Em uma perspectiva evolutiva, a emergência e a manutenção de comportamentos altruísticos e de cooperação não é de fácil entendimento. O impulso por ajudar um indivíduo desconhecido não pode significar um prejuízo na capacidade reprodutiva, o que muitas vezes parece ser o mais óbvio. Muito se tem feito no sentido de compreender os ganhos indiretos da cooperação, ou o que se espera em retorno por este comportamento. A espera por reciprocidade é um dos modos de se tornar a cooperação atraente. Os seres humanos possuem uma capacidade singular de expandir a reciprocidade para interações organizadas em que não necessariamente se recebe a retribuição de um favor, mas sim o favor de um terceiro indivíduo. Para estes sistemas, de reciprocidade indireta, são necessários elaborados processos cognitivos que sustentam uma capacidade para linguagem, julgamentos morais e organização social. Entende-se que esta forma de cooperação é um fator essencial para a evolução do intelecto e da estrutura social atuais dos seres humanos. A teoria dos jogos evolucionária é uma ferramenta matemática muito utilizada na sistematização analítica dos problemas envolvendo cooperação e processos evolutivos no geral. A capacidade reprodutiva é traduzida em termos de funções matemáticas, sendo possível realizar dinâmicas populacionais que modelam a pressão seletiva. Neste trabalho, utilizamos métodos de teoria dos jogos evolucionária para explorar modelos de reciprocidade indireta, expandindo o tratamento de um modelo para interações opcionais envolvendo estratégias de cooperadores condicionais. Mostramos que a presença de cooperadores incondicionais ameaça a estabilidade da cooperação e que erros de execução podem ser uma solução. / At an evolutionary perspective, the emergence and maintenance of altruistic and cooperative behaviours is of no easy understanding. The impulse of helping an unrelated individual cannot mean a loss of reproductive fitness, as many times may seem the obvious. Much has been done in the way of knowing the indirect benefits of cooperation, or what to expect in retribution for this behaviour. To expect reciprocity is one way of looking at cooperation as more attractive. Human beings have a singular capacity of expanding reciprocity to organized interactions where retribution of a favour is not necessary, but one can expect the favour of a third-party. For these systems, of indirect reciprocity, elaborate cognitive processes are necessary, ones that maintain the capacity for language, moral judgements and social organization. One can understand this form of cooperation as an essential factor for the evolution of humans nowadays´ intellect and social structure. Evolutionary game theory is a mathematical tool that is largely used in the analytical systematization of problems involving cooperation and evolutionary processes in general. Reproductive fitness is understood in terms of mathematical functions, making possible the work on population dynamics that model selective pressure. In this work, we use methods in evolutionary game theory to explore models of indirect reciprocity, expanding the treatment of a model for optional interactions involving conditional cooperators strategies. We show that the presence of unconditional cooperators threatens the stability of cooperation and that execution errors might be a solution.

Em uma perspectiva evolutiva, a emergência e a manutenção de comportamentos altruísticos e de cooperação não é de fácil entendimento. O impulso por ajudar um indivíduo desconhecido não pode significar um prejuízo na capacidade reprodutiva, o que muitas vezes parece ser o mais óbvio. Muito se tem feito no sentido de compreender os ganhos indiretos da cooperação, ou o que se espera em retorno por este comportamento. A espera por reciprocidade é um dos modos de se tornar a cooperação atraente. Os seres humanos possuem uma capacidade singular de expandir a reciprocidade para interações organizadas em que não necessariamente se recebe a retribuição de um favor, mas sim o favor de um terceiro indivíduo. Para estes sistemas, de reciprocidade indireta, são necessários elaborados processos cognitivos que sustentam uma capacidade para linguagem, julgamentos morais e organização social. Entende-se que esta forma de cooperação é um fator essencial para a evolução do intelecto e da estrutura social atuais dos seres humanos. A teoria dos jogos evolucionária é uma ferramenta matemática muito utilizada na sistematização analítica dos problemas envolvendo cooperação e processos evolutivos no geral. A capacidade reprodutiva é traduzida em termos de funções matemáticas, sendo possível realizar dinâmicas populacionais que modelam a pressão seletiva. Neste trabalho, utilizamos métodos de teoria dos jogos evolucionária para explorar modelos de reciprocidade indireta, expandindo o tratamento de um modelo para interações opcionais envolvendo estratégias de cooperadores condicionais. Mostramos que a presença de cooperadores incondicionais ameaça a estabilidade da cooperação e que erros de execução podem ser uma solução. / At an evolutionary perspective, the emergence and maintenance of altruistic and cooperative behaviours is of no easy understanding. The impulse of helping an unrelated individual cannot mean a loss of reproductive fitness, as many times may seem the obvious. Much has been done in the way of knowing the indirect benefits of cooperation, or what to expect in retribution for this behaviour. To expect reciprocity is one way of looking at cooperation as more attractive. Human beings have a singular capacity of expanding reciprocity to organized interactions where retribution of a favour is not necessary, but one can expect the favour of a third-party. For these systems, of indirect reciprocity, elaborate cognitive processes are necessary, ones that maintain the capacity for language, moral judgements and social organization. One can understand this form of cooperation as an essential factor for the evolution of humans nowadays´ intellect and social structure. Evolutionary game theory is a mathematical tool that is largely used in the analytical systematization of problems involving cooperation and evolutionary processes in general. Reproductive fitness is understood in terms of mathematical functions, making possible the work on population dynamics that model selective pressure. In this work, we use methods in evolutionary game theory to explore models of indirect reciprocity, expanding the treatment of a model for optional interactions involving conditional cooperators strategies. We show that the presence of unconditional cooperators threatens the stability of cooperation and that execution errors might be a solution.

Diss. Stockholm : Handelshögskolan, 2009 Sammanfattning jämte 6 uppsatser

- / Doctorat en Sciences / info:eu-repo/semantics/nonPublished

Le fichiers qui accompagnent mon document ont été réalisés avec le logiciel Mathematica / Dans cette thèse, nous étudions la théorie des jeux évolutionnaires dans quelques exemples de populations structurées. En particulier, nous analysons l’évolution de la coopération en déterminant des conditions qui la favorisent dans le cas des interactions par paire. On s’intéresse à l’évolution de la coopération dans un espace phénotypique de dimension quelconque. Puis on étudie la coopération dans une population finie, subdivisée en groupes de même quelconques avec une hiérarchie entre les groupes. Finalement, on présente l’effet de l’aspiration sur le processus évolutif dans une population finie répartie sur un cercle où il y a des positions à occuper. / In this thesis, we study some examples of structured populations. In particular, we analyze the evolution of cooperation in the sense of determining conditions that favor it. We study the evolution of cooperation in a phenotype space of any size. We study also the evolution of cooperation in a finite population subdivided into hierarchical groups of any size. Finally, we study the effect of aspiration on the evolutionary process in a finite population distributed on a circle with only a local interaction by pairwise.

Cette thèse s'intéresse aux liens entre l'évolution de la coopération et la sélection de second ordre. Dans une première partie, nous montrons comment des organismes digitaux adaptent leurs génomes pour encoder les gènes liées à la coopération d'une manière plus contrainte (suppression d'évolvabilité), notamment à l'aide d'opérons et d'overlaps impliquant aussi des gènes essentiels. Dans une deuxième partie, nous testons expérimentalement cette vision des overlaps de gènes comme "contrainte évolutive" grâce à des outils d'algorithmique et de biologie synthétique que nous avons développés. Dans une troisième partie, nous utilisons des simulations par agents pour montrer comment une forme de division du travail peut être interprétée comme un système coopératif à la lumière de la théorie évolutive moderne. Dans une dernière partie, nous montrons que la dispersion spatiale des allèles coopératives obtenue par des phénomènes de "genetic hitchiking" joue un rôle important dans l'évolution de la coopération, quand bien même ce mécanisme de dispersion s'applique aussi à des allèles non coopératives, grâce à la "relatedness" (aux loci codant pour la coopération) crée par l'invasion locale de mutations bénéfiques (à des loci non liés à la coopération) et par l'équilibre complexe entre ces mutations bénéfiques et la robustesse mutationnelle. L'ensemble de ces résultats appelle à une prise en compte plus importante des pressions sélectives de second ordre dans l'étude de l'évolution sociale, et au développement de modèles plus réalistes qui permettraient d'intégrer de telles forces évolutives. Nous insistons également sur l'importance du paysage mutationnel dans l'étude des populations bactériennes, et montrons le potentiel croissant de la biologie synthétique comme outil d'étude de ce paysage et de l'évolution microbienne en général. / In the first part, I show how digital organisms adapt their genomes to encode cooperation-related genes in a more constrained way (evolvability suppression), especially using operons and overlaps also involving essential genes. In the second part, we experimentally test this view of gene overlaps as an evolutionary constraint, using both algorithmic and synthetic biology tools that we have developed. In the third part, I use agent-based simulations to show how a form of division of labour can be interpreted as a cooperative system in the light of modern evolutionary theory. In the final part, I show that the patterns of dispersal of cooperative alleles due to hitchhiking phenomena play an important role in the evolution of cooperation. The last result holds even though the hitchhiking mechanisms also applies to non-cooperative alleles, thanks to the relatedness (at cooperation-related loci) created by the local invasion of beneficial mutations (at loci not related to cooperation). The beneficial mutations form a complex and interesting equilibrium with mutational robustness, which I investigate using in silico evolution. On the whole, these results call for a more careful consideration of the second-order selection pressures in the study of social evolution, and show the necessity for more realistic models allowing to integrate such evolutionary forces. My thesis research specifically highlights the importance of the mutational landscape in the study of microbial populations and shows the increasing potential of synthetic biology as a tool to study such landscape and microbial evolution in general.

Dans cette thèse, nous étudions les effets d'un environnement stochastique et de l'utilisation d'une stratégie d'opting-out sur l'évolution de la coopération dans les jeux évolutionnaires. La thèse contient 8 articles, dont 6 sont déjà publiés dans des revues avec comité de lecture. Outre l'introduction, la thèse est divisée en deux parties, la partie 1 composée de 5 articles et la partie 2 de 3 articles. La partie 1 étudie l'impact de gains randomisés dans les jeux évolutionnaires. L'article 1 introduit les concepts de stabilité pour les jeux avec matrice de paiement aléatoire 2x2 dans des populations infinies avec des générations discrètes sans chevauchement dans un environnement stochastique. On y donne les conditions pour qu'un équilibre, sur la frontière ou à l'intérieur du simplexe des fréquences des stratégies, soit stochastiquement localement stable ou instable. L'article 2 étend les résultats de l'article 1 au cas où la valeur sélective est une fonction exponentielle du gain attendu suite à des interactions aléatoires par paires et montre que, de manière inattendue, le bruit aléatoire environnemental peut rompre un cycle périodique et favoriser la stabilité d'un équilibre intérieur. L'article 3 discute des effets de la sélection faible. Alors que les conditions de stabilité dans un environnement aléatoire reviennent aux conditions du cas déterministe lorsque l'intensité de la sélection diminue, les fluctuations aléatoires des gains peuvent accélérer la vitesse de convergence vers un équilibre stable sous une sélection plus faible. L'article 4 applique la théorie de la stabilité évolutive stochastique à un jeu randomisé de dilemme du prisonnier. On y montre que l'augmentation de la variance des gains de défection est propice à l'évolution de la coopération. L'article 5 étudie les jeux matriciels randomisés dans des populations finies et donne les conditions pour que la sélection favorise l'évolution de la coopération dans le contexte du jeu randomisé de dilemme du prisonnier. La partie 2 considère un jeu répété de dilemme du prisonnier dans le cas où un comportement d'opting-out est adopté par chaque joueur dans les interactions par paires. L'article 6 étudie la dynamique évolutive de la coopération et de la défection dans ce contexte et montre une possible coexistence à long terme, en supposant une population infinie et un équilibre rapide (en fait, instantané) dans les fréquences des paires. L'article 7 rapporte des résultats expérimentaux avec 264 étudiants universitaires utilisant la stratégie d'opting-out qui soutiennent la prédiction théorique d'une coexistence à long terme de coopération et de défection. L'article 8 étend l'analyse du modèle avec la stratégie d'opting-out au cas d'une population finie et fournit une preuve rigoureuse des deux échelles de temps pour les fréquences de coopération et de défection d'une part et les fréquences de paires de stratégies d'autre part. / In this thesis, we study the effects of a stochastic environment and the use of an opting-out strategy on the evolution of cooperation in evolutionary games. The thesis contains 8 articles, among which 6 are already published in peer-reviewed journals. Apart from the introduction, the thesis is divided into two parts, Part 1 made with 5 articles and Part 2 with 3 articles. Part 1 studies randomized payoffs in evolutionary games. Article 1 introduces stability concepts for 2x2 matrix games in infinite populations undergoing discrete, non-overlapping generations in a stochastic environment and gives conditions for an equilibrium, either on the boundary or in the interior of the simplex of all strategy frequencies, to be stochastically locally stable or unstable. Article 2 extends the results of Article 1 to the case where fitness is an exponential function of expected payoff in random pairwise interactions and shows that, unexpectedly, environmental random noise can break a periodic cycle and promote stability of an interior equilibrium. Article 3 discusses the effects of weak selection. While stability conditions in a random environment return to conditions in the deterministic case as selection intensity diminishes, random fluctuations in payoffs can accelerate the speed of convergence toward a stable equilibrium under weaker selection. Article 4 applies stochastic evolutionary stability theory to a randomized Prisoner's dilemma game and shows that increasing the variance in payoffs for defection is conducive to the evolution of cooperation. Article 5 studies randomized matrix games in finite populations and gives conditions for selection to favor the evolution of cooperation in the context of a randomized Prisoner's dilemma. Part 2 considers a repeated Prisoner's dilemma game with an opting-out behavior adopted by every player in pairwise interactions. Article 6 studies the evolutionary dynamics of cooperation and defection in this context and shows possible long-term coexistence, assuming an infinite population and fast (actually, instantaneous) equilibrium in the pair frequencies. Article 7 reports experimental results with 264 university students using the opting-out strategy that support the theoretical prediction of a long-term coexistence of cooperation and defection. Article 8 extends the analysis of the model with the opting-out strategy to the case of a finite population and provides a rigorous proof of the two-time scales for the frequencies of cooperation and defection on one hand and the frequencies of strategy pairs on the other.