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Nitrososphaeria

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Nitrososphaeria is a distinct class of Archaea belonging to the phylum Thermoproteota. These microscopic organisms play vital roles in the Earth's biogeochemical cycles. They are primary drivers in both the nitrogen and carbon cycles. Most species in this class function as chemolithoautotrophic ammonia-oxidizers. This means they derive energy from chemical reactions involving ammonia. They also create their own food from inorganic carbon. Because of these roles, they are essential to the health of aquatic and terrestrial environments.

The way these organisms process energy is highly specialized. Most members use an ammonia oxidation pathway to gain energy. This process allows them to thrive in oligotrophic conditions, where nutrients are very scarce. They can function at much lower ammonia concentrations than ammonia-oxidizing bacteria. Furthermore, their pathway requires less oxygen to work. This allows them to dominate in low-oxygen environments like sediments or hot springs. Some species can even use urea as a substrate for nitrification. This allows them to compete with phytoplankton for resources.

Nitrososphaeria also possess a unique method for fixing carbon. They use the hydroxypropionate/hydroxybutyrate cycle to turn bicarbonate into organic matter. This specific cycle appears to have evolved independently from other Thermoproteota. Notably, this carbon-fixation pathway is more efficient than any other known aerobic autotrophic pathway. This high efficiency is why they succeed in nutrient-poor areas. Some species, such as Nitrosopumilus maritimus, exhibit mixotrophy. Mixotrophy is the ability to use both organic and inorganic carbon sources. At least two isolated strains are obligate mixotrophs, meaning they must have organic carbon to grow.

The history of discovering Nitrososphaeria is a journey of changing scientific classifications. In 1996, biologists at the University of California found Cenarchaeum symbiosum inside a sponge near Santa Barbara. Initially, researchers thought it was related to the Crenarchaeota group. However, genetic analysis of ribosomal RNA and DNA polymerase suggested otherwise. In 2005, biologists at the University of Washington identified Nitrosopumilus maritimus in Seattle. By 2008, Nitrososphaera gargensis was discovered in a Siberian hot spring. Because these organisms were so genetically different from known phyla, scientists created the phylum Thaumarchaeota in 2008.

Taxonomy continued to evolve as new data emerged. In 2014, Michaela Stieglmeier and her colleagues discovered Nitrososphaera viennensis in Austrian garden soil. They established the class Nitrososphaeria and several other taxonomic ranks. In 2021, the phylum Nitrososphaerota was formalized by Aharon Oren and George M. Garrity. However, a team of Australian scientists led by Christian Rinke and Philip Hugenholtz published a new classification. They merged Nitrososphaerota into the phylum Thermoproteota. This change demoted Nitrososphaeria from a phylum to a class level.

These organisms serve as incredible tools for understanding Earth's history. They produce unique membrane-spanning lipids called glycerol dialkyl glycerol tetraethers, or GDGTs. One specific lipid, crenarchaeol, is found only in Nitrososphaeria. These lipids act as a potential biomarker for the class. Because the structure of these lipids varies with temperature, scientists use them as a proxy. This method is known as the TEX86 paleotemperature proxy. By studying GDGTs in marine sediments, researchers can reconstruct ancient ocean temperatures. Additionally, their ability to fix CO2 allows their lipids to record past carbon cycle changes.

Beyond climate records, Nitrososphaeria support massive biological networks. They are likely the dominant producers of vitamin B12 in many environments. Vitamin B12 is critical for DNA synthesis and the citric acid cycle. Many eukaryotic phytoplankton are auxotrophic, meaning they cannot make B12 themselves. They must acquire it from the environment. Therefore, Nitrososphaeria may influence algal blooms and global atmospheric CO2 levels. In the ocean, many species live in the subphotic zone between 100 and 350 meters deep. They even exist in extreme places like tropical mangrove swamps. Some species there, like Candidatus Giganthauma, form very large filaments.

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