The phylogenetic position of Xenoturbella bocki has been a matter of controversy since its description in 1949. We sequenced a second complete mitochondrial genome of this species and performed phylogenetic analyses based on the amino acid sequences of all 13 mitochondrial protein-coding genes and on its gene order. Our results confirm the deuterostome relationship of Xenoturbella. However, in contrast to a recently published study (Bourlat et al. in Nature 444:85–88, 2006), our data analysis suggests a more basal branching of Xenoturbella within the deuterostomes, rather than a sister-group relationship to the Ambulacraria (Hemichordata and Echinodermata).

In this paper, we present a method and a corresponding tool called Tarzan for cophylogeny analysis of phylogenetic trees where the nodes are labelled with divergence timing information. The tool can be used for example to infer the common history of hosts and their parasites, of insect-plant relations or symbiotic relationships. Our method does the reconciliation analysis using an event-based concept where each event is assigned a cost and cost minimal solutions are sought. The events that are used by Tarzan are cospeciations, sortings, duplications, and (host) switches. Different from existing tools, Tarzan can handle more complex timing information of the phylogenetic trees for the analysis. This is important because several recent studies of cophylogenetic relationship have shown that timing information can be very important for the correct interpretation of results from cophylogenetic analysis. We present two examples (one host-parasite system and one insect-plant system) that show how divergence timing information can be integrated into reconciliation analysis and how this influences the results.

Part I: A remarkable feature of life on Earth is that despite the apparent observed diversity, the underlying chemistry that powers it is highly conserved. From the level of the nucleobases, through the amino acids and proteins they encode, to the metabolic pathways of chemical reactions catalyzed by these proteins, biology often utilizes identical solutions in vastly disparate organisms. This universality is intriguing as it raises the question of whether these recurring features exist because they represent some truly optimal solution to a given problem in biology, or whether they simply exist by chance, having arisen very early in life's history. In this project we consider the universality of metabolism { the set of chemical reactions providing the energy and building blocks for cells to grow and divide. We develop an algorithm to construct the complete network of all possible biochemically feasible compounds and reactions, including many that could have been utilized by life but never were. Using this network we investigate the most highly conserved piece of metabolism in all of biology, the trunk pathway of glycolysis. We design a method which allows a comparison between the large number of alternatives to this pathway and which takes into account both thermodynamic and biophysical constraints, finding evidence that the existing version of this pathway produces optimal metabolic fluxes under physiologically relevant intracellular conditions. We then extend our method to include an evolutionary simulation so as to more fully explore the biochemical space. Part II: Studies of population dynamics have a long history and have been used to understand the properties of complex networks of ecological interactions, extinction events, biological diversity and the transmission of infectious disease. One aspect of these models that is known to be of great importance, but one which nonetheless is often neglected, is spatial structure. Various classes of models have been proposed with each allowing different insights into the role space plays. Here we use a lattice-based approach. Motivated by gene transfer and parasite dynamics, we extend the well-studied contact process of statistical physics to include multiple levels. Doing so generates a simple model which captures in a general way the most important features of such biological systems: spatial structure and the inclusion of both vertical as well as horizontal transmission. We show that spatial structure can produce a qualitatively new effect: a coupling between the dynamics of the infection and of the underlying host population, even when the infection does not affect the fitness of the host. Extending the model to an arbitrary number of levels, we find a transition between regimes where both a finite and infinite number of parasite levels are sustainable, and conjecture that this transition is related to the roughening transition of related surface growth models.

W.D. Hamilton has been celebrated as the twentieth-century Darwin. His extension of evolutionary theory to explain social behaviours has been extensively documented. Current accounts, however, have often overlooked the extent to which his early research goals were tied to his desire to see that a better world was created through a scientific understanding of society. In fact, when his interests in humans, and especially his eugenic concerns, have been acknowledged, they have been distanced from his scientific achievements and treated separately. Using new sources to reexamine the development of Hamilton’s most famous idea, the theory of inclusive fitness, we may better understand how his perception of cultural upheaval shaped his reading of social behaviours as evolved characters following universal laws. Understanding this, we may see that however successful Hamilton was, he never realized his original dream, which was to devise a theory that would inform the human world, replacing religious and ideological beliefs. As he sought to solidify his career in the 1970s, he moved away from publicly disclosing his more controversial ideas. This meant that by the time the science of social behaviour inspired heated debates, he was almost always absolved from political critiques. Many assumed that his theory was derived from observations of insects, and his eugenic ideas were forgotten, ignored, or not understood. He was therefore well positioned to become the objective figurehead of a new discipline, sociobiology. This does not mean that his desire to understand society as the result of genetic laws subsided, and by placing inclusive fitness against its social and political background, we might reimagine its trajectory and its impact in new ways. We might also begin to see Hamilton not as an isolated scholar unengaged with society but as an individualist whose extra-scientific beliefs paralleled his scientific theories in meaningful ways.

Palaeontological data are essential for determining patterns of biological diversity through geological time, enabling the investigation of important macroevolutionary events such as mass extinctions and explosive radiations. Most studies utilise proxies of taxonomic diversity. A more complex undertaking is to assess patterns of morphological variety (disparity) through time, revealing the manner in which groups evolved through their ‘design space’. Many published studies indicate clades tend to reach their maximum disparity early in their evolutionary history. Whether this is a real biological pattern has yet to be tested. Chapter 1 tackles the evolution of disparity in metazoans across the Phanerozoic. The results of a meta-analysis of disparity in 98 extinct clades indicate early high disparity is the most prevalent pattern across the Phanerozoic but finds no clear trends through the Phanerozoic. Mass extinction ended clades were the exception, tending to result in late high disparity. Chapters 2-4 focus on the clade Bivalvia for disparity and diversity analysis. Bivalves are ecologically and taxonomically diverse and have an excellent fossil record but have not been scrutinised using the latest diversity techniques, and have been untouched by disparity analysis. Chapter 2 uses the most up to date stratigraphic ranges and techniques to revise the bivalve Phanerozoic diversity curve. The results show bivalve Phanerozoic diversity is robust to the sampling and fossil record biases examined. Chapter 3 uses data provided as part of collaboration between Martin Hughes, Dr Joseph Carter (University of North Carolina) and Dr Matthew Wills (University of Bath) to address the disparity of bivalves across the Palaeozoic. The results find disparity rises across time but not decreased by mass extinctions. Chapter 4 conducts the first large scale analysis of disparity across latitude. The results find that bivalve disparity across latitude is unchanging and stable compared to the steep gradient of bivalve diversity.

Les piRNAs sont une population de petit ARNs très diverse, que l'on retrouve dqns la lignée germinales des animaux pour réprimer les éléments génétiques mobiles : agissant de pair avec les protéines Piwi, ils guident le clivage des transposons actif. Chez la Drosophile, 3 protéines Piwi sont présentes, dont deux d'entre elles, AUB et AGO3, sont cytoplasmique et la dernière, PIWI, est nucléaire cependant son mécanisme d'action reste inconnu. La source principale de piRNAs sont des régions du génome bien particulière, appelé cluster de piRNAs. Cependant, il n'est pas encore connu a ce jour qu'est ce qui différentie ces région du reste du génome. Durant mon doctorant mon travail s'est focalisé sur ces deux questions centrales :Quel est le rôle de PIWI dans le noyau? Nous avons montré que PIWI était responsable de répression transcriptionnelle des transposons par l'intermédiaire de la déposition de marques chromatiniennes répressive, H3K9me3, grâce à la spécificité des piRNAs.Comment sont définit les régions générant des piRNA et comment sont régules leur expression ?Nous avons trouvé que les piRNAs qui sont transmis par la mère aux progénitures sont responsables de l'identification des régions génomiques donnant naissances à de nouveau piRNAs, grâce à la déposition de H3K9me3 dans le noyau et par l'initiation du cycle ping-pong dans le cytoplasme.Nous avons aussi mis en évidence les régions promoteurs des clusters de piRNAs, et trouve qu'elles sont nécessaires pour la production de piRNAs. / PiRNAs are a diverse population of small RNA found in the animal germline to silence mobile genetic elements: loaded into Piwi proteins, they guide homology-dependent cleavage of active transposon mRNAs. In Drosophila, three Piwi proteins are expressed, from which two, AUB and AGO3, are known to destroy transposon transcripts in the cytoplasm. The third one, Piwi itself, is nuclear and the molecular mechanism of its function remains unknown. The main sources of piRNAs are discrete genomic loci called piRNA clusters, however it is not known what differentiate them from non-piRNA producing loci. During my PhD, I focused my work on two central questions:1) What is the role of Piwi in the nucleus? We showed that Piwi is responsible for transcriptional silencing by mediating installment of repressive marks, especially H3K9me3, over active transposons copies in a piRNA dependent manner.2) How are piRNA clusters defined, and what regulates their expression? Analyzing what features differentiate a piRNA producing loci from any non-producing loci in the genome, we were able to single out some specific characteristics: . We showed that maternally inherited piRNAs are responsible to define germline clusters at the next generation through two mechanisms: in the nucleus, by deposition of H3K9me3 onto complementary genomic sequence, and, in the cytoplasm, by initiating the ping-pong cycle using cluster transcripts as substrates, leading to their processing into mature piRNAs.. We found that cluster promoters are essential to mediate full cluster transcription, which is allowed thanks to a very specific chromatin signature necessary to ensure piRNA production.

Cette étude analyse la quasi-totalité du cycle de vie de deux espèces de puffins sympatriques, le puffin tropical (Puffinus bailloni, PT) et le puffin du Pacific (P. pacificus, PP) nichant sur Aride Island (Seychelles). L’objectif est de comprendre les interactions existantes entre ces espèces, ainsi que l’influence de l’environnement, au sein de la colonie et dans le milieu marin. Le nombre de couples ont était estime à 15,000 pour le PP et 30,000 pour le PT (où la population est probablement en déclin). Ils sélectionnent davantage des zones escarpées avec des végétations basses. Les PP montrent également une préférence pour les zones rocheuses avec un sol profond. Les PT nichent pendant toute l’année mais de manière plus intensive en fin de saison de reproduction des PP et le taux d'échec de la reproduction est maximal durant la période où les deux espèces nichent en même temps. Dans le milieu marin, il-y-a une ségrégation spatiale entre les deux espèces, exploitant des zones d’alimentation bien distinctes hors saison de reproduction et se distinguant également dans leur comportement alimentaire (profondeur de plonge). Dans la colonie les deux espèces sont en concurrence sur le site de reproduction mais les PP semblent être de meilleurs compétiteurs. Nous proposons la mise en place de mesures de conservation pour les deux espèces, mais favorisant spécialement les PT. / The procellariiformes are one of the most endangered orders in the world. In addition, information on population estimates, trends, movements at sea and population dynamic is lacking for many species. We analysed almost the full life cycle of two sympatric species: the tropical (Puffinus bailloni, TS) and the wedge-tailed shearwater (P. pacificus, WS) breeding at Aride Island (Seychelles), in order to understand the interactions between them and the environment both at sea and in the colony. We assessed habitat selection, abundance, distribution and breeding success of the two species at the colony and the habitat selection and distribution at sea. The population was 15,000 and 30,000 pairs for WS and TS respectively; the TS is possibly in decline. Both species selected steep and lowly vegetated areas; the WS also selected boulder-like zones. The TS nests year-round but it has a peak of breeding at the end of the WS breeding season and its failure rate is higher when the two species co-exist. We found spatial, rather than ecological, segregation between the two species at sea in both foraging behaviour (diving depth) and inter-breeding areas. On land they are likely to compete for breeding sites with the WS often over competing the TS. We then suggest future conservation plans focused on both species but especially on the TS.

Central notions in evolutionary biology are intrinsically topological. This claim is maybe most obvious for the discontinuities associated with punctuated equilibria. Recently, a mathematical framework has been developed that derives the concepts of phenotypic characters and homology from the topological structure of the phenotype space. This structure in turn is determined by the genetic operators and their interplay with the properties of the genotype-phenotype map.

Background Coevolutionary systems like hosts and their parasites are commonly used model systems for evolutionary studies. Inferring the coevolutionary history based on given phylogenies of both groups is often done by employing a set of possible types of events that happened during coevolution. Costs are assigned to the different types of events and a reconstruction of the common history with a minimal sum of event costs is sought. Results This paper introduces a new algorithm and a corresponding tool called CoRe-PA, that can be used to infer the common history of coevolutionary systems. The proposed method utilizes an event-based concept for reconciliation analyses where the possible events are cospeciations, sortings, duplications, and (host) switches. All known event-based approaches so far assign costs to each type of cophylogenetic events in order to find a cost-minimal reconstruction. CoRe-PA uses a new parameter-adaptive approach, i.e., no costs have to be assigned to the coevolutionary events in advance. Several biological coevolutionary systems that have already been studied intensely in literature are used to show the performance of CoRe-PA. Conclusion From a biological point of view reasonable cost values for event-based reconciliations can often be estimated only very roughly. CoRe-PA is very useful when it is difficult or impossible to assign exact cost values to different types of coevolutionary events in advance.

This thesis investigates how increased global mean temperatures on Earth, induced by the increase in the luminosity of the Sun as it ages, change the types of habitable environments on the planet at local scales over the next 3 Gyr. Rising temperatures enhance silicate weathering rates, reducing atmospheric CO₂ levels to below the threshold for photosynthesis, while simultaneously pushing environments past the temperature tolerances of plant and animal species. This leads to the end of all plant life and animal life (due to reduced food, O₂ and H₂O availability, as well as higher temperatures) within the next 1 Gyr. The reduction in the extent of the remaining microbial biosphere due to increasing temperatures and rapid ocean evaporation is then modelled, incorporating orbital parameter changes until all known types of life become extinct; a maximum of 2.8 Gyr from the present. The biosignatures associated with these changes are determined and the analysis extended to Earth-like extrasolar planets nearing the end of their habitable lifetimes. In particular, the stages in the main sequence evolutions of Sun-like stars within 10 pc are evaluated and used to extrapolate the stage that an Earth-analogue planet would be at in its habitable evolution, to determine the best candidate systems for a far-future Earth-analogue biosphere, highlighting the Beta Canum Venaticorum system as a good target. One of the most promising biosignatures for a microbial biosphere on the far-future Earth (and similar planets) may be CH₄, which could reach levels in the atmosphere that make it more readily detectable than it is for a present-day Earth-like atmosphere. Determining these biosignatures will help expand the search for life to the wider range of environments that will be found as the habitable exoplanet inventory grows and planets are found at different stages in their habitable evolution.