Nitrososphaerota

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Nitrososphaerota
ThaspiviridaeFig1rv2.png
Nitrosopumilus maritimus , partially with virions of Nitrosopumilus spindle-shaped virus 1 ( Thaspiviridae ) attached.
Scientific classification
Domain:
Superphylum:
Phylum:
Nitrososphaerota

Brochier-Armanet et al. 2021 [1]
Class:
Order
Synonyms
  • "Nitrososphaerota" Whitman et al. 2018
  • "Nitrososphaeraeota" Oren et al. 2015
  • "Thaumarchaeota" Brochier-Armanet et al. 2008 [2]

The Nitrososphaerota (syn. Thaumarchaeota) are a phylum of the Archaea proposed in 2008 after the genome of Cenarchaeum symbiosum was sequenced and found to differ significantly from other members of the hyperthermophilic phylum Thermoproteota (formerly Crenarchaeota). [3] [2] [4] Three described species in addition to C. symbiosum are Nitrosopumilus maritimus , Nitrososphaera viennensis , and Nitrososphaera gargensis . [2] The phylum was proposed in 2008 based on phylogenetic data, such as the sequences of these organisms' ribosomal RNA genes, and the presence of a form of type I topoisomerase that was previously thought to be unique to the eukaryotes. [2] [5] This assignment was confirmed by further analysis published in 2010 that examined the genomes of the ammonia-oxidizing archaea Nitrosopumilus maritimus and Nitrososphaera gargensis , concluding that these species form a distinct lineage that includes Cenarchaeum symbiosum. [6] The lipid crenarchaeol has been found only in Nitrososphaerota, making it a potential biomarker for the phylum. [7] [8] Most organisms of this lineage thus far identified are chemolithoautotrophic ammonia-oxidizers and may play important roles in biogeochemical cycles, such as the nitrogen cycle and the carbon cycle. Metagenomic sequencing indicates that they constitute ~1% of the sea surface metagenome across many sites. [9]

Contents

Nitrososphaerota-derived membrane-spanning tetraether lipids (glycerol dialkyl glycerol tetraethers; GDGTs) from marine sediments can be used to reconstruct past temperatures via the TEX86 paleotemperature proxy, as these lipids vary in structure according to temperature. [10] Because most Nitrososphaerota seem to be autotrophs that fix CO2, their GDGTs can act as a record for past Carbon-13 ratios in the dissolved inorganic carbon pool, and thus have the potential to be used for reconstructions of the carbon cycle in the past. [7]

Taxonomy

Phylogeny of Nitrososphaerota [11] [12] [13]
Conexivisphaeria
Conexivisphaerales
Conexivisphaeraceae

Conexivisphaera

Nitrososphaeria
Nitrososphaerales
Nitrososphaeraceae

Nitrososphaera

Nitrosopumilales
Nitrosopumilaceae

Nitrosarchaeum

Nitrosopumilus

Phylogeny of Nitrososphaerota [14] [15] [16]
Nitrososphaeria
"Geothermarchaeales"

"Geothermarchaeaceae"

Conexivisphaerales
Conexivisphaeraceae

Conexivisphaera

Nitrososphaerales
"Nitrosocaldaceae"

"Ca. Nitrosothermus"

"Ca. Nitrosocaldus"

Nitrososphaeraceae

"Ca. Nitrosocosmicus"

"Ca. Nitrosopolaris"

Nitrososphaera

Nitrosopumilaceae

"Ca. Nitrosotalea"

"Ca. Nitrosotenuis"

"Ca. Nitrosopelagicus"

" Cenarchaeum "

Nitrosarchaeum

Nitrosopumilus

The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN) [17] and National Center for Biotechnology Information (NCBI) [18]

Metabolism

Nitrososphaerota are important ammonia oxidizers in aquatic and terrestrial environments, and are the first archaea identified as being involved in nitrification. [32] They are capable of oxidizing ammonia at much lower substrate concentrations than ammonia-oxidizing bacteria, and so probably dominate in oligotrophic conditions. [8] [33] Their ammonia oxidation pathway requires less oxygen than that of ammonia-oxidizing bacteria, so they do better in environments with low oxygen concentrations like sediments and hot springs. Ammonia-oxidizing Nitrososphaerota can be identified metagenomically by the presence of archaeal ammonia monooxygenase (amoA) genes, which indicate that they are overall more dominant than ammonia oxidizing bacteria. [8] In addition to ammonia, at least one Nitrososphaerota strain has been shown to be able to use urea as a substrate for nitrification. This would allow for competition with phytoplankton that also grow on urea. [34] One study of microbes from wastewater treatment plants found that not all Nitrososphaerota that express amoA genes are active ammonia oxidizers. These Nitrososphaerota may be capable of oxidizing methane instead of ammonia, or they may be heterotrophic, indicating a potential for a diversity of metabolic lifestyles within the phylum. [35] Marine Nitrososphaerota have also been shown to produce nitrous oxide, which as a greenhouse gas has implications for climate change. Isotopic analysis indicates that most nitrous oxide flux to the atmosphere from the ocean, which provides around 30% of the natural flux, may be due to the metabolic activities of archaea. [36]

Many members of the phylum assimilate carbon by fixing HCO3 . [9] This is done using a hydroxypropionate/hydroxybutyrate cycle similar to the Thermoproteota but which appears to have evolved independently. All Nitrososphaerota that have been identified by metagenomics thus far encode this pathway. Notably, the Nitrososphaerota CO2-fixation pathway is more efficient than any known aerobic autotrophic pathway. This efficiency helps explain their ability to thrive in low-nutrient environments. [33] Some Nitrososphaerota such as Nitrosopumilus maritimus are able to incorporate organic carbon as well as inorganic, indicating a capacity for mixotrophy. [9] At least two isolated strains have been identified as obligate mixotrophs, meaning they require a source of organic carbon in order to grow. [34]

A study has revealed that Nitrososphaerota are most likely the dominant producers of the critical vitamin B12. This finding has important implications for eukaryotic phytoplankton, many of which are auxotrophic and must acquire vitamin B12 from the environment; thus the Nitrososphaerota could play a role in algal blooms and by extension global levels of atmospheric carbon dioxide. Because of the importance of vitamin B12 in biological processes such as the citric acid cycle and DNA synthesis, production of it by the Nitrososphaerota may be important for a large number of aquatic organisms. [37]

Environment

Many Nitrososphaerota, such as Nitrosopumilus maritimus, are marine and live in the open ocean. [9] Most of these planktonic Nitrososphaerota, which compose the Marine Group I.1a, are distributed in the subphotic zone, between 100m and 350m. [7] Other marine Nitrososphaerota live in shallower waters. One study has identified two novel Nitrososphaerota species living in the sulfidic environment of a tropical mangrove swamp. Of these two species, Candidatus Giganthauma insulaporcus and Candidatus Giganthauma karukerense, the latter is associated with Gammaproteobacteria with which it may have a symbiotic relationship, though the nature of this relationship is unknown. The two species are very large, forming filaments larger than ever before observed in archaea. As with many Nitrososphaerota, they are mesophilic. [38] Genetic analysis and the observation that the most basal identified Nitrososphaerota genomes are from hot environments suggests that the ancestor of Nitrososphaerota was thermophilic, and mesophily evolved later. [32]

See also

Related Research Articles

<i>Nanoarchaeum equitans</i> Species of archaeon

Nanoarchaeum equitans is a species of marine archaea that was discovered in 2002 in a hydrothermal vent off the coast of Iceland on the Kolbeinsey Ridge by Karl Stetter. It has been proposed as the first species in a new phylum, and is the only species within the genus Nanoarchaeum. Strains of this microbe were also found on the Sub-polar Mid Oceanic Ridge, and in the Obsidian Pool in Yellowstone National Park. Since it grows in temperatures approaching boiling, at about 80 °C (176 °F), it is considered to be a thermophile. It grows best in environments with a pH of 6, and a salinity concentration of 2%. Nanoarchaeum appears to be an obligate symbiont on the archaeon Ignicoccus; it must be in contact with the host organism to survive. Nanoarchaeum equitans cannot synthesize lipids but obtains them from its host. Its cells are only 400 nm in diameter, making it the smallest known living organism, and the smallest known archaeon.

<span class="mw-page-title-main">Nitrification</span> Biological oxidation of ammonia/ammonium to nitrate

Nitrification is the biological oxidation of ammonia to nitrate via the intermediary nitrite. Nitrification is an important step in the nitrogen cycle in soil. The process of complete nitrification may occur through separate organisms or entirely within one organism, as in comammox bacteria. The transformation of ammonia to nitrite is usually the rate limiting step of nitrification. Nitrification is an aerobic process performed by small groups of autotrophic bacteria and archaea.

<span class="mw-page-title-main">Korarchaeota</span> Proposed phylum within the Archaea

The Korarchaeota is a proposed phylum within the Archaea. The name is derived from the Greek noun koros or kore, meaning young man or young woman, and the Greek adjective archaios which means ancient. They are also known as Xenarchaeota. The name is equivalent to Candidatus Korarchaeota, and they go by the name Xenarchaeota or Xenarchaea as well.

Methanotrophs are prokaryotes that metabolize methane as their source of carbon and chemical energy. They are bacteria or archaea, can grow aerobically or anaerobically, and require single-carbon compounds to survive.

Methanosarcina acetivorans is a versatile methane producing microbe which is found in such diverse environments as oil wells, trash dumps, deep-sea hydrothermal vents, and oxygen-depleted sediments beneath kelp beds. Only M. acetivorans and microbes in the genus Methanosarcina use all three known metabolic pathways for methanogenesis. Methanosarcinides, including M. acetivorans, are also the only archaea capable of forming multicellular colonies, and even show cellular differentiation. The genome of M. acetivorans is one of the largest archaeal genomes ever sequenced. Furthermore, one strain of M. acetivorans, M. a. C2A, has been identified to possess an F-type ATPase along with an A-type ATPase.

<span class="mw-page-title-main">Nitrosopumilales</span> Order of archaea

The Nitrosopumilales are an order of the Archaea class Nitrososphaeria.

<i>Nitrosopumilus</i> Genus of archaea

Nitrosopumilus maritimus is an extremely common archaeon living in seawater. It is the first member of the Group 1a Nitrososphaerota to be isolated in pure culture. Gene sequences suggest that the Group 1a Nitrososphaerota are ubiquitous with the oligotrophic surface ocean and can be found in most non-coastal marine waters around the planet. It is one of the smallest living organisms at 0.2 micrometers in diameter. Cells in the species N. maritimus are shaped like peanuts and can be found both as individuals and in loose aggregates. They oxidize ammonia to nitrite and members of N. maritimus can oxidize ammonia at levels as low as 10 nanomolar, near the limit to sustain its life. Archaea in the species N. maritimus live in oxygen-depleted habitats. Oxygen needed for ammonia oxidation might be produced by novel pathway which generates oxygen and dinitrogen. N. maritimus is thus among organisms which are able to produce oxygen in dark.

<span class="mw-page-title-main">Archaea</span> Domain of single-celled organisms

Archaea is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria, but this term has fallen out of use.

<i>Cenarchaeum symbiosum</i> Species of archaeon

Cenarchaeum symbiosum is a species of Archaea in the genus Cenarchaeum, in the phylum Nitrososphaerota in the domain Archaea. C. symbiosum is psychrophilic and is found inhabiting marine sponges.

Nanohaloarchaea is a clade of diminutive archaea with small genomes and limited metabolic capabilities, belonging to the DPANN archaea. They are ubiquitous in hypersaline habitats, which they share with the extremely halophilic haloarchaea.

Nitrososphaera is a mesophilic genus of ammonia-oxidizing Crenarchaeota. The first Nitrososphaera organism was discovered in garden soils at the University of Vienna leading to the categorization of a new genus, family, order and class of Archaea. This genus is contains three distinct species: N. viennensis, Ca. N. gargensis, and Ca N. evergladensis. Nitrososphaera are chemolithoautotrophs and have important biogeochemical roles as nitrifying organisms.

The "Aigarchaeota" are a proposed archaeal phylum of which the main representative is Caldiarchaeum subterraneum. It is not yet clear if this represents a new phylum or a Nitrososphaerota order, since the genome of Caldiarchaeum subterraneum encodes several Nitrososphaerota-like features. The name "Aigarchaeota" comes from the Greek αυγή, avgí, meaning "dawn" or "aurora", for the intermediate features of hyperthermophilic and mesophilic life during the evolution of its lineage.

<span class="mw-page-title-main">Lokiarchaeota</span> Phylum of archaea

Lokiarchaeota is a proposed phylum of the Archaea. The phylum includes all members of the group previously named Deep Sea Archaeal Group, also known as Marine Benthic Group B. Lokiarchaeota is part of the superphylum Asgard containing the phyla: Lokiarchaeota, Thorarchaeota, Odinarchaeota, Heimdallarchaeota, and Helarchaeota. A phylogenetic analysis disclosed a monophyletic grouping of the Lokiarchaeota with the eukaryotes. The analysis revealed several genes with cell membrane-related functions. The presence of such genes support the hypothesis of an archaeal host for the emergence of the eukaryotes; the eocyte-like scenarios.

Nitrososphaera gargensis is a non-pathogenic, small coccus measuring 0.9 ± 0.3 μm in diameter. N. gargensis is observed in small abnormal cocci groupings and uses its archaella to move via chemotaxis. Being an Archaeon, Nitrososphaera gargensis has a cell membrane composed of crenarchaeol, its isomer, and a distinct glycerol dialkyl glycerol tetraether (GDGT), which is significant in identifying ammonia-oxidizing archaea (AOA). The organism plays a role in influencing ocean communities and food production.

<span class="mw-page-title-main">DPANN</span> A superphylum of Archaea grouping taxa that display various environmental and metabolic features

DPANN is a superphylum of Archaea first proposed in 2013. Many members show novel signs of horizontal gene transfer from other domains of life. They are known as nanoarchaea or ultra-small archaea due to their smaller size (nanometric) compared to other archaea.

"Candidatus Thorarchaeota", or simply Thorarchaeota, is a phylum within the superphylum Asgard archaea. The Asgard superphylum represents the closest prokaryotic relatives of eukaryotes. Since there is such a close relation between the two different domains, it provides further evidence to the two-domain tree of life theory which states that eukaryotes branched from the archaeal domain. Asgard archaea are single cell marine microbes that contain branch like appendages and have genes that are similar to eukarya. The asgard archaea superphylum is composed of Thorarchaeota, Lokiarchaeota, Odinarchaeota, and Heimdallarchaeota. Thorarchaeota were first identified from the sulfate-methane transition zone in tidewater sediments. Thorarcheota are widely distributed in marine and freshwater sediments.

<span class="mw-page-title-main">TACK</span> Clade of Archaea

TACK is a group of archaea, its name an acronym for Thaumarchaeota, Aigarchaeota, Crenarchaeota, and Korarchaeota, the first groups discovered. They are found in different environments ranging from acidophilic thermophiles to mesophiles and psychrophiles and with different types of metabolism, predominantly anaerobic and chemosynthetic. TACK is a clade that is sister to the Asgard branch that gave rise to the eukaryotes. It has been proposed that the TACK clade be classified as Crenarchaeota and that the traditional "Crenarchaeota" (Thermoproteota) be classified as a class called "Sulfolobia", along with the other phyla with class rank or order.

Crenarchaeol is a glycerol biphytanes glycerol tetraether (GDGT) biological membrane lipid. Together with archaeol, crenarcheol comprises a major component of archaeal membranes. Archaeal membranes are distinct from those of bacteria and eukaryotes because they contain isoprenoid GDGTs instead of diacyl lipids, which are found in the other domains. It has been proposed that GDGT membrane lipids are an adaptation to the high temperatures present in the environments that are home to extremophile archaea

Christa Schleper is a German microbiologist known for her work on the evolution and ecology of Archaea. Schleper is Head of the Department of Functional and Evolutionary Biology at the University of Vienna in Austria.

References

  1. Oren A, Garrity GM (2021). "Valid publication of the names of forty-two phyla of prokaryotes". Int J Syst Evol Microbiol. 71 (10): 5056. doi: 10.1099/ijsem.0.005056 . PMID   34694987.
  2. 1 2 3 4 Brochier-Armanet C, Boussau B, Gribaldo S, Forterre P (March 2008). "Mesophilic Crenarchaeota: Proposal for a third archaeal phylum, the Thaumarchaeota". Nature Reviews Microbiology. 6 (3): 245–52. doi:10.1038/nrmicro1852. PMID   18274537. S2CID   8030169.
  3. Tourna M, Stieglmeier M, Spang A, Könneke M, Schintlmeister A, Urich T, Engel M, Schloter M, Wagner M, Richter A, Schleper C (May 2011). "Nitrososphaera viennensis, an ammonia oxidizing archaeon from soil". Proceedings of the National Academy of Sciences of the United States of America. 108 (20): 8420–5. Bibcode:2011PNAS..108.8420T. doi: 10.1073/pnas.1013488108 . PMC   3100973 . PMID   21525411.
  4. DeLong EF (1992-06-15). "Archaea in coastal marine environments". Proceedings of the National Academy of Sciences. 89 (12): 5685–5689. Bibcode:1992PNAS...89.5685D. doi: 10.1073/pnas.89.12.5685 . ISSN   0027-8424. PMC   49357 . PMID   1608980.
  5. Brochier-Armanet C, Gribaldo S, Forterre P (December 2008). "A DNA topoisomerase IB in Thaumarchaeota testifies for the presence of this enzyme in the last common ancestor of Archaea and Eucarya". Biology Direct. 3: 54. doi: 10.1186/1745-6150-3-54 . PMC   2621148 . PMID   19105819.
  6. Spang A, Hatzenpichler R, Brochier-Armanet C, Rattei T, Tischler P, Spieck E, Streit W, Stahl DA, Wagner M, Schleper C (August 2010). "Distinct gene set in two different lineages of ammonia-oxidizing archaea supports the phylum Thaumarchaeota". Trends in Microbiology. 18 (8): 331–40. doi:10.1016/j.tim.2010.06.003. PMID   20598889.
  7. 1 2 3 Pearson A, Hurley SJ, Walter SR, Kusch S, Lichtin S, Zhang YG (2016). "Stable carbon isotope ratios of intact GDGTs indicate heterogeneous sources to marine sediments". Geochimica et Cosmochimica Acta. 181: 18–35. Bibcode:2016GeCoA.181...18P. doi:10.1016/j.gca.2016.02.034.
  8. 1 2 3 Pester M, Schleper C, Wagner M (June 2011). "The Thaumarchaeota: an emerging view of their phylogeny and ecophysiology". Current Opinion in Microbiology. 14 (3): 300–6. doi:10.1016/j.mib.2011.04.007. PMC   3126993 . PMID   21546306.
  9. 1 2 3 4 Walker CB, de la Torre JR, Klotz MG, Urakawa H, Pinel N, Arp DJ, Brochier-Armanet C, Chain PS, Chan PP, Gollabgir A, Hemp J, Hügler M, Karr EA, Könneke M, Shin M, Lawton TJ, Lowe T, Martens-Habbena W, Sayavedra-Soto LA, Lang D, Sievert SM, Rosenzweig AC, Manning G, Stahl DA (May 2010). "Nitrosopumilus maritimus genome reveals unique mechanisms for nitrification and autotrophy in globally distributed marine crenarchaea". Proceedings of the National Academy of Sciences of the United States of America. 107 (19): 8818–23. Bibcode:2010PNAS..107.8818W. doi: 10.1073/pnas.0913533107 . PMC   2889351 . PMID   20421470.
  10. Schouten S, Hopmans EC, Schefuß E, Damste JS (2002). "Distributional variations in marine crenarchaeotal membrane lipids: a new tool for reconstructing ancient sea water temperatures?". Earth and Planetary Science Letters. 204 (1–2): 265–274. Bibcode:2002E&PSL.204..265S. doi:10.1016/S0012-821X(02)00979-2. S2CID   54198843.
  11. "The LTP" . Retrieved 10 May 2023.
  12. "LTP_all tree in newick format" . Retrieved 10 May 2023.
  13. "LTP_06_2022 Release Notes" (PDF). Retrieved 10 May 2023.
  14. "GTDB release 08-RS214". Genome Taxonomy Database . Retrieved 10 May 2023.
  15. "ar53_r214.sp_label". Genome Taxonomy Database . Retrieved 10 May 2023.
  16. "Taxon History". Genome Taxonomy Database . Retrieved 10 May 2023.
  17. J.P. Euzéby. "Thaumarchaeota". List of Prokaryotic names with Standing in Nomenclature (LPSN). Retrieved 2021-03-20.
  18. Sayers, et al. "Thaumarchaeota". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 2021-03-20.
  19. Stieglmeier M, Klingl A, Alves RJ, Rittmann SK, Melcher M, Leisch N, et al. (August 2014). "Nitrososphaera viennensis gen. nov., sp. nov., an aerobic and mesophilic, ammonia-oxidizing archaeon from soil and a member of the archaeal phylum Thaumarchaeota". International Journal of Systematic and Evolutionary Microbiology. 64 (Pt 8): 2738–52. doi:10.1099/ijs.0.063172-0. PMC   4129164 . PMID   24907263.
  20. Muller F, Brissac T, Le Bris N, Felbeck H, Gros O (August 2010). "First description of giant Archaea (Thaumarchaeota) associated with putative bacterial ectosymbionts in a sulfidic marine habitat". Environmental Microbiology. 12 (8): 2371–83. Bibcode:2010EnvMi..12.2371M. doi: 10.1111/j.1462-2920.2010.02309.x . PMID   21966926.
  21. Zhalnina KV, Dias R, Leonard MT, Dorr de Quadros P, Camargo FA, Drew JC, et al. (7 July 2014). "Genome sequence of Candidatus Nitrososphaera evergladensis from group I.1b enriched from Everglades soil reveals novel genomic features of the ammonia-oxidizing archaea". PLOS ONE. 9 (7): e101648. Bibcode:2014PLoSO...9j1648Z. doi: 10.1371/journal.pone.0101648 . PMC   4084955 . PMID   24999826.
  22. Könneke M, Bernhard AE, de la Torre JR, Walker CB, Waterbury JB, Stahl DA (September 2005). "Isolation of an autotrophic ammonia-oxidizing marine archaeon". Nature. 437 (7058): 543–6. Bibcode:2005Natur.437..543K. doi:10.1038/nature03911. PMID   16177789. S2CID   4340386.
  23. Lehtovirta-Morley LE, Stoecker K, Vilcinskas A, Prosser JI, Nicol GW (September 2011). "Cultivation of an obligate acidophilic ammonia oxidizer from a nitrifying acid soil". Proceedings of the National Academy of Sciences of the United States of America. 108 (38): 15892–7. Bibcode:2011PNAS..10815892L. doi: 10.1073/pnas.1107196108 . PMC   3179093 . PMID   21896746.
  24. Lebedeva EV, Hatzenpichler R, Pelletier E, Schuster N, Hauzmayer S, Bulaev A, Grigor'eva NV, Galushko A, Schmid M, Palatinszky M, Le Paslier D, Daims H, Wagner M (2013). "Enrichment and genome sequence of the group I.1a ammonia-oxidizing Archaeon "Ca. Nitrosotenuis uzonensis" representing a clade globally distributed in thermal habitats". PLOS ONE. 8 (11): e80835. Bibcode:2013PLoSO...880835L. doi: 10.1371/journal.pone.0080835 . PMC   3835317 . PMID   24278328.
  25. Li Y, Ding K, Wen X, Zhang B, Shen B, Yang Y (March 2016). "A novel ammonia-oxidizing archaeon from wastewater treatment plant: Its enrichment, physiological and genomic characteristics". Scientific Reports. 6: 23747. Bibcode:2016NatSR...623747L. doi:10.1038/srep23747. PMC   4814877 . PMID   27030530.
  26. Santoro AE, Dupont CL, Richter RA, Craig MT, Carini P, McIlvin MR, et al. (January 2015). "Genomic and proteomic characterization of "Candidatus Nitrosopelagicus brevis": an ammonia-oxidizing archaeon from the open ocean". Proceedings of the National Academy of Sciences of the United States of America. 112 (4): 1173–8. Bibcode:2015PNAS..112.1173S. doi: 10.1073/pnas.1416223112 . PMC   4313803 . PMID   25587132.
  27. Blainey PC, Mosier AC, Potanina A, Francis CA, Quake SR (February 2011). "Genome of a low-salinity ammonia-oxidizing archaeon determined by single-cell and metagenomic analysis". PLOS ONE. 6 (2): e16626. Bibcode:2011PLoSO...616626B. doi: 10.1371/journal.pone.0016626 . PMC   3043068 . PMID   21364937.
  28. Kim BK, Jung MY, Yu DS, Park SJ, Oh TK, Rhee SK, Kim JF (October 2011). "Genome sequence of an ammonia-oxidizing soil archaeon, "Candidatus Nitrosoarchaeum koreensis" MY1". Journal of Bacteriology. 193 (19): 5539–40. doi:10.1128/JB.05717-11. PMC   3187385 . PMID   21914867.
  29. Park SJ, Kim JG, Jung MY, Kim SJ, Cha IT, Kwon K, Lee JH, Rhee SK (December 2012). "Draft genome sequence of an ammonia-oxidizing archaeon, "Candidatus Nitrosopumilus koreensis" AR1, from marine sediment". Journal of Bacteriology. 194 (24): 6940–1. doi:10.1128/JB.01857-12. PMC   3510587 . PMID   23209206.
  30. Mosier AC, Allen EE, Kim M, Ferriera S, Francis CA (April 2012). "Genome sequence of "Candidatus Nitrosopumilus salaria" BD31, an ammonia-oxidizing archaeon from the San Francisco Bay estuary". Journal of Bacteriology. 194 (8): 2121–2. doi:10.1128/JB.00013-12. PMC   3318490 . PMID   22461555.
  31. Bayer B, Vojvoda J, Offre P, Alves RJ, Elisabeth NH, Garcia JA, Volland JM, Srivastava A, Schleper C, Herndl GJ (May 2016). "Physiological and genomic characterization of two novel marine thaumarchaeal strains indicates niche differentiation". The ISME Journal. 10 (5): 1051–63. Bibcode:2016ISMEJ..10.1051B. doi:10.1038/ismej.2015.200. PMC   4839502 . PMID   26528837.
  32. 1 2 Brochier-Armanet C, Gribaldo S, Forterre P (February 2012). "Spotlight on the Thaumarchaeota". The ISME Journal. 6 (2): 227–30. Bibcode:2012ISMEJ...6..227B. doi:10.1038/ismej.2011.145. PMC   3260508 . PMID   22071344.
  33. 1 2 Könneke M, Schubert DM, Brown PC, Hügler M, Standfest S, Schwander T, Schada von Borzyskowski L, Erb TJ, Stahl DA, Berg IA (June 2014). "Ammonia-oxidizing archaea use the most energy-efficient aerobic pathway for CO2 fixation". Proceedings of the National Academy of Sciences of the United States of America. 111 (22): 8239–44. Bibcode:2014PNAS..111.8239K. doi: 10.1073/pnas.1402028111 . PMC   4050595 . PMID   24843170.
  34. 1 2 Qin W, Amin SA, Martens-Habbena W, Walker CB, Urakawa H, Devol AH, Ingalls AE, Moffett JW, Armbrust EV (2014). "Marine ammonia-oxidizing archaeal isolates display obligate mixotrophy and wide ecotypic variation". Proceedings of the National Academy of Sciences. 111 (34): 12504–12509. Bibcode:2014PNAS..11112504Q. doi:10.1073/PNAS.1324115111. ISSN   0027-8424. PMC   4151751 . PMID   25114236.
  35. Mussmann M, Brito I, Pitcher A, Sinninghe Damsté JS, Hatzenpichler R, Richter A, Nielsen JL, Nielsen PH, Müller A, Daims H, Wagner M, Head IM (October 2011). "Thaumarchaeotes abundant in refinery nitrifying sludges express amoA but are not obligate autotrophic ammonia oxidizers". Proceedings of the National Academy of Sciences of the United States of America. 108 (40): 16771–6. Bibcode:2011PNAS..10816771M. doi: 10.1073/pnas.1106427108 . PMC   3189051 . PMID   21930919.
  36. Santoro AE, Buchwald C, McIlvin MR, Casciotti KL (2011-09-02). "Isotopic Signature of N2O Produced by Marine Ammonia-Oxidizing Archaea". Science. 333 (6047): 1282–1285. Bibcode:2011Sci...333.1282S. doi:10.1126/science.1208239. ISSN   0036-8075. PMID   21798895. S2CID   36668258.
  37. Doxey AC, Kurtz DA, Lynch MD, Sauder LA, Neufeld JD (February 2015). "Aquatic metagenomes implicate Thaumarchaeota in global cobalamin production". The ISME Journal. 9 (2): 461–71. Bibcode:2015ISMEJ...9..461D. doi:10.1038/ismej.2014.142. PMC   4303638 . PMID   25126756.
  38. Muller F, Brissac T, Le Bris N, Felbeck H, Gros O (August 2010). "First description of giant Archaea (Thaumarchaeota) associated with putative bacterial ectosymbionts in a sulfidic marine habitat". Environmental Microbiology. 12 (8): 2371–83. Bibcode:2010EnvMi..12.2371M. doi: 10.1111/j.1462-2920.2010.02309.x . PMID   21966926.

Further reading