The nitrogen cycle on Earth today is driven by a complex network of microbially-mediated transformations. Atmospheric N2 is fixed into biologically available forms that can either be incorporated into biomass or utilized for bioenergetic redox reactions. The cycle is kept in balance by the return of fixed nitrogen to the atmospheric N2 pool by anammox and denitrification. The early evolution and history of the nitrogen cycle is not well resolved, particularly before the evolution of oxygenic photosynthesis and rise of atmospheric oxygen ca. 2.3 Gya. Ammonia oxidation is a biochemically difficult reaction requiring activation of ammonia using O2 or oxidized nitrogen species that are produced using O2. Before the rise of oxygen, when O2 was largely unavailable, nitrification could not proceed, trapping fixed nitrogen in reduced forms such as ammonia and biomass. Without production of nitrite and nitrate, anammox and denitrification could not occur, preventing return of fixed nitrogen to the N2 pool and leaving the nitrogen cycle unclosed. While it has been hypothesized that ammonia oxidation could be driven anaerobically by processes such as phototrophy or iron reduction, these metabolisms have not been recovered in extant microorganisms, and would require complex unknown biochemical mechanisms. Furthermore, phylogenetic data for the key organisms and biochemical pathways involved in denitrification and anammox suggest that these metabolisms postdate the rise of oxygen. This is particularly clear for steps utilizing enzymes in the Heme-Copper Oxidoreductase superfamily, which appear to have originally evolved for O2 reduction at non-negligible substrate concentrations. Together, this suggests that the Archean nitrogen cycle was not closed, and that nitrogen fixed to reduced forms—either through biological nitrogen fixation or abiotic processes—was not easily returned to the atmospheric N2 pool. In principle, this could have stripped the atmosphere of N2 over timescales of hundreds of Myr, which is consistent with recent paleopressure estimates that suggest < 0.5 bar by late Archean time. The modern, N2-rich atmosphere and (largely) closed biological nitrogen cycle may therefore not have evolved until Proterozoic time, after the rise of oxygen.
AGU Fall Meeting Abstracts
- Pub Date:
- December 2016
- 4845 Nutrients and nutrient cycling;
- OCEANOGRAPHY: BIOLOGICAL AND CHEMICALDE: 4912 Biogeochemical cycles;
- and modeling;