Evolvability and organismal architecture:The blind watchmaker and the reminiscent architect

Organisms are constantly faced with the challenge of adapting to new circumstances. In this thesis, I argue that the ability to adapt to new circumstances, “evolvability”, is deeply ingrained in the genetic, developmental, morphological, and physiological architecture of organisms. Using a blend of conceptual research, theoretical modelling, and multidisciplinary studies, I demonstrate how organismal architecture can evolve so that organisms can cope better and better with future environmental challenges. As a first step, I systematically classify the many factors contributing to evolvability. Then I use a simulation approach to show how evolvability-enhancing structures can readily evolve in gene-regulatory networks. This happens via the evolution of "mutational transformers" - structural elements that convert random mutations at the genetic level into adaptation-enhancing mutations at the phenotypic level. In another thesis chapter, I demonstrate that even if selection acts only sporadically, complex adaptations can evolve and persist over long time periods. In other words, complex adaptations do not require constant selection pressure. In an interdisciplinary contribution, I apply biological insights regarding the properties of an evolvability-enhancing mutation structure to the design of algorithms used in Artificial Intelligence. The result is the “Facilitated Mutation” method which enhances the performance of the algorithms in various respects, highlighting the potential for leveraging biological principles in computational sciences. Finally, I embed my research findings in a philosophical context. I emphasise the importance of organismal architecture in retaining evolutionary memories and suggest future research directions to further enhance our understanding of evolvability

Evolution of sex-specific pace-of-life syndromes: genetic architecture and physiological mechanisms

Sex differences in life history, physiology, and behavior are nearly ubiquitous across taxa, owing to sex-specific selection that arises from different reproductive strategies of the sexes. The pace-of-life syndrome (POLS) hypothesis predicts that most variation in such traits among individuals, populations, and species falls along a slow-fast pace-of-life continuum. As a result of their different reproductive roles and environment, the sexes also commonly differ in pace-of-life, with important consequences for the evolution of POLS. Here, we outline mechanisms for how males and females can evolve differences in POLS traits and in how such traits can covary differently despite constraints resulting from a shared genome. We review the current knowledge of the genetic basis of POLS traits and suggest candidate genes and pathways for future studies. Pleiotropic effects may govern many of the genetic correlations, but little is still known about the mechanisms involved in trade-offs between current and future reproduction and their integration with behavioral variation. We highlight the importance of metabolic and hormonal pathways in mediating sex differences in POLS traits; however, there is still a shortage of studies that test for sex specificity in molecular effects and their evolutionary causes. Considering whether and how sexual dimorphism evolves in POLS traits provides a more holistic framework to understand how behavioral variation is integrated with life histories and physiology, and we call for studies that focus on examining the sex-specific genetic architecture of this integration

Antecipação na tomada de decisão com múltiplos critérios sob incerteza

Orientador: Fernando José Von ZubenTese (doutorado) - Universidade Estadual de Campinas, Faculdade de Engenharia Elétrica e de ComputaçãoResumo: A presença de incerteza em resultados futuros pode levar a indecisões em processos de escolha, especialmente ao elicitar as importâncias relativas de múltiplos critérios de decisão e de desempenhos de curto vs. longo prazo. Algumas decisões, no entanto, devem ser tomadas sob informação incompleta, o que pode resultar em ações precipitadas com consequências imprevisíveis. Quando uma solução deve ser selecionada sob vários pontos de vista conflitantes para operar em ambientes ruidosos e variantes no tempo, implementar alternativas provisórias flexíveis pode ser fundamental para contornar a falta de informação completa, mantendo opções futuras em aberto. A engenharia antecipatória pode então ser considerada como a estratégia de conceber soluções flexíveis as quais permitem aos tomadores de decisão responder de forma robusta a cenários imprevisíveis. Essa estratégia pode, assim, mitigar os riscos de, sem intenção, se comprometer fortemente a alternativas incertas, ao mesmo tempo em que aumenta a adaptabilidade às mudanças futuras. Nesta tese, os papéis da antecipação e da flexibilidade na automação de processos de tomada de decisão sequencial com múltiplos critérios sob incerteza é investigado. O dilema de atribuir importâncias relativas aos critérios de decisão e a recompensas imediatas sob informação incompleta é então tratado pela antecipação autônoma de decisões flexíveis capazes de preservar ao máximo a diversidade de escolhas futuras. Uma metodologia de aprendizagem antecipatória on-line é então proposta para melhorar a variedade e qualidade dos conjuntos futuros de soluções de trade-off. Esse objetivo é alcançado por meio da previsão de conjuntos de máximo hipervolume esperado, para a qual as capacidades de antecipação de metaheurísticas multi-objetivo são incrementadas com rastreamento bayesiano em ambos os espaços de busca e dos objetivos. A metodologia foi aplicada para a obtenção de decisões de investimento, as quais levaram a melhoras significativas do hipervolume futuro de conjuntos de carteiras financeiras de trade-off avaliadas com dados de ações fora da amostra de treino, quando comparada a uma estratégia míope. Além disso, a tomada de decisões flexíveis para o rebalanceamento de carteiras foi confirmada como uma estratégia significativamente melhor do que a de escolher aleatoriamente uma decisão de investimento a partir da fronteira estocástica eficiente evoluída, em todos os mercados artificiais e reais testados. Finalmente, os resultados sugerem que a antecipação de opções flexíveis levou a composições de carteiras que se mostraram significativamente correlacionadas com as melhorias observadas no hipervolume futuro esperado, avaliado com dados fora das amostras de treinoAbstract: The presence of uncertainty in future outcomes can lead to indecision in choice processes, especially when eliciting the relative importances of multiple decision criteria and of long-term vs. near-term performance. Some decisions, however, must be taken under incomplete information, what may result in precipitated actions with unforeseen consequences. When a solution must be selected under multiple conflicting views for operating in time-varying and noisy environments, implementing flexible provisional alternatives can be critical to circumvent the lack of complete information by keeping future options open. Anticipatory engineering can be then regarded as the strategy of designing flexible solutions that enable decision makers to respond robustly to unpredictable scenarios. This strategy can thus mitigate the risks of strong unintended commitments to uncertain alternatives, while increasing adaptability to future changes. In this thesis, the roles of anticipation and of flexibility on automating sequential multiple criteria decision-making processes under uncertainty are investigated. The dilemma of assigning relative importances to decision criteria and to immediate rewards under incomplete information is then handled by autonomously anticipating flexible decisions predicted to maximally preserve diversity of future choices. An online anticipatory learning methodology is then proposed for improving the range and quality of future trade-off solution sets. This goal is achieved by predicting maximal expected hypervolume sets, for which the anticipation capabilities of multi-objective metaheuristics are augmented with Bayesian tracking in both the objective and search spaces. The methodology has been applied for obtaining investment decisions that are shown to significantly improve the future hypervolume of trade-off financial portfolios for out-of-sample stock data, when compared to a myopic strategy. Moreover, implementing flexible portfolio rebalancing decisions was confirmed as a significantly better strategy than to randomly choosing an investment decision from the evolved stochastic efficient frontier in all tested artificial and real-world markets. Finally, the results suggest that anticipating flexible choices has lead to portfolio compositions that are significantly correlated with the observed improvements in out-of-sample future expected hypervolumeDoutoradoEngenharia de ComputaçãoDoutor em Engenharia Elétric

The Nurture of Nature: Biology, Psychology and Culture

In this thesis I explore what consequences taking development seriously in evolutionary considerations will have for how we understand the evolution of psychology and culture. I first explicate the relationship between development and evolution that informs a number of approaches to evolution, including neo-Darwinian evolutionary biology and evolutionary developmental biology. I argue that, to a greater or lesser extent, developmental processes have been misconstrued in these accounts and that the full role of development, from an evolutionary point of view, has not always been acknowledged. Instead, I suggest that a better model of the relationship between development and evolution can be found in developmental systems theory. I explore the neo-Darwinian underpinnings of a number of accounts of the evolution of culture and psychology, including the branch of evolutionary psychology associated with the work of, among others, John Tooby and Leda Cosmides, and the gene-culture co-evolutionary account of Peter Richerson and Robert Boyd. I argue that as well as being vulnerable to the same sorts of problems that plague neo-Darwinian evolutionary biology, they face other difficulties. These accounts suppose an internalist model of the mind, and this model is neither justified nor useful. The extended mind hypothesis offers a different model of the mind whereby cognitive processes can be partially constituted by structures in the environment. I sketch an alternative account of what the evolution of human psychology and culture by combining a developmental systems approach to evolution and development with the extended mind hypothesis. This will result in a very different understanding of the relationship between biology, psychology and culture

Leaving a solitary life behind: Evolutionary processes leading to sociality in animals

Die Entwicklung stabiler Tiergruppen ist ein wichtiger Übergang in der Evolution, der aufgrund des selektiven Drucks, der mit sozialen Interaktionen verbunden ist, Veränderungen in der Populationsstruktur und in den aufkommenden Eigenschaften mit sich bringt. Die Sozialität basiert auf Kooperation, ein evolutionäres Puzzle in der darwinistischen Theorie, das auf der Konkurrenz um begrenzte Ressourcen beruht. Im ersten Kapitel stellen wir die Bedeutung der Verwandtschaftsselektion (i.e. Nepotismus) in Frage, um das Auftreten von Tiergruppen zu erklären, das das aktuelle Paradigma darstellt. Diese Theorie legt nahe, dass genetische Ähnlichkeit notwendig ist, um die Konkurrenz zwischen Individuen zu reduzieren, die eine Kooperation ermöglichen. Wir schlagen einen alternativen Rahmen vor, der die zahlreichen und unterschiedlichen Arten berücksichtigt, in denen die elterliche Fürsorge die Entwicklung des Gruppenlebens katalysiert haben könnte. Wir betonen die Bedeutung koevolutiver Prozesse zwischen Parasiten und Raubtieren mit elterlichen Investitionsstrategien, lange bevor ein Übergang zur Sozialität stattfinden kann. Aufbauend auf empirischen und theoretischen Erkenntnissen aus einem breiten Spektrum von Taxa, einschließlich Wirbeltieren und wirbellosen Tieren, schlagen wir vor, dass direkte Fitnessvorteile, die sich aus dem selektiven Druck ergeben, der mit der Evolution der elterlichen Fürsorge verbunden ist, die Kraft hinter dem Auftreten von Tiergruppen sind. In diesem Rahmen ist die Verwandtenselektion eher ein Verstärker oder sogar ein Nebenprodukt aus evolutionären Prozessen, die mit der elterlichen Fürsorge in Verbindung stehen, und nicht die Hauptvoraussetzung für die Entwicklung der Zusammenarbeit. Im zweiten Kapitel konzentrieren wir uns auf die Untersuchung fakultativ sozialer Spezies, um die Prozesse zu verstehen, die eine einsame Spezies zu einem Gruppenleben führen. In diesem Sinne beschreiben wir das Sozialsystem eines fakultativ sozialen Primaten mit gemeinschaftlicher Zucht, Microcebus murinus, anhand von Daten über mehr als 200 Individuen aus einer Wildpopulation. Durch die Untersuchung der gemeinsamen Schlafplatznutzung bei dieser einsamen Futtersuche wollen wir die soziale Flexibilität sowohl auf der Ebene der Art als auch auf der Ebene des Individuums charakterisieren. Wir finden Belege für die soziale Flexibilität bei philopatrischen Weibchen und zerstreuenden Männchen. Darüber hinaus zeigen wir, im Gegensatz zu früheren Ergebnissen, eine höhere Fähigkeit zur Sozialität und sozialen Flexibilität bei den Männchen. Unsere Ergebnisse deuten also darauf hin, dass die weibliche Gemeinschaftszucht möglicherweise nicht die einzige treibende Kraft für die Sozialität bei dieser Art ist, was den in Kapitel 1 dargelegten Rahmen kritisiert; und dass nicht verwandte Männchen genauso anfällig für die Bildung sozialer Gruppen sind wie verwandte Weibchen, was darauf hindeutet, dass die Verwandtschaftsauswahl auch nicht in der Lage ist, die Entwicklung der sozialen Systeme des Mausmakis zu erklären. Während wir in den ersten beiden Kapiteln die Übergänge zur Sozialität aus einer anpassungsorientierten Perspektive diskutiert haben, untersuchen wir in Kapitel 3.1 die Möglichkeit, dass die Sozialität bei Microcebus murinus ein passives Ergebnis der heterogenen Verteilung von Nahrungsressourcen und Schlafplätzen sein könnte. Wir finden keine Belege für einen Effekt der Verfügbarkeit von Nahrungsressourcen oder der Einschränkung der Nistplätze auf individuelle Sozialstrategien. Daher könnten die intrinsischen Vorteile, die mit dem gemeinsamen Schlafen und der gemeinsamen Nutzung eines Heimbereichs mit anderen verbunden sind, bei dieser Art im Spiel sein. In Kapitel 3.2 entwickeln wir die in Kapitel 3.1 angewandte Methode zur Beurteilung der Zuverlässigkeit der für jedes Individuum gesammelten Informationen, um mit Hilfe der Michaelis-Menten-Modellierung Heimatorte zu bauen. Wir glauben, dass dies ein potenziell nützliches Instrument für Studien in der freien Natur sein könnte, wo sowohl die Knappheit der Daten als auch die individuellen Unterschiede in der Menge der gesammelten Daten bewegungsökologische Analysen erschweren können. Abschließend betonen wir, dass die soziale Evolution ein vielfältiger Prozess ist, der mehrere Ebenen der Lebenskomplexität in sich birgt und miteinander verflochten ist und sich den Versuchen einer einheitlichen Erklärung ihrer Ursprünge widersetzt.The evolution of stable animal groups is a major transition in evolution entailing changes in population structure and emerging properties due to the selective pressures associated with social interactions. Sociality is based on cooperation, an evolutionary puzzle in Darwinian theory that is grounded on competition for limited resources. In the first chapter, we challenge the importance of kin selection (i.e. nepotism) to explain the appearance of animal groups, which is the current paradigm. This theory suggests that genetic similarity is needed to reduce competition between individuals allowing cooperation to be selected. We propose an alternative framework that takes into account the numerous and diverse ways in which parental care may have catalyzed the evolution of group living. We emphasize the importance of coevolutionary processes between parasites and predators with parental investment strategies long before transitions to sociality may occur. Building on empirical and theoretical evidence from a wide range of taxa, including vertebrates and invertebrates, we suggest that direct fitness benefits arising from selective pressures associated with parental care evolution are the force behind the appearance of animal groups. Under this framework, kin-selection is rather an enhancer or even a by-product derived from evolutionary processes related to parental care and not the main prerequisite for cooperation to evolve. In the second chapter, we focus on studying facultatively social species to understand the processes that lead a solitary species to become group-living. In this sense, we describe the social system of a facultatively social primate with communal breeding, Microcebus murinus, using data on more than 200 individuals from a wild population. By studying sleeping site sharing in this solitary foraging species, we aim to characterize the social flexibility both at the species as well as at the individual levels. We find evidence for social flexibility in philopatric females and dispersing males. Moreover, contrary to previous findings, we show a higher capacity for sociality and social flexibility in males. Thus, our results suggest that female communal breeding may not be the only force driving sociality in this species, criticizing the framework exposed in chapter 1; and that unrelated males may be as prone as related females to form social groups, which suggests that kin-selection is also unable to explain the evolution of mouse lemurs’ social systems. While in the first two chapters, we discussed transitions to sociality from an adaptationist perspective, in Chapter 3.1, we examine the possibility that sociality in Microcebus murinus may be a passive result of heterogeneous distribution of food resources and sleeping sites. We find no evidence for an effect of food resource availability or nesting limitation on individual social strategies. Thus, intrinsic benefits associated with sleeping together and sharing a home range with others may be at play in this species. In chapter 3.2, we develop the method used in chapter 3.1 to assess the reliability of information gathered per individual to construct home ranges using Michaelis-Menten modeling. We believe this might be a potentially useful tool for studies in the wild where scarcity of data as well as between-individual variation in the amount of data collected may hamper movement ecology analyses. We end by emphasizing that social evolution is a manifold process that embeds and intertwines several layers of life complexity, resisting attempts for unitary explanations of its origins