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Nitrogen fixation and its impact in the early evolution of life of Earth Supervisors: Dr Patricia Sanchez-Baracaldo (School of Biological Sciences and School of Geographical Sciences, University of Bristol) – Main supervisor Dr Davide Pisani (School of Biological Sciences and School of Earth Sciences, University of Bristol) Prof Timothy Lenton (Department of Geographical Sciences, University of Exeter) Host institution: University of Bristol Project description: Nitrogen is an essential nutrient whose availability limits the productivity of the biosphere1, and life on Earth would not have had a chance of diversify without the evolution of organisms capable of reducing N 2 (which is extremely abundant in the atmosphere but extremely stable and therefore unusable by the largest majority of organisms) to ammonia, which on the other hand, can be easily assimilated into living systems. The process of nitrogen reduction to ammonia is generally known as nitrogen fixation. This key process is carried out by a small number of diverse prokaryotes including several Eubacteria and the methanogenic Archaeabacteria using enzymes known as nitrogenases. Several different nitrogenases exist and from a strictly functional point of view, they differ because they use alternative metallic central atoms (Iron, molybdenum, or vanadium) in the heterometallic complex defining their active site. While there is strong evidence to suggest that extant nitrogenases can be traced back to a single ancestor2, it is still unclear which nitrogenease (Iron, molybdenum, or vanadium) dependent evolved first, and in what order the other nitrogenases originated. Further to that, it is still unclear when and in what organisms these genes first emerged and how they spread across the prokaryotes as there is strong evidence that they have been laterally transferred across distantly related lineages. Objectives: This project aims to understand how the nitrogenase enzyme complex evolved, elucidating the origin of nitrogen fixation and the establishment of a key global bigeochemical cycle (the nitrogen cycle). In particular we shall (1) Investigate relationships among nitrogenase enzymes and delineate evolutionary patterns (both vertical and horizontal) across prokaryotes. Further to that (2) we shall attempt establishing what were the first organisms able to carry out Nitrogen fixation and (3) when in Earth History this process was established by looking at what is known of the geochemical record (e.g. Nitrogen isotopes). Gruber, N., and Galloway, J. N. (2008). An Earth-system perspective of the global nitrogen cycle. Nature 451, 293–296. Falkowski, P. G. (1997). Evolution of the nitrogen cycle and its influence on the biological sequestration of CO 2 in the ocean. Nature 387, 272–275