Univ., Diss.--Düsseldorf.

Darwin's (1859) theory of evolution by natural selection accounts for the adaptations of organisms, but, as Fisher (1930) famously said, 'natural selection is not evolution.' Evolutionary theory has two major components: i) natural selection, which involves the underlying dynamics of populations; and ii) adaptive evolutionary change, which involves the optimisation of phenotypes for fitness maximisation. Many of the traditional theoretical frameworks in evolutionary theory have focussed on studying optimisation processes that generate biological adaptations. In recent years, however, a number of evolutionary theorists have turned to using frameworks such as the 'replicator dynamics' or 'eco-evolutionary dynamics', to explore the dynamics of natural selection. There has, however, been little attempt to explore how these dynamical systems frameworks relate to more traditional frameworks in evolutionary theory or how they incorporate the principles that embody the process of evolution by natural selection, namely, phenotypic variation, differential reproductive success, and heritability. In this thesis, I use these principles to provide the formal foundations of a general framework - a mathematical synthesis - in which the future state of an evolutionary system can be predicted from its present state; what I will call a 'dynamical systems theory of natural selection.' Given the state of an existing biological system, and a set of assumptions about how individuals within the system interact, the job of the dynamical systems theory of natural selection is no less than to predict the future in its entirety.

Natural selection favours those individuals that respond best to novel features of their selective environment. For many, a critical challenge is responding to evolutionary change in mutualistic species. These responses create complex feedbacks, so only coevolutionary approaches are able to fully answer key questions about the maintenance or disruption of mutualistic behaviour, and explain the range of mechanisms that allow individuals to benefit from these associations. I first consider the hypothesis that economic models studying multiple classes of traders, where each trader seeks to optimise its own payoffs will yield insights into mutualistic systems. I show that individuals can be favoured to discriminate amongst potential partners based on the price for which they provide resources. Then, I show that market mechanisms can maintain cooperation and drive specialisation in mutualistic systems. I extend this market model to allow individuals to restrict a mutualistic partner's access to resources, and show that this strategy can stabilise cooperation and increase the fitness of both partners. I also explicitly incorporate relatedness in my market model. I show that high relatedness sometimes increases cooperativeness in members of a mutualistic species, but sometimes decreases cooperativeness as it narrow the scope for partner choice to maintain cooperation. Having studied market mechanisms, I consider indiscriminate costly help to members of another species. I discover that this trait can be favoured by natural selection and can be classified as either altruism between or altruism within species. Finally, I consider a framework for analysing coevolved phenotypic responses to a partner's cooperativeness, a challenging process to model. I demonstrate that this framework can yield firm predictions about behaviour whenever partners hold private information about their costs and benefits.

Les bactéries constituent le socle de presque tous les écosystèmes et l’étude de leurs dynamiques face aux perturbations biotiques et abiotiques est essentielle à la compréhension de leur maintien, de leur évolution et de leur diversification. Cette thèse vise à une meilleure appréhension de l’impact des bactériophages et des antibiotiques sur l’écologie évolutive des populations bactériennes et, plus particulièrement, sur l’évolution de leurs comportements sociaux. Dans une première partie, nous avons étudié comment les antibiotiques (Chapitres 1 et 2) et les phages (Chapitre 3) affectent les interactions fondées sur la production de biens publics ainsi que l’évolution de la résistance dans les populations de Pseudomonas aeruginosa, en combinant modélisation mathématique et évolution expérimentale. Nous avons montré que les phages et les antibiotiques favorisent les tricheurs face aux coopérateurs dans les environnements homogènes. Alors que l’avantage des tricheurs permet la croissance de la population et augmente la fréquence de résistance à court terme (Chapitre 1), les populations dominées par les tricheurs finissent par décliner en présence de phages, vraisemblablement suite aux pressions combinées des phages et des tricheurs (Chapitre 3). Dans une seconde partie, nous avons exploré in vitro les interactions complexes entre les phages et les antibiotiques dans le contexte des thérapies combinées. Conformément à la prédiction de la théorie de l’évolution selon laquelle plusieurs moyens de contrôle combinés sont plus efficaces que chacun séparément, nous avons montré que l’usage simultané de phages et d’antibiotiques réduit davantage la survie et la résistance des populations. Si ce résultat principal peut être modulé par différents facteurs tels que la dose d’antibiotiques (Chapitres 4 et 5), le moment d’inoculation (Chapitre 4), et le mode d’action des antibiotiques (Chapitre 5), il persiste sur le long terme (Chapitre 5). Nos résultats soulignent la complexité des interactions entre les effets négatifs des phages et des antibiotiques et l’écologie évolutive des populations bactériennes et apportent de nouveaux éléments à la fois à la compréhension de l’évolution de la socialité et à l’usage thérapeutique potentiel des phages et des antibiotiques. / Bacteria are the basis of virtually all ecosystems and examining their dynamics in the face of biotic and abiotic perturbations is essential to understanding their persistence, evolution and diversification. This thesis is directed towards a better understanding of the impact of phage and antibiotic pressure on the evolutionary ecology of bacterial populations and, in particular, on the evolution of bacterial social behaviours. First, using a combination of mathematical modelling and experimental evolution, we studied how antagonisms in the form of antibiotics (Chapters 1 and 2) and phages (Chapter 3) affect the dynamics of public goods production and strategies, and the evolution of resistance in populations of the bacterium Pseudomonas aeruginosa. We found that both phages and antibiotics favour cheats over cooperators in well-mixed environments. While the advantage to cheats leads to population growth and even increased resistance frequency in the short-term (Chapter 1), the cheat-dominated populations eventually declined in the presence of phage predators, arguably due to the combination of antagonist pressure and cheating load (Chapter 3). Second, based on the evolutionary prediction that multiple control agents will be more efficient at controlling bacterial populations and reducing the evolution of resistance, we investigated in vitro the complex interactions between phages and antibiotics in the context of combined therapies. We showed that the combination of phages and antibiotics decreased population survival and resistance evolution significantly more than either alone. While this main result may be mitigated by several factors such as antibiotic dose (Chapters 4 and 5), the timing of inoculation (Chapter 4), and antibiotic mode of action (Chapter 5), it is also obtained in longer-term assays (Chapter 5). Our results highlight the complexity of the interplay between the negative effects exerted by antibiotics and phages and the evolutionary ecology of bacterial populations, and bring new insights both to the understanding of social evolution and for the potential therapeutic use of phages and antibiotics.

The core aim of this interdisciplinary research is to provide a critical analysis of the influence of Darwinism and Social Darwinism on a sample of modern Chinese writings. To achieve these aims, the researcher uses a range of both Chinese and English sources to explore their close affinities with Darwinism and Social Darwinism. Following this course, the research examines how Darwinian thought was introduced to the Chinese reading public in the late nineteenth century through a translation of Thomas Henry Huxley’s Evolution and Ethics by Yen Fu, and the subsequent impact of this work and Darwinian thought in general on seven literary and political figures: K'ang Yu-wei, Liang Qichao, Lu Xun, Hu Shih, Chen Duxiu, Sun Yat-sen and Mao Zedong. From an historical perspective, the Opium Wars and imperial invasions of China in the nineteenth century severely weakened the country’s political, economic, diplomatic, military, educational and cultural power. For these reasons and others, from 1840 to 1949, China experienced a tumultuous period of social and political transformation, which has eventually led to her revival in the twenty-first century. It will be seen that each of the literary figures examined here used evolutionary thought to justify revolution at various points on China’s long march to modernity. Progressive Darwinian ideas sharply contrasted with the old Confucian values upheld within Chinese communities. Nevertheless, the faults and weaknesses of Qing China awakened many pioneering revolutionaries who sought to reverse the status quo by initiating a series of radical reforms and revolutionary movements. Many within the Chinese intellectual elite looked to the tide of change and progress coming from the West, which they hoped might replace the recent historical stagnation and Confucian dogma embedded in Chinese culture and society. In this vein, many of these pioneering revolutionaries set about driving the historical transformation of China by selecting, translating and interpreting Darwinian ideas in their own writings. From Yen Fu in the nineteenth century to Mao Zedong in the twentieth century, evolutionary thought went hand in hand with China’s modernization.

Cumulative culture is rare, if not altogether absent in non-human species. At the foundation of cumulative learning is the ability to flexibly modify, relinquish or build upon prior behaviours to make them more productive or efficient. Within the primate literature, a failure to optimise solutions in this way is often proposed to derive from low-fidelity copying of witnessed behaviours, sub-optimal social learning heuristics, or a lack of relevant socio-cognitive adaptations. However, humans can also be markedly inflexible in their behaviours, perseverating with, or becoming fixated on outdated or inappropriate responses. Humans show differential patterns of flexibility as a function of cognitive load, exhibiting difficulties with inhibiting suboptimal behaviours when there are high demands on working memory. Here I present a series of studies on captive chimpanzees which show that not only is inhibitory control compromised in chimpanzees, but indicate ape behavioural conservatism may be underlain by similar constraints as in humans; chimpanzees show relatively little conservatism when behavioural optimisation involves the inhibition of a well-established but simple solution, or the addition of a simple modification to a well-established but complex solution. In contrast, when behavioural optimisation involves the inhibition of a well-established but complex solution, and especially when the alternative solution is also complex, chimpanzees show evidence of behavioural conservatism. I propose that conservatism is linked to behavioural complexity, potentially mediated by cognitive resource availability, and may be an important factor in the evolution of cumulative culture.

本論文從系統分化的社會演化觀點嘗試提出對現代民主之內涵與民主化開展之可能性的另種論述。本文中所提及的「民主」與民主化不一定跳脫西方世界對相關概念的想像與定義，但在嘗試將「民主」視為政治系統用以描述自身之方式的這個主張而言，無論是西方式民主或者是其他種類的民主均能夠含括於其中。此外，民主化的開展僅被視為政治系統朝向「民主」的一種系統分化的社會演化過程。西方國家對民主的界定則是在此分化的演化發展中，透過宗教與政治、宗教與法律，以及政治與法律等系統彼此之間、以及從社會中分化出來而取得之形式「之一」。這是一個對系統而言不斷增加複雜性的過程，也正是在此過程中，對於民主或民主化之內容的界定變得越來越不容易。在本論文對西方與台灣民主化過程的探究中，嘗試指出的是一個類似的功能分化的社會演化與系統內部分化的相互辯證過程。這個辯證過程使現代「民主」概念得以作為一個「未預期之後果」而出現。 宗教對信仰自由的維護與對人權的爭取毋寧是來自於自身的信仰元素。換句話說，與之有關的論述是以一種自我指涉的運作方式而獲得實現。此種不斷回到自身之信仰以尋求論述之正當性的作法在操作上賦予了這些論述正當性，這便是系統的自我指涉運作。就此而言，系統毋寧是封閉的，其在自身之中不斷地自我生產有關信仰自由與人權的論述。然而，也正是因為其運作上的封閉，使得宗教（長老教會）所生產出來的論述與接續之行動均能夠在內部獲得支持，從而對自身而言具有正當性。這個正當性也成為其能夠進行觀察與自我描述的基礎。從系統論的觀點而言，長老教會在政治與社會關懷的論述與行動上，便是以封閉為基礎的開放性自我指涉運作。若非以此自我指涉的封閉且開放之運作為其系統形成的特性，那麼論述與行動的正當性便易於受到質疑，甚至無法獲得系統內部的支持。據此，現代社會以功能為主要分化形式的系統形成，至少在時間面向上均必須取得自我指涉或自主性運作上的正當性。這個正當性決定於系統在多大程度上能夠持續地維持與其環境的界線。長老教會對人權與信仰自由的主張與論述的建構，或許是與政治系統對相同論題的溝通存在著無法跨越的差異。這個差異同時也標示的是宗教與政治系統之間的那個界線，而這個差異的維持在系統分化的社會演化過程中，使得「民主（化）」作為政治系統乃至於社會描述自身的語意是可能的。 / This thesis draws on Niklas Luhmann’s theory of social systems, and examines the relationship between religion and democratization in Taiwan. In order to advance the analysis and to make some helpful suggestions, two points are firstly highlighted. One is the concept of democracy itself, and the other is the descriptions of it by the political system and by other social systems within society. The word ‘democracy’ has long history since ancient Greek city-state, however, its modern use and the meaning of it can possibly date back to the Middle ages, especially to the sixteenth century. Modern concept of democracy relates to the tradition of monotheism, in this case referring to the Reformation and later to the formation of Western political philosophy. The second point argues that ‘democracy’ is simply a self-description of society, and its realization lies in the transformation of the form of differentiation from hierarchical to functional. Luhmann would like to link the semantics to social evolution, by which he can investigate and explore the dialectical process between semantics and structures. The analysis of the relationship between democracy and democratization refers to this dialectical process, concerning systems differentiation and social evolution. Issues relating to democratization, the consolidation of democracies, and their discontented consequences, etc. need to be reconsidered in the sense that the concept of democracy and its modern descriptions utilize a ‘paradox’ which must be hidden while forming and maintaining identities, whether they be societies, nations, or individuals. This constitutes both positive and negative sides of the development of democracy, and also leads to crises gradually confronted by those ‘matured’ democratic Western countries and also by some third-wave democratizing regions. Among them exists a similar problem which will be discussed in this thesis: the concept of democracy and its modern developments are increasingly eroding its own foundation when the differentiation of subsystems and the evolution of society both go too far. This by no means indicates that there will not be any democracy in the near future. Instead, as an observer observing observations, this thesis inquires our present situations while at the same time attempting at offering some possible and also suitable questions from systems theory perspective.

Der Zusammenhang von technischer Entwicklung und gesellschaftlicher Modernisierung wird mit Mitteln der systemtheoretischen Soziologie untersucht. Zu diesem Zweck werden die Konzepte "Technik" und "soziale Evolution" reformuliert. "Technik" ist ein spezieller Typ der Grenzbildung von Systemen, die Ereignisse ihrer Umwelt als kausal verknüpft beobachten; Technik wird demnach definiert als fixierte Erwartung, die sich auf kausal verknüpfte Umweltereignisse bezieht. "Soziale Evolution" ist der Typ des Strukturwandels von sozialen Systemen, der sich aus der Selektion und Restabilisierung von Strukturvariationen ergibt. Aus soziologischer Sicht ist jede technische Entwicklung als ein spezifischer Aspekt der sozialen Evolution zu verstehen. Dies wird mit der Konfrontation des Begriff des "Fortschritts" mit dem der "Modernisierung" gezeigt. Eine Gesellschaft versucht durch das Verwenden und das Beschränken von Technik über ihre eigene Evolution zu verfügen. Es erweist sich, daß diese Versuche zwar komplexer, aber nicht unbedingt erfolgreicher werden. Die einer jeden Evolution eigene Entwicklungsoffenheit kann nicht außer Kraft gesetzt werden, auch dann nicht, wenn eine Gesellschaft Evolutionstheorien entwickelt und reflexiv auf sich selbst anwendet. Anders als vormoderne Gesellschaften zeichnet sich die moderne Gesellschaft durch ihre hochgradig frei kontextualisierbare und damit riskante Technik aus. Kontextualisierungen und Entscheidungsnotwendigkeiten im Umgang mit moderner Technik bedingen Reflexivität. Technische Entwicklungen lassen Determinationen und Optionen in gleichem Maße wachsen. Komplex ist die soziale Seite von Technik, nicht die Kompliziertheit ihrer technischen Komponenten. Reflexive Technologiepolitik und Technikfolgenabschätzung sind gezwungen, die soziale Dynamik technischer Entwicklungen zu berücksichtigen. / There are several means to analyse the intrinsic connection of technological development and social evolution in sociology today. The thesis examines technology and its development as part of social evolution from a system-theoretical point of view. Using this approach, terms like "technology" and "social evolution" are reformulated. "Technology" is a particular type of border of social systems, using the mode of causality in observing its environment; thus Technology is defined as the fixation of causal expectations. "Social evolution" is a social systems' structural change by the selection and reinforcement of variations, creating a singular history. In terms of sociology, any technological development is part of a wider developing social context. This is visualized by confronting the terms "progress" and "modernization"; both being concepts of sociology as well as of debates on technology and social development. By using technology, by giving way to innovation, and by limiting the use of technology, a given society tries to monitor and influence its own evolution. The thesis shows that these attempts become more complex, but not necessarily more effective. The fundamental properties of any evolution cannot be annulated --- the social evolution always remains unpredictable, even when evolutionary theories allow reflexive observations. In contrast to its predecessors the modern society constitutes a type of technology which is highly independent of any given social context. But this freedom is risky. Contextualizations and decision-makings imply a higher degree of reflexivity. In the case of technological development, both determinations and options increase at the same time. What is complex in technological development is its social, not its mere technical side. Any reflexive debate on technology has to reflect technology as a social issue.

O objetivo deste trabalho é analisar os elementos que influenciaram a transformação do modo de conhecer do ser social ao longo da história. Partimos da indicação de Marx e Engels, acerca da relação entre a consciência e o mundo material, em que afirmam que é o ser social que determina a consciência. Sendo o ser social determinado pelas atividades práticas que realiza, e sendo a práxis produtiva a atividade fundamental de produção e reprodução da vida material, delimitamos nosso objeto a dois elementos: a divisão do trabalho e a troca mercantil. Nossa hipótese se expressa por meio de uma metáfora: antes o conhecimento era um espelho inteiro, porém embaçado. Depois, ele se torna um espelho quebrado, porém com cada pedaço mais nítido. Argumentaremos que a divisão do trabalho “quebra o espelho” transformando o conhecimento de totalidade (espelho inteiro) em unilateralidade (espelho quebrado em vários pedaços). E a troca mercantil “limpa o espelho” transformando o conhecimento que antes era superficial (embaçado) em um conhecimento mais profundo (límpido), por meio da relação entre abstração real e abstração do pensamento. Concluímos que a divisão do trabalho e a troca mercantil transformaram o modo de conhecer de algo mais empírico para algo mais abstrato. / The objective of this study is to analyze the factors that influenced the transformation of the social being mode of knowing throughout history. We start from the statement of Marx and Engels on the relationship between consciousness and the material world, affirming that it is the social being that determines consciousness. If the social being is determined by the practical activities, and if the productive practice is the fundamental activity of production and reproduction of material life, we delimit our object to two elements: the division of labor and commodity exchange. Our hypothesis is expressed through a metaphor: before the knowledge was a full mirror, but blurry. Then it becomes a broken mirror, but with each piece clearer. We will argue that the division of labor "break the mirror" transforming the totality of knowledge (full mirror) in one-sidedness (mirror broken into several pieces). And the commodity exchange "clean the mirror" transforming knowledge that was superficial (blurry) on a deeper knowledge (clear), through the relationship between real abstraction and abstraction of thought. We conclude that the division of labor and commodity exchange turned the mode of knowing something more empirical to something more abstract.

O objetivo deste trabalho é analisar os elementos que influenciaram a transformação do modo de conhecer do ser social ao longo da história. Partimos da indicação de Marx e Engels, acerca da relação entre a consciência e o mundo material, em que afirmam que é o ser social que determina a consciência. Sendo o ser social determinado pelas atividades práticas que realiza, e sendo a práxis produtiva a atividade fundamental de produção e reprodução da vida material, delimitamos nosso objeto a dois elementos: a divisão do trabalho e a troca mercantil. Nossa hipótese se expressa por meio de uma metáfora: antes o conhecimento era um espelho inteiro, porém embaçado. Depois, ele se torna um espelho quebrado, porém com cada pedaço mais nítido. Argumentaremos que a divisão do trabalho “quebra o espelho” transformando o conhecimento de totalidade (espelho inteiro) em unilateralidade (espelho quebrado em vários pedaços). E a troca mercantil “limpa o espelho” transformando o conhecimento que antes era superficial (embaçado) em um conhecimento mais profundo (límpido), por meio da relação entre abstração real e abstração do pensamento. Concluímos que a divisão do trabalho e a troca mercantil transformaram o modo de conhecer de algo mais empírico para algo mais abstrato. / The objective of this study is to analyze the factors that influenced the transformation of the social being mode of knowing throughout history. We start from the statement of Marx and Engels on the relationship between consciousness and the material world, affirming that it is the social being that determines consciousness. If the social being is determined by the practical activities, and if the productive practice is the fundamental activity of production and reproduction of material life, we delimit our object to two elements: the division of labor and commodity exchange. Our hypothesis is expressed through a metaphor: before the knowledge was a full mirror, but blurry. Then it becomes a broken mirror, but with each piece clearer. We will argue that the division of labor "break the mirror" transforming the totality of knowledge (full mirror) in one-sidedness (mirror broken into several pieces). And the commodity exchange "clean the mirror" transforming knowledge that was superficial (blurry) on a deeper knowledge (clear), through the relationship between real abstraction and abstraction of thought. We conclude that the division of labor and commodity exchange turned the mode of knowing something more empirical to something more abstract.