<p>Modern microbialites and microbial mats are the focus of ongoing research as they provide an opportunity to understand microbial-mineral interactions during carbonate precipitation and the generation of biosignatures that can inform our interpretation of the geological record. This study determined the natural abundance isotopic compositions ([13]C, [14]C) of the primary carbon pools and microbial communities associated with modern freshwater microbialites located in Pavilion Lake and in carbonate rich microbial mats on the nearby Cariboo Plateau in British Columbia, Canada. </p> <p> Natural abundance [14]C analysis of carbon pools associated with the Pavilion Lake microbialites demonstrated that structures were actively growing and that groundwater carbon inputs to the lake and microbialites were minimal. Rather, ambient dissolved inorganic carbon (DIC) was the primary carbon source for both microbial communities and recent carbonate. </p> <p> Isotopic enrichment of calcium carbonate within microbial communities associated with the microbialites was identified as a biosignature of microbial photosynthetic influence driving precipitation. Elevated oxygen concentrations and pH within the microenvironment of small, sporadic nodular microbial surface communities was concurrent with in situ precipitation of carbonate with δ[13]C values higher than predicted abiotic values and δ[13]C of bulk organic matter and phospholipid fatty acids (PLFA) that were consistent with a photosynthetically dominated community. Elevated carbonate δ[13]C values were also noted in the thin surface microbial mat recovered from shallow (11m) microbialites. These samples showed increased biomass during summer sampling periods as compared to deeper samples, consistent with expected high rates of photosynthetic activity due to higher light levels and temperature at these depths. These results contrast other recent studies of modern microbialite systems that identified biosignatures of heterotrophic influences on precipitation of carbonates. PLFA profiles demonstrated that the surface microbial mat community consisting of both photosynthetic and heterotrophic microbes was stable over seasonal and spatial changes in light and temperature. However, changes in microbial biomass with depth and season indicated that microbial activity and growth plays an important role in the development of isotopic biosignatures. </p> <p> Biosignatures of high levels of photosynthetic activity were also observed in carbonate, rich microbial mats that exhibited undersaturated p CO2 concentrations during the summer and DIC δ[13]C values enriched above values predicted for isotopic equilibrium with atmospheric CO2. Seasonal and annual shifts in the balance of heterotrophy and autotrophy in the lakes and microenvironment of the mat accounted for observed variations in DIC and associated carbonate δ[13]C values. In contrast to other organic rich microbial mats, bulk organic δ[13]C values were not enriched and the systems did not show evidence of CO2 limitation. Rather, these results indicated that low bulk organic δ[13]C values and large isotopic discriminations can exist under conditions of high DIC concentrations and carbonate content that provide a non limiting carbon source to replenish photosynthetic drawdown. </p> / Thesis / Doctor of Philosophy (PhD)

Ubiquitaires et très anciens, les communautés des tapis microbiens font preuve de capacités métaboliques et adaptatives très importantes. Situés en zone côtières, ces écosystèmes peuvent être soumis à des contaminations pétrolières. Dans ce contexte, cette étude vise d’une part à décrire la structure et le fonctionnement de tapis microbiens et d’autre part à comprendre l’impact d’une contamination pétrolière sur ces écosystèmes. Cette étude porte sur deux tapis microbiens de l’étang de Berre aux paramètres physico-Chimiques proches mais présentant des contaminations pétrolières contrastées. Le fonctionnement du système étant tributaire d’autres facteurs tels que la lumière et les saisons, les variations saisonnières et nycthémérales ont été prises en compte dans cette étude. Un accent particulier a également été porté sur le cycle du soufre de par son importance en milieu marin. Les résultats de cette étude mettent en évidence des structures de communautés différentes entre les deux tapis au niveau global, la séparation spatiale prévalant sur la séparation saisonnière. La fraction active de la communauté du site contaminé présente une évolution linéaire tandis que celle du site témoins suit pour sa part les variations saisonnières. Au niveau du site contaminé une augmentation de l’expression des gènes impliqués dans la dégradation des hydrocarbures couplée à une biodégradation des hydrocarbures suggère que le tapis contaminé est adapté à la contamination pétrolière. Malgré les différences de structures et d’activités de dégradation, des profils métaboliques très semblables sont cependant observables entre les deux tapis, avec des fonctions similaires laissant supposer une redondance fonctionnelle. Des variations saisonnières et nycthémérales ont également été observées avec notamment des Desulfobulbaceae plus abondantes au printemps et plus actives en journée. Des études culturales ont été réalisées en parallèle. Elles permettront d’appréhender de manière complémentaire la dynamique des communautés des sulfato-Réducteurs au sein du tapis et de mieux comprendre les variations mises en évidence dans cette étude. / Ubiquist and very ancient, the microbial mats communities demonstrate very important metabolic and adaptive capacities. Located in the coastal area, these ecosystems may be subject to oil contamination. In this context, the aim of this study is on one hand to describe the structure and functioning of microbial mats and on the other to understand the impact of oil contamination on these ecosystems. This study focused on two microbial mats from the Berre lagoon with close physical chemical parameters but with contrasted hydrocarbon contamination levels. The functioning of the system is dependent on other factors such as light and seasons, diurnal and seasonal variations were taken into account in this study. Special emphasis was placed on the sulfur cycle due to its importance in the marine environments. The results of this study highlighted different communities’ structures at the global level between both mats, the spatial variation prevailed on seasonal variation. The active part of the community from the contaminated site shows a linear trend while that one of the uncontaminated site follows the seasonal variations. The contaminated site shows genes involved in hydrocarbon degradation more expressed coupled to a hydrocarbon biodegradation suggesting that the contaminated mat is adapted to the petroleum contamination. Despite these differences in the structure and the degradation capacities, very similar metabolic profiles are observed between the two mats with similar functions, suggesting functional redundancy. Seasonal and diurnal variation was also observed, the Desulfobulbaceae were particularly more abundant in spring and more active during the day. A complementary cultural approach will allow to better understanding the dynamics of sulfate-Reducers communities in the mat and comprehending these variations.

The current research project investigated the anoxygenic phototrophic and metal(loid) transforming bacteria of hypersaline and deep ocean hydrothermal environments. The East German Creek brine springs, an unusual flowing hypersaline system, was enumerated using classical techniques. Subterranean sulfide supported purple sulfur and nonsulfur bacteria, but at the highly oxygenated surface, aerobic anoxygenic phototrophs (AAP) were numerically dominant (up to 16-36% of cultivable bacteria). Strains (EG8, EG13, EG17, EG19) with unusual phylogenetic affiliation and novel photosynthetic and metal(loid) reducing traits were described taxonomically. Chromocurvus halotolerans gen. nov., sp. nov. was proposed as a second example of a gammaproteobacterial AAP. It exhibited bent rod-shaped cells, unusual among AAP. Facultatively anaerobic Charonomicrobium ambiphototrophicum gen. nov., sp. nov. was capable of both aerobic and anaerobic anoxygenic photosynthesis, and incapable of photoautotrophy, distinguishing it from both AAP and purple nonsulfur bacteria. Roseovarius vanadiphilum sp. nov. surprisingly produced 4.5 times more biomass and 2 times more bacteriochlorophyll (BChl) at extremely high NaVO3 concentration (7.5 g/l) than in metal-free medium. A second novel metabolic mode, anaerobic respiration on the toxic metalloid tellurate, was described for a relative of non-phototrophic Shewanella frigidimarina (ER-Te-48), from deep ocean hydrothermal vent Paralvinella worms at Explorer Ridge in the Pacific Ocean. Other strains respired on SeO32- (ER-Se-17L), VO3- (ER-V-6), and VO43- (AV-V-25). These organisms provided the first examples of anaerobic respiration on Te, Se and V at hydrothermal vents. High level resistance of AAP to metal(loid)s prompted investigation of the influence of TeO32- on photosynthetic pigment production in species including Erythromicrobium ramosum (from a terrestrial hydrothermal system) and Erythrobacter litoralis (from a hypersaline supralittoral system). Tellurite enhanced photosynthetic pigment production up to 3.4 times, consistent with an antioxidant carotenoid-based defense mechanism. However, in E. litoralis BChl precursors such as Mg protoporphyrin or its monomethyl ester also accumulated, indicating biosynthetic pathway interruption. In hydrothermal and hypersaline ecosystems, largely devoid of eukaryotic phototrophs but often enriched in metal(loid)s, AAP and metal(loid) reducers are key modulators of nutrient and toxin availability. The presented results on their ecology, physiology and biochemistry have important implications for theoretical understanding of extreme environments and hold potential for biotechnological applications.

The current research project investigated the anoxygenic phototrophic and metal(loid) transforming bacteria of hypersaline and deep ocean hydrothermal environments. The East German Creek brine springs, an unusual flowing hypersaline system, was enumerated using classical techniques. Subterranean sulfide supported purple sulfur and nonsulfur bacteria, but at the highly oxygenated surface, aerobic anoxygenic phototrophs (AAP) were numerically dominant (up to 16-36% of cultivable bacteria). Strains (EG8, EG13, EG17, EG19) with unusual phylogenetic affiliation and novel photosynthetic and metal(loid) reducing traits were described taxonomically. Chromocurvus halotolerans gen. nov., sp. nov. was proposed as a second example of a gammaproteobacterial AAP. It exhibited bent rod-shaped cells, unusual among AAP. Facultatively anaerobic Charonomicrobium ambiphototrophicum gen. nov., sp. nov. was capable of both aerobic and anaerobic anoxygenic photosynthesis, and incapable of photoautotrophy, distinguishing it from both AAP and purple nonsulfur bacteria. Roseovarius vanadiphilum sp. nov. surprisingly produced 4.5 times more biomass and 2 times more bacteriochlorophyll (BChl) at extremely high NaVO3 concentration (7.5 g/l) than in metal-free medium. A second novel metabolic mode, anaerobic respiration on the toxic metalloid tellurate, was described for a relative of non-phototrophic Shewanella frigidimarina (ER-Te-48), from deep ocean hydrothermal vent Paralvinella worms at Explorer Ridge in the Pacific Ocean. Other strains respired on SeO32- (ER-Se-17L), VO3- (ER-V-6), and VO43- (AV-V-25). These organisms provided the first examples of anaerobic respiration on Te, Se and V at hydrothermal vents. High level resistance of AAP to metal(loid)s prompted investigation of the influence of TeO32- on photosynthetic pigment production in species including Erythromicrobium ramosum (from a terrestrial hydrothermal system) and Erythrobacter litoralis (from a hypersaline supralittoral system). Tellurite enhanced photosynthetic pigment production up to 3.4 times, consistent with an antioxidant carotenoid-based defense mechanism. However, in E. litoralis BChl precursors such as Mg protoporphyrin or its monomethyl ester also accumulated, indicating biosynthetic pathway interruption. In hydrothermal and hypersaline ecosystems, largely devoid of eukaryotic phototrophs but often enriched in metal(loid)s, AAP and metal(loid) reducers are key modulators of nutrient and toxin availability. The presented results on their ecology, physiology and biochemistry have important implications for theoretical understanding of extreme environments and hold potential for biotechnological applications.

Les microbialithes sont des dépôts organosédimentaires benthiques résultant de la minéralisation et de la lithification de tapis microbiens, et dont les plus anciennes formes, se développant il y a 3.4 Ga, constituèrent les premiers écosystèmes terrestres. Ils forment alors une archive sédimentaire unique incluant des périodes clés de l’histoire géologique. Ce travail de thèse se propose d’analyser et de comparer les processus et produits de minéralisation dans les tapis microbiens actuels de deux environnements contrastés : un exemple de lac intracontinental hypersalin, le Grand Lac salé (GSL) aux USA ; une lagune hypersaline à alimentation marine, à Cayo Coco (Cuba) (CCLN). Le devenir des minéraux au cours de la diagenèse précoce, ainsi que leur potentiel d’enregistrement de biosignatures seront particulièrement analysés. Cette thèse se focalisera spécialement sur l’influence de trois facteurs majeurs contrôlant la minéralogie et la fabrique des microbialithes : (i) le rôle de la chimie du milieu (ii) le rôle des métabolismes microbiens (le moteur de l’alcalinité) ; (iii) le rôle de la production et de la dégradation des matrices organiques extracellulaires (EOM). Les deux cas d’études démontrent un rôle prépondérant de la production d’EOM par les cyanobactéries et leur dégradation par les bactéries hétérotrophes dans la minéralisation : (1) Dans les deux systèmes, la première phase minérale a précipiter sur les EOM alvéolaires est une phase riche en magnésium et en silicium. Ce type de minéraux nécessite des pH>8.6-8.7 pour cristalliser. (2) Une autre observation commune est que les carbonates cristallisent souvent dans des zones de forte activité des bactéries sulfato-réductrices (SRB). Notre hypothèse est que les SRB dégradent les EOM, libérant des cations (Mg2+ et Ca2+) disponibles pour la cristallisation des carbonates. Dans les tapis du CCLN et contrairement au GSL, nos résultats démontrent une forte activité de photosynthèse anoxygénique par les bactéries pourpres sulfureuses (PSB). La limite entre la zone oxique et la zone anoxique est caractérisée par un pH maximum et coïncide avec la formation d’une lamine de carbonates. Deux différences majeures sont observées entre les paragenèses du GSL et du CCLN : (1) le locus initial de la précipitation des carbonates. Dans le GSL, l’aragonite précipite dans les cyanobactéries, perminéralise leur paroi et enfin la matrice organique. Pour Cuba, une calcite magnésienne péloïdale précipite sur les EOM puis rempli les bactéries ; (2) la minéralogie et l’évolution des carbonates lors de la diagenèse précoce. Les microbialithes du GSL montrent une aragonite partiellement dissoute et une dolomite venant se développer à sa périphérie. Au CCLN, de l’aragonite se développe en surcroissance des peloïdes de HMC précédemment formés. Les différences minéralogiques des carbonates entre les deux systèmes pourraient s’expliquer par un changement du rapport Mg/Ca. Les résultats pourront être utilisés afin de mieux interpréter les conditions paléoenvironnementales et les processus microbiens en jeu dans des microbialithes de registres fossiles analogues. / Microbialites are benthic organosedimentary deposits resulting of the mineralization and lithification of the microbial mats, and the most ancient forms, developing there are 3.4 Ga, are the first earthly ecosystem. They form a unique sedimentary archive including key periods of the geological history. This study proposes to analyze and compare the processes and the products of mineralization in the modern microbial mats of two different environments: an example of intracontinental modern lake, the Great Salt Lake (USA; GSL); a lagoonal marine sea fed in Cayo Coco (Cuba; CCLN). The mineral product during of the primary diagenesis, as that them potential of biosignatures recording will be particularly detailed. This work will focus on the influence of three major factors controlling the mineralogy and the fabric of the microbialites: (i) environment chemistry role, (ii) microbial metabolisms role, (iii) role of the production and degradation of the extracellular organic matrix (EOM). Both environments studied show a high role of the EOM production by cyanobacteria and them degradation by the heterotrophic bacteria in the mineralization: (1) In both systems, the first phase to precipitate on the alveolar EOM is a rich magnesium and silica phase. This type of mineral needs pH around 8.6/8.7 to precipitate. (2) An other common observation is that carbonate precipitate generally in the high sulfate-reducing activity zones. Our hypothesis is that the sulfato-reducing bacteria (SRB) degrade the EOM, releasing cations (Mg2+ and Ca2+) available for carbonate crystallization. The limit between the oxic and anoxic zones is characterized by maximum pH coinciding with the precipitation of carbonate lamina. Two mains differences have been observed between the paragenesis both systems: (1) initial locus of the carbonate precipitation. In the GSL, the aragonite precipitates in the bacteria and then permineralizes the wall of bacteria and then the EOM network. In Cuba, the peloidal magnesian calcite precipitates on the EOM then fill the bacteria; (2) the mineralogy and the evolution of the carbonate during the preliminary diagenesis. The microbialithes of GSL show the aragonite partly dissolved and a dolomite developing next to the aragonite. In the CCLN, aragonite developing around the magnesian calcite peloids. The mineralogical carbonate differences between both systems could explain by a change of the Mg/Ca. The results could be used to better understand and interpret the paleoenvironmental conditions and the microbial processes stake in ancient microbialite analogs.

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