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REDOX METABOLISM AND LIFE

in the Archaean period could have been used for aerobic respiration in the last common universal ancestor of Bacteria and Archaea. In fact, even today, cytochrome oxidases in some organisms operate in niches containing very low oxygen pressure. Interestingly, nitric oxide reductase, which catalyzes a similar reaction (Eq. (1.7)), is speculated to have been a progenitor of cytochrome c oxidases, which is homologous. 2NO + 2H+ + 2e− → N2 O + H2 O (1.7)

The “Great Oxidation Event” thus led to an explosion of genomic and metabolic complexity culminating in the rise of metazoans in the Cambrian period (∼600 million years ago). Based on the long “oxygenation time” on Earth—the time it took for atmospheric oxygen levels to reach present values (i.e., <1 billion years ago) needed to sustain large metazoan organisms—astrobiologists have argued that evolution of complex life forms on other habitable planets would be limited by the lifetime of the stars they orbit.

1.2

GLOBAL REDOX CYCLES

The evolution of life meant changes for the planetary environment and, today, nonequilibrium redox cycling of six principal elements—carbon, hydrogen, nitrogen, oxygen, phosphorus, and sulfur—shape and are, in turn, shaped by life on Earth. These biogeochemical cycles are interdependent and the major cycles are briefly discussed here (Fig. 1.2). Atmospheric carbon exists primarily in the form of carbon dioxide in the atmosphere (∼0.04% but rising). Atmospheric carbon dioxide is captured by photosynthetic plants in a process that releases oxygen while the carbon is reduced to biomass that is utilized by animals and microbes (Fig. 1.2A) or becomes buried in organic sediments. In the oxidative half cycle, the energy captured in the biomass is extracted via respiration that consumes oxygen and returns carbon dioxide to the atmosphere. The vast majority of global carbon reserves is trapped in the lithosphere in sedimentary rock deposits and does not cycle while a smaller proportion is found in terrestrial and aquatic fossil fuel reserves that are being rapidly depleted. Deforestation and fossil fuel burning have had a major impact on atmospheric carbon levels in the postindustrial history of humankind, increasing levels of greenhouse gases that lead to global warming. In aquatic environments, fermentative microbes utilize anaerobic organic detritus that is converted to a variety of small molecules that bubble upward. Some of these products can be captured by acetogens to produce acetate or methanogens to produce methane. The latter can be converted to carbon dioxide by methanotrophs or can escape into the atmosphere, where it is a potent greenhouse gas, or it can be trapped in giant gas hydrates in the ocean that may in fact constitute the largest fuel reserves on the planet. Nitrogen is the most abundant element in the atmosphere (78%); however, soluble and biologically available nitrogenous compounds are scarce commodities in natural environments. Nitrogen’s entry into the biogeochemical cycle results primarily from the activity of nitrogen-fixing bacteria that generate ammonia, which is utilized for the

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cultivation of leguminous crops. and fossil fuel burning are contributing in major ways to large scale changes in the global nitrogen cycle. H2 Decomposers.GLOBAL REDOX CYCLES 5 A Sun Photosynthesis Atomospheric O2 CO2 + H2O Respiration O2. synthesis of organic nitrogen compounds (viz. Acetogens Organic Sediments Combustion aerobic Fossil Fuels anaerobic B Nitrification NH2OH NO2− NO3− O2 Nitrogen Fixation N2 NH3 Denitrification NO2− NO N2O NO2− Nitrate Assimilation Organic Nitrogen Figure 1. CO2.2B). amino acids) (Fig. Simplified global biogeochemical cycles for carbon/hydrogen/oxygen (A) and nitrogen (B). O2 CO2 + H2O Photosynthesis Methanotrophs O CH4 2 Methanogens CH4 H2/CO2 or CH3CO2− CO. Chemical synthesis of fertilizers. Plants and some microbes capable of nitrate assimilation can use nitrite and nitrate to generate ammonia.. Anthropogenic alterations to the nitrogen biogeochemical cycle have resulted in an almost twofold increase in the transfer of inert nitrogen to biologically available forms. Ammonia can also be used as an energy source by soil bacteria and be oxidized sequentially to nitrite and nitrate in a process known as nitrification. Biomass Biomass. Search . 1.2. CH3CO2−.

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