation between species. My results also underscore the need to do full analyses of multiple mechanisms of social evolution to uncover their interactions to uncover the interactions between them.

Evolutionary Biology↗

bioRxiv2016

The genealogical decomposition of a matrix population model with applications to the aggregation of stages

Matrix projection models are a central tool in many areas of population biology. In most applications, one starts from the projection matrix to quantify the asymptotic growth rate of the population (the dominant eigenvalue), the stable stage distribution, and the reproductive values (the dominant right and left eigenvectors, respectively). Any primitive projection matrix also has an associated ergodic Markov chain that contains information about the genealogy of the population. In this paper, we show that these facts can be used to specify any matrix population model as a triple consisting of the ergodic Markov matrix, the dominant eigenvalue and one of the corresponding eigenvectors. This decomposition of the projection matrix separates properties associated with lineages from those associated with individuals. It also clarifies the relationships between many quantities commonly used to describe such models, including the relationship between eigenvalue sensitivities and elasticities. We illustrate the utility of such a decomposition by introducing a new method for aggregating classes in a matrix population model to produce a simpler model with a smaller number of classes. Unlike the standard method, our method has the advantage of preserving reproductive values and elasticities. It also has conceptually satisfying properties such as commuting with changes of units.

Ecology↗

bioRxiv2016

Selective pressures on C4 photosynthesis evolution in grasses through the lens of optimality

CO2, temperature, water availability and light intensity were all potential selective pressures to propel the initial evolution and global expansion of C4 photosynthesis over the last 30 million years. To tease apart how the primary selective pressures varied along this evolutionary trajectory, we coupled photosynthesis and hydraulics models while optimizing photosynthesis over stomatal resistance and leaf/fine-root allocation. We further examined the importance of resource (e.g. nitrogen) reallocation from the dark to the light reactions during and after the initial formation of C4 syndrome. We show here that the primary selective pressures--all acting upon photorespiration in C3 progenitors--changed through the course of C4 evolution. The higher stomatal resistance and leaf-to-root allocation ratio enabled by the C4 carbon-concentrating mechanism led to a C4 advantage without any change in hydraulic properties, but selection on nitrogen reallocation varied. Water limitation was the primary driver for the initial evolution of C4 25-32 million years ago, and could positively select for C4 evolution with atmospheric CO2 as high as 600 ppm. Under these high CO2 conditions, nitrogen reallocation was necessary. Low CO2 and light intensity, but not nitrogen reallocation, were the primary drivers during the global radiation of C4 5-10 MYA. Finally, our results suggest that identifying the predominate selective pressures at the time C4 first evolved within a lineage should help explain current biogeographical distributions.\n\nStatement of authorshipHZ, BH and EA conceptualized the study. HZ and EA built the model, HZ and BH put the idea in a general evolutionary context, HZ performed the modeling work and analyzed output data. HZ wrote the first draft, BH and EA contributed substantially to revisions.\n\nSignificance StatementC4 photosynthesis pathway had evolved more than 60 times independently across the terrestrial plants through mid-Oligocene (~30 MYA) and diversified at late Miocene (5 to 10 MYA). We use an optimal physiology model to examine the primary selective pressures along the evolutionary history. Water limitation was the primary driver for C4 evolution from the initial evolutionary events 25-32 MYA until CO2 became low enough to, along with light intensity, drive the global radiation of C4 5-10 MYA. This modeling framework can be used to investigate evolution of other physiological traits (e.g. N reallocation, hydraulics) after the initial formation of C4 syndrome, which contributed to further increasing productivity of C4 in historical and current environmental conditions.

Evolutionary Biology↗

bioRxiv2016

Invasion fitness, inclusive fitness, and reproductive numbers in heterogeneous populations

How should fitness be measured to determine which phenotype or \"strategy\" is uninvadable when evolution occurs in subdivided populations subject to local demographic and environmental heterogeneity? Several invasion fitness measures, such as basic reproductive number, lifetime dispersal success of a local lineage, or inclusive fitness have been proposed to address this question, but the relationships between them and their generality remains unclear. Here, we ascertain uninvadability (all mutant strategies always go extinct) in terms of the growth rate of a mutant allele arising as a single copy in a population. We show from this growth rate that uninvadability is equivalently characterized by at least three conceptually distinct invasion fitness measures: (i) lineage fitness, giving the average personal fitness of a randomly sampled mutant lineage member; (ii) inclusive fitness, giving a reproductive value weighted average of the direct fitness cost and relatedness weighted indirect fitness benefits accruing to a randomly sampled mutant lineage member; and (iii) three types of reproductive numbers, giving lifetime success of a local lineage. Our analysis connects approaches that have been deemed different, generalizes the exact version of inclusive fitness to class-structured populations, and provides a biological interpretation of selection on a mutant allele under arbitrary strength of selection.

Preprint↗

bioRxiv2015

Social inheritance can explain the structure of animal social networks

The social network structure of animal populations has major implications for survival, reproductive success, sexual selection, and pathogen transmission of individuals. But as of yet, no general theory of social network structure exists that can explain the diversity of social networks observed in nature, and serve as a null model for detecting species and population-specific factors. Here we propose a simple and generally applicable model of social network structure. We consider the emergence of network structure as a result of social inheritance, in which newborns are likely to bond with maternal contacts, and via forming bonds randomly. We compare model output to data from several species, showing that it can generate networks with properties such as those observed in real social systems. Our model demonstrates that important observed properties of social networks, including heritability of network position or assortative associations, can be understood as consequences of social inheritance.

Animal Behavior and Cognition↗

bioRxiv2013

Pathways to social evolution: reciprocity, relatedness, and synergy

Many organisms live in populations structured by space and by class, exhibit plastic responses to their social partners, and are subject to non-additive ecological and fitness effects. Social evolution theory has long recognized that all of these factors can lead to different selection pressures but has only recently attempted to synthesize how these factors interact. Using models for both discrete and continuous phenotypes, we show that analyzing these factors in a consistent framework reveals that they interact with one another in ways previously overlooked. Specifically, behavioral responses (reciprocity), genetic relatedness, and synergy interact in non-trivial ways that cannot be easily captured by simple summary indices of assortment. We demonstrate the importance of these interactions by showing how they have been neglected in previous synthetic models of social behavior both within and between species. These interactions also affect the level of behavioral responses that can evolve in the long run; proximate biological mechanisms are evolutionarily stable when they generate enough responsiveness relative to the level of responsiveness that exactly balances the ecological costs and benefits. Given the richness of social behavior across taxa, these interactions should be a boon for empirical research as they are likely crucial for describing the complex relationship linking ecology, demography, and social behavior.

Evolutionary Biology↗