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There are ~12 supergroups of complex-celled organisms (eukaryotes), but relationships among them (including the root) remain elusive. For Paper I, I developed a dataset of 37 eukaryotic proteins of bacterial origin (euBac), representing the conservative protein core of the proto-mitochondrion. This gives a relatively short distance between ingroup (eukaryotes) and outgroup (mitochondrial progenitor), which is important for accurate rooting. The resulting phylogeny reconstructs three eukaryote megagroups and places one, Discoba (Excavata), as sister group to the other two (neozoa). This rejects the reigning “Unikont-Bikont” root and highlights the evolutionary importance of Excavata. For Paper II, I developed a 150-gene dataset to test relationships in supergroup SAR (Stramenopila, Alveolata, Rhizaria). Analyses of all 150-genes give different trees with different methods, but also reveal artifactual signal due to extremely long rhizarian branches and illegitimate sequences due to horizontal gene transfer (HGT) or contamination. Removing these artifacts leads to strong consistent support for Rhizaria+Alveolata. This breaks up the core of the chromalveolate hypothesis (Stramenopila+Alveolata), adding support to theories of multiple secondary endosymbiosis of chloroplasts. For Paper III, I studied the evolution of cox15, which encodes the essential mitochondrial protein Heme A synthase (HAS). HAS is nuclear encoded (nc-cox15) in all aerobic eukaryotes except Andalucia godoyi (Jakobida, Excavata), which encodes it in mitochondrial DNA (mtDNA) (mt-cox15). Thus the jakobid gene was postulated to represent the ancestral gene, which gave rise to nc-cox15 by endosymbiotic gene transfer. However, our phylogenetic and structure analyses demonstrate an independent origin of mt-cox15, providing the first strong evidence of bacteria to mtDNA HGT. Rickettsiales or SAR11 often appear as sister group to modern mitochondria. However these bacteria and mitochondria also have independently evolved AT-rich genomes. For Paper IV, I assembled a dataset of 55 mitochondrial proteins of clear α-proteobacterial origin (including 30 euBacs). Phylogenies from these data support mitochondria+Rickettsiales but disagree on the placement of SAR11. Reducing amino-acid compositional heterogeneity (resulting from AT-bias) stabilizes SAR11 but moves mitochondria to the base of α-proteobacteria. Signal heterogeneity supporting other alternative hypotheses is also detected using real and simulated data. This suggests a complex scenario for the origin of mitochondria.

The Proterozoic period saw some of the most fundamental revolutions in the biological and geological world. During this period, life diversified and set the stage for the radiation of multicellular life, altering the face of the planet in the process. The fossil record of this time is not yet fully understood, and a revisitation of a historically reported fossil deposit in the 1 Ga Torridon rocks of northwest Scotland shows that they host the fossils of the earliest non-marine eukaryotes, as well as a full and diverse fossil assemblage preserved in sedimentary phosphates and shales. Fine scale sedimentology of the fluvio-lacustrine rocks of the Torridon Group reveals them to be laid down in a laterally variable basin with distinctly different palaeoenvironments. The resident biota is seen to be similarly variable between lithofacies. New criteria for classifying taphonomic effects are presented, and used to characterise assemblages from different palaeoenvironments, with broader applications beyond this study. The Torridon rocks are also host to macrostructures on the surfaces and soles of beds, and these are interpreted as of likely biological origin, with their variability mapped between different lithofacies. High-resolution studies of both the preserved biota and the mineralogy of the preserving medium reveal in detail not only the fine scale structure of the fossil organisms, but also the reasons for their exceptional preservation. Phosphate is analysed in detail to explain its enigmatic occurrence in Proterozoic lakes. To place the Torridon deposits in context, both older and younger rocks were examined in comparison, from the 2 Ga Gunflint Formation of Ontario, Canada, and the Precambrian-Cambrian successions of eastern Newfoundland respectively. New finds of phosphate in these rocks help to reveal biochemical interactions and evolution on the early Earth, with implications for further understanding life on our own planet and elsewhere.

No / The genome of Caenorhabditis elegans encodes multiple homologues of the two major families of mammalian equilibrative and concentrative nucleoside transporters. As part of a programme aimed at understanding the biological rationale underlying the multiplicity of eukaryote nucleoside transporters, we have now demonstrated that the nematode genes ZK809.4 (ent-1) and K09A9.3 (ent-2) encode equilibrative transporters, which we designate CeENT1 and CeENT2 respectively. These transporters resemble their human counterparts hENT1 and hENT2 in exhibiting similar broad permeant specificities for nucleosides, while differing in their permeant selectivities for nucleobases. They are insensitive to the classic inhibitors of mammalian nucleoside transport, nitrobenzylthioinosine, dilazep and draflazine, but are inhibited by the vasoactive drug dipyridamole. Use of green fluorescent protein reporter constructs indicated that the transporters are present in a limited number of locations in the adult, including intestine and pharynx. Their potential roles in these tissues were explored by using RNA interference to disrupt gene expression. Although disruption of ent-1 or ent-2 expression alone had no effect, simultaneous disruption of both genes yielded pronounced developmental defects involving the intestine and vulva.

Recent studies suggest that horizontal gene transfer (HGT) plays an important role in niche adaptation in some eukaryotes and may be a major evolutionary force in unicellular lineages. One subcategory of HGT is endosymbiotic gene transfer (EGT), which is characterized by a large influx of genes from endosymbiont to host nuclear genome and is a critical step in the establishment of permanent organelles, such as plastids. The dinoflagellates are a diverse group of mostly marine eukaryotes that have a propensity for both HGT and plastid endosymbiosis. Many dinoflagellates are predators and can acquire both genes and plastids from prey, blurring the distinction between HGT and EGT. Here, I measure genome mosaicism in dinoflagellates to investigate how HGT has impacted gene innovation and plastid endosymbiosis in this group. Because analysis of HGT depends on accurate phylogenetic trees, I first assessed the sensitivity of automated phylogenomic methods to variation in taxon sampling due to homolog selection parameters. Using methods based on this analysis, I showed that a large amount of HGT has occurred in dinoflagellates, particularly from bacterial donors. Further, I demonstrated that the dinoflagellate Alexandrium tamarense has the largest number of genes gained relative to related eukaryotes using ancestral gene content reconstruction. Additionally, dinoflagellates have lost several ubiquitous eukaryotic metabolic genes, but missing genes have been functionally replaced by xenologs from many evolutionarysources. Other transferred genes are involved in diverse functions. These results suggest that dinoflagellate genomes are heavily impacted by HGT. Also, I investigated the timing and consequences of HGT in plastid endosymbiosis. Using the dinflagellate Dinophysis acuminata, a mixotrophic species that sequesters and maintains prey plastids, I identified plastid-targeted proteins that function in photosystem stabilization and metabolite transport. Dinophysis acuminate may be able to extend the useful life of the stolen plastid by protecting the photosystem and replacing damaged transporters. Phylogenetic analyses showed that genes are derived from multiple sources indicating a complex evolutionary history involving HGT. Dinophysis acuminate can acquire both genes and plastids from prey, which suggests that HGT could play an important role in plastid acquisition during the earliest stages of this transition.

Les introns sont des portions de gènes transcrites dans l’ARN messager, mais retirées pendant l’épissage avant la synthèse des produits du gène. Chez les eucaryotes, on rencontre les introns splicéosomaux, qui sont retirés de l’ARN messager par des splicéosomes. Les introns permettent plusieurs processus importants, tels que l'épissage alternatif, la dégradation des ARNs messagers non-sens, et l'encodage d'ARNs fonctionnels. Leurs rôles nous interrogent sur l'influence de la sélection naturelle sur leur évolution. Nous nous intéressons aux mutations qui peuvent modifier les produits d'un gène en changeant les sites d'épissage des introns. Ces mutations peuvent influencer le fonctionnement d'un organisme, et constituent donc un sujet d'étude intéressant, mais il n'existe actuellement pas de logiciels permettant de les étudier convenablement. Le but de notre projet était donc de concevoir une méthode pour détecter et analyser les changements des sites d'épissage des introns splicéosomaux. Nous avons finalement développé une méthode qui repère les évènements évolutifs qui affectent les introns splicéosomaux dans un jeu d'espèces données. La méthode a été exécutée sur un ensemble d'espèces d'oomycètes. Plusieurs évènements détectés ont changé les sites d’épissage et les protéines, mais de nombreux évènements trouvés ont modifié les introns sans affecter les produits des gènes. Il manque à notre méthode une étape finale d'analyse approfondie des données récoltées. Cependant, la méthode actuelle est facilement reproductible et automatise l'analyse des génomes pour la détection des évènements. Les fichiers produits peuvent ensuite être analysés dans chaque étude pour répondre à des questions spécifiques. / Introns are portions of genes transcribed into messenger RNA, but removed during RNA splicing. In eukaryotes, they are called spliceosomal introns as they are removed by spliceosomes. Introns allow many important processes such as alternative splicing, nonsense-mediated decay and functional-RNA coding. These roles leads to the question of the influence of natural selection on evolution of introns. We focus on mutations that are able to change gene products by modifing introns splice sites. These mutations seems to be an interesting topic as they can affect proteins, but there is currently no software to study them properly. The aim of our project was to design a method to detect and analyze changes in splice sites of spliceosomal introns. We finally developed a method that locates the evolutionary events on splice sites of spliceosomal introns in a given species set. The method was performed on a set of oomycetes. Several detected events change splice sites and proteins, but there is also many events that seems to modify introns without affecting gene products. Our method lacks a final step for thorough analysis of the collected events. However, the current method is easily reusable and automates genome analysis for the detection of events. The resulting files can then be analyzed in each study to answer specific questions.